The Seven Sins of Evolutionary Psychology - Konrad Lorenz Institute

ContentsEvolutionary Psychology: An ExchangeJaak Panksepp/Jules B. Panksepp 2 A Synopsis of“The Seven Sins of Evolutionary Psychology”Ullica Segerstråle 6 Minding the Brain:The Continuing Conflict about Models and RealityLinda Mealey 14 “La plus ça change…”:Response to a Critique of Evolutionary PsychologyCarlos Stegmann 20 Comments on Jaak Panksepp and Jules B. Panksepp’sPaper “The Seven Sins of Evolutionary Psychology”Russell Gardner, Jr. 25 Affective Neuroscience, Psychiatry, and SociophysiologyGerhard Meisenberg 31 Degrees of ModularityIan Pitchford 39 No Evolution. No CognitionScott Atran 46 The Case for Modularity: Sin or Salvation?Jaak Panksepp/Jules B. Panksepp 56 A Continuing Critique of Evolutionary Psychology:Seven Sins for Seven Sinners, Plus or Minus TwoGeneral ArticlesShulamith Kreitler 81 An Evolutionary Perspective on Cognitive OrientationChris MacDonald 98 Evolutionary Ethics:Value, Psychology, Strategy and Conventions106 Zusammenfassungen der Artikel in deutscher SpracheImpressumEvolution and Cognition: ISSN: 0938-2623 Published by: Konrad Lorenz Institut fürEvolutions- und Kognitionsforschung, Adolf-Lorenz-Gasse 2, A-3422 Altenberg/Donau.Tel.: 0043-2242-32390; Fax: 0043-2242-323904; e-mail: sec@kla.univie.ac.at; WorldWide Web: http://www.univie.ac.at/evolution/kli/ Editors: Rupert Riedl, ManfredWimmer Layout: Alexander Riegler Aim and Scope: “Evolution and Cognition” is aninterdisciplinary forum devoted to all aspects of research on cognition in animals and humans. The major emphasis ofthe journal is on evolutionary approaches to cognition, reflecting the fact that the cognitive capacities of organisms resultfrom biological evolution. Empirical and theoretical work from both fields, evolutionary and cognitive science, is accepted,but particular attention is paid to interdisciplinary perspectives on the mutual relationship between evolutionary andcognitive processes. Submissions dealing with the significance of cognitive research for the theories of biological andsociocultural evolution are also welcome. “Evolution and Cognition” publishes both original papers and review articles.Period of Publication: Semi-annual Price: Annuals subscription rate (2 issues): ATS 500; DEM 70, US$ 50; SFr 60;GBP 25. Annual subscriptions are assumed to be continued automatically unless subscription orders are cancelled bywritten information. Single issue price: ATS 300; DEM 43; US$ 30; SFr 36; GBP 15 Publishing House: WUV-Universitätsverlag/ViennaUniversity Press, Berggasse 5, A-1090 Wien, Tel.: 0043/1/3105356-0, Fax: 0043/1/3197050 Bank: Ersteösterreichische Spar-Casse, Acct.No. 073-08191 (Bank Code 20111) Advertising: Vienna University Press, Berggasse 5,A-1090 Wien. Supported by Cultural Office of the City of Vienna, Austrian Federal Ministry of Science/ Transportationand the Section Culture and Science of the Lower Austrian State Government.

Jaak Panksepp and Jules B. PankseppA Synopsis of“The Sevens Sins of Evolutionary Psychology”Our target article (PANKSEPP/PANKSEPP 2000) summarizedsome common and uncommon concerns thatneuroscientifically oriented investigators of themind/brain may have with currently fashionableversions of evolutionary psychology. Our main thesiswas that an evolutionary psychology that seeksto understand “human nature” will be woefully incompleteand perhaps misguided if it does not incorporatethe lessons from the past century of researchon subcortical emotional and motivationalsystems that all mammals share. It is advisable tobegin building an evolutionary viewpoint of mindbased on comparative concepts that incorporate theintrinsic, evolutionarily provide systems found inall mammalian brains.The more evangelical varieties of psycho–evolutionarythought that arose from social-science traditionshave typically not been concerned with howthe human mind emerges from specific brain functions.Thus, our argument was motivated by threeconverging and overlapping issues: (1) The failure ofevolutionary psychology (e.g., BUSS 1999, and manyothers) to address the evolved, adaptive mechanismsin the mammalian brain that have been andare being studied by many investigators in the neurosciences.(2) The prolific tendency of evolutionarypsychology to generate adaptive stories, especiallywith regard to human socio–emotional processes,without much apparent concern about how suchpresumed adaptations might be instantiated in neuralsystems through a great diversity of brain–environmentinteractions (e.g., COSMIDES/TOOBY 2000).(3) The concurrent claim that evolutionary psychologyis now providing one of the most robust lines ofthought for identifying the types of evolved mechanismsthat actually exist in the human brain(TOOBY/COSMIDES 2000).Our discussion was framed with the recognitionthat several distinct schools of thought are currentlyproviding conceptual frameworks to help guide futureempirical inquiries in the ongoing attempt tounderstand how evolution constructed the mind/brain of humans and other animals. Also, our articlewas premised on the full acceptance that evolutionaryviewpoints are essential for specifying the neuropsychologicalnatural kinds that actually exist, asancestral birthrights, within the brains of humansas well as other animals.Our specific concerns were as follows:1. Are there really Pleistocene sources of currenthuman social adaptations? Evolutionary Psychologistsseem to be backing off from this assumptionas they begin to realize that neocortex is the maintype of cerebral tissue that emerged two millionyears ago, when hominid brains diverged from the“chimp-sized” brains of our ancestral stock. Thus,the critical question is whether special-purposeprograms emerged in neocortical tissues primarilythrough genetic changes that occurred in the humanline during the past few million years. That issueprobably cannot be answered unambiguouslywithout concurrent behavioral neuroscience andmolecular biology research. If such adaptationsemerged, they are likely to reflect quite general informationhandling capacities, permitting abilitiesranging from art to human languages, fromthought to culture. Most evolved, special-purposesystems may be devoted to functions we still sharewith other animals.2. Excessive species-centrism in Evolutionary Psychology.Completion of the human genomeproject has unequivocally highlighted the likelihoodfor genetic similarities across all mammalianspecies. We suggest that without a clear recognitionof what we share with other animals, it will be impossibleto specify what is truly unique in the intrinsic,genetically-provided capacities of the humanmind/brain. The neuroanatomical andneurochemical homologies that exist in all mammalianbrains presently provide the most substantiveway to specify the types of evolved faculties inour own minds.Evolution and Cognition ❘ 2 ❘ 2001, Vol. 7, No. 1

A Synopsis of “The Sevens Sins of Evolutionary Psychology”3. The sin of adaptationism. It is well recognizedthat one must always discriminate between adaptationsarising from natural selection and those arisingfrom the emergence of flexible abilities to learnnew skills and strategies. The cortex obviously becomesspecialized for a large number of special purposeskills and abilities during development (manyof which may seem superfically “encapsulated” becausemost of our thoughts are constrained byprimitive emotional systems that regulate our feelings),but current evolutionary psychology providesno special inroads to understanding whichadaptations have truly emerged through natural selection.4. The sin of massive modularity. From a neuroscienceperspective, “modularity” is an obsoleteconcept, resembling the “centers” concept that wasdiscarded by scientists doing brain research severaldecades ago. There are many special-purpose circuitsand systems in subcortical areas of the brain,but those systems are not encapsulated and appearto interact with each other extensively. For instance,during every emotion, the activities of widespreadbrain areas are modified. Obviously, a criticalscientific issue is the degree to which the specializedfunctions of higher cerebral abilities exist as birthrightsand how much they reflect the operation ofdevelopmental programs.5. On the conflation of emotions and cognitions.The cortex mediates many specialized cognitiveabilities. Most of the key systems for mammalianemotionality are subcortically organized, and theneglect of these systems can only lead to a very mistakenview of how emotionality is truly elaboratedin the human brain. Disciplines that focus simplyon the higher cortico–cognitive aspects of emotionalprocessing are bound to make many mistakesabout the fundamental nature of emotions and affectiveprocesses (PANKSEPP 2001).6. The absence of credible neural perspective inEvolutionary Psychology. This is a general criticismof all mind sciences that fail to take real brainissues into consideration. Although the full potentialof the mind emerges from organismic–environmentalinteractions, all foundational issues need tobe linked to our gradually emerging understandingof how specific neural systems function. The neurosciencerevolution has provided enough raw evidencefor everyone to begin thinking more clearlyabout those underlying brain issues.7. Anti-organic bias or the computationalist/representationalistmyth. There have always beentwo grand perspectives for explaining the organizationof mind—mind as an organic process and mindas a “spiritual” process that can exist independentlyof the material world. The modern computationalview still tends to subscribe to a modern variant ofthe latter view. From a non-dualistic, materialisticperspective, that may be highly misleading. Allminds are probably strongly constrained by the organic–neuralprocesses by which they are instantiated.To neglect those issues, in preference for a strictcomputationalist view, is likely, we believe, to retarddeep understanding. In our reply to the commentators,we highlight a recent critique by FODOR (2000),one of the great promoters of the computationalism,who now agrees that grave errors have beenmade during the cognitive revolution.Our criticisms do not apply equally to all existinglines of psycho–evolutionary thought, but mainlyto the rhetorically most audible social psychologicalvariants that arose from the ashes of the contentious“sociobiology wars” of the late 1970s and early80s. Additional issues could have been raised, suchas the massive interactions of certain well-characterizedbrain systems (e.g., dopamine, norepinephrine,serotonin, acetylcholine, GABA and glutamate)which participate heavily in all of the processesabout which evolutionary psychologists opine. Severalrelated “sinful” tendencies could have beenhighlighted, but they all ultimately relate to the failureof Evolutionary Psychology to assume a deeplyorganic stance that fully respects the role of developmentalsystems in the epigenetic creation ofhigher mental faculties.Our overall view is that the developmental interactionsbetween ancient, special-purpose circuitsand more recent, general-purpose brain systems,such as those found in the neocortex, can generatemany of the “modularized” human abilities thatEvolutionary Psychology has entertained. By simplyaccepting the remarkable degree of neocortical plasticity,especially during human development, genetically-dictated,sociobiological “modules” beginto resemble products of dubious human ambitionrather than sound scientific inquiry. We believe thatif Evolutionary Psychology attempts to construct aview of human nature based upon inclusive-fitnessissues and improbable neurological assumptions,while continuing to avoid a substantive confrontationwith our ancient animalian heritage, it is aslikely to retard lasting scientific progress as to advanceit. In short, the only place many concepts ofEvolution and Cognition ❘ 3 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. Pankseppevolutionary psychology may be encapsulated arein the minds of scholars who, all too often, havelittle wish to immerse themselves in the essentialneuroscientific and genetic issues.We recognize that there are recent strands ofthought in evolutionary psychology that take positionsmore congenial to our own. For instance, weadmire BADCOCK’s (2000) confrontation with key biologicalissues, as well as his willingness to try tolink evolutionary thought to earlier intellectual traditions.We especially appreciate the way he handlesthe possibility that cultural issues, obviouslyimportant in their own right, may be linked to someimportant heritability findings such as genomic imprinting.Stunning discoveries such as that of KEV-ERNE and colleagues (1996), indicating that the developmentof neocortex may be influenced more bymaternally-imprinted genes while the developmentof key emotional areas such as the hypothalamusare governed more by paternally-imprinted genes,provide novel ancestral–genetic tethers over brainorganization that may have profound implicationsfor the nature–nurture controversy.Of course, we remain as intellectually stimulatedby evolutionary psychological stories as anyone. Wehave been especially entertained by the recent willingnessof some theorists to recognize the importanceof the emergence of artistic endeavors in thesocio–cortical emergence of our species (MILLER2000) and of course, the view of language as partly asocial-grooming adaptation (DUNBAR 1995) whichfits nicely with some animal social-systems data(PANKSEPP 1998a). However, we would again adviseevolutionists to view the neocortex as the generalpurposeplayground for our basic attentional, emotionaland motivational systems, as opposed to theirsources. Indeed, it may well be that such a generalpurposeassociational spaces are the ideal playgroundsfor the many dimensions of human creativityand entertainment, both abundantly seriousand humorous depending on the depth of our emotionalpersonalities as muchAuthors’ addressJaak Panksepp and Jules B. Panksepp,J.P. Scott Center for Neuroscience, Mind &Behavior, Departments of Psychology andBiological Sciences, Bowling Green StateUniversity. Emails: jpankse@bgnet.bgsu.eduor julesp@caspar.bgsu.eduas anything else. With ourneocortex we can do whateverwe can imagine—and becauseof the mass of generalpurpose associational spacethat we possess, there are fewobvious limits to our imagination.This affords us great libertiesin our humanistic endeavorsand helps create great havoc for ourscientific ones.We would simply note that the expansion of instinctsat the turn of the previous century is nowbeing matched by the postulation of geneticallymodularized cognitive functions. The “expansionof instincts” failed scientifically because they werenot tethered to brain systems, and the expansion of“modularized” functions may have a comparablefate for similar reasons. A recent, and more guardedattempt to bring instinctual systems back to theforefront of our thinking (e.g., PANKSEPP 1998a) hasbeen premised on only entertaining basic psycho–behavioral entities for which there exist robust andconverging neuroscientific evidence. This also givesus considerable confidence in defining what typesof systems we need to study more earnestly in humans,while preventing mere human creativityfrom being the major guide of what does or does notexist in the natural organization of our minds. If wecarefully work out the fundamental neuro-psychologicalprocesses that we share with other animals,we will be in a better position to comprehend whatis truly unique in our own mind/brains.Our implicit aim in the target article was tostrongly encourage similar constraints in evolutionarypsychological thought. Obviously, there aremany stories that remain to be told around this newintellectual campfire, and the likelihood that primitiveforms of affective self-representation still link usto a deep animalian past may be a worthy concept formainstream thought in evolutionary psychology(PANKSEPP 1998a,b). Indeed, this view may allow us tobetter understand and harmonize scientific and humanisticendeavors. We do hope that an increasingnumber of scholars will begin to incorporate the evolutionarypassages of deep-time into their considerationsof the evolved nature of human mind-flesh…most especially in ways that can lead to the deep empiricalevaluation of ideas. Being inheritors of a vastand to some extent, general-purpose cerebral canopy,the “demoralizing” fact thatour mental activities are alsoconstrained by our instinctualtendencies need no longer beseen as being at odds with ourmore open-ended and nobledesires. With a modest degreeof mental flexibility, theycould serve as a foundation fora new humanism.Evolution and Cognition ❘ 4 ❘ 2001, Vol. 7, No. 1

A Synopsis of “The Sevens Sins of Evolutionary Psychology”ReferencesBadcock, C. (2000) Evolutionary Psychology: A Critical Introduction.Polity Press: Cambridge UK.Buss, D. M. (1999) Evolutionary Psychology: The New Scienceof the Mind. Allyn and Bacon: Boston.Cosmides, L./Tooby, J. (2000) Evolutionary psychology andthe emotions. In: Lewis, M./Haviland, J. (eds) The Handbookof Emotions, 2nd edition. Guilford: New York, pp.91–116.Dunbar, R. (1995) Grooming, Gossip and The Evolution ofLanguage. Harvard University Press: Cambridge, MA.Fodor, J. (2000) The Mind Doesn’t Work That Way, TheScope and Limits of Computational Psychology. The MITPress: Cambridge, MA.Keverne, E. B./Fundele, R., et al. (1996) Genomic imprintingand the differential roles of parental genomes in brain development.Developmental Brain Research 92: 91–100.Miller, G. (2000) The Mating Mind: How Sexual ChoiceShaped the Evolution of Human Nature. William Heinemann:London UK.Panksepp, J. (1998a) Affective Neuroscience, The Foundationsof Human and Animal Emotions. Oxford UniversityPress: New York.Panksepp, J. (1998b) The periconscious substrates of consciousness:Affective states and the evolutionary origins ofthe SELF. Journal of Consciousness Studies 5: 566–582.Panksepp, J. (2001) At the interface between the affective,behavioral and cognitive neurosciences. Decoding theemotional feelings of the brain. Brain and Cognition, inpress.Panksepp, J./Panksepp, J. B. (2000) The seven sins of evolutionarypsychology. Evolution and Cognition 6(2): 108–131.Tooby, J./Cosmides, L. (2000) Toward mapping the evolvedfunctional organization of mind and brain. In: Gazzaniga,M.S. (ed) The New Cognitive Neuroscience, 2nd edition.MIT Press: Cambridge MA, pp. 1167–1178.Evolution and Cognition ❘ 5 ❘ 2001, Vol. 7, No. 1

Ullica SegerstråleMinding the BrainThe Continuing Conflict about Models and RealityTHERE IS THE FAMOUSjoke about a travelerasking a local if he knewhow far it was to a particulartown. Yes, said theman. But, he added, youcan’t get to there fromhere! This may be ratherlike the way Jaak andJules PANKSEPP view theprospect of elucidatingthe mind with the help ofevolutionarily informedcognitive neuroscience.Theirs is one of the strongestscientific objectionsso far to the whole enterpriseof evolutionarypsychology. For thePANKSEPPs, there is onlyone correct way to go,and that is through thefield of affective neuroscience.You cannot findout about the functioningof the mind/brain byasking questions aboutadaptation and design,and by reconstructingthe kinds of problemsour Pleistocene ancestorsfaced, they say. Evolutionarypsychologists areon the wrong track. Ifyou want to find outabout functions, youshould not be looking atevolutionary biology, but rather at neuroanatomy.Lining up ‘seven deadly sins’ of evolutionarypsychology, as they do, may sound a bit harsh.(Well, Konrad LORENZ listed eight deadly sins, butthat was of a much broader population, ‘civilizedAbstractThe battle for ‘good science’, so evident in the sociobiologycontroversy, now continues in the realm of neuroscience.Modelers again clash with hardnosed scientific‘realists’. From the PANKSEPPs’ point of view, in order tobe scientific and ground their own enterprise, evolutionarypsychologists ought to pay attention to anatomyand the evolutionary history of the mind.Moreover, the PANKSEPPs’ criticism of cognitive neuroscienceparallels traditional ethological objections tosociobiology, emphasizing not only the ultimate function,but also the development, evolutionary history,and proximate causes of behaviors. This may signalan emerging conflict between evolutionary psychologyand ethology. Meanwhile, the PANKSEPPs are in goodcritical company with many others, such as developmentallyoriented scientists, who argue that the modularityin the adult brain does not reflect a preexistingdomain specificity, but is rather a product ofepigenesis. Indeed, a new interdisciplinary alliancemay be currently emerging, as the critics of sociobiologyand evolutionary psychology, in their search for amore ‘holistic’ and ‘pluralistic’ paradigm, have recentlydiscovered ethology!Finally, we should not forget that evolutionary psychologyis an evolving research program. Most likelywe will see an eventual rapprochement between cognitiveand affective neuroscience. A possible mediatorcould be the interdisciplinary field of nonverbal communication.Key wordsSociobiology controversy, neuroscience, ethology, evolutionarypsychology, development, nonverbal communication,human nature, modularity.Evolution and Cognition ❘ 6 ❘ 2001, Vol. 7, No. 1man’ as a whole). I believethat the PANKSEPPs’stance may be reflectingsomething more than anopposition between acognitive and an affectiveapproach to neuroscience.It could be an expressionof a deeperconflict between twoneighboring fields: evolutionarypsychologyand ethology. In thiscommentary I note howtheir criticism often coincideswith an ethologicalviewpoint and someof the objections thatethologists have typicallyhad to sociobiology.I also find the PANKSEPPs’views to be surprisinglyin tune with those of thecritics of sociobiology.Indeed, a new ‘criticalsyndrome’ appears to beforming in academia,bringing together researchersfrom differentfields with different typesof objections to evolutionarypsychology andgene-selectionist sociobiology.Finally I discussthe role of the interdisciplinaryfield of nonverbalcommunication as apossible mediator between cognitive and affectiveneuroscience, or between an information orientedand an anatomical approach.The PANKSEPPs are convinced that the best scientificprogress in elucidating the mind will be made

Minding the Brainon the basis of real, physical knowledge rather thanthrough modeling. In the sociobiology controversy,experimentally oriented scientists voiced a similarcriticism of the efforts of sociobiologists, who wereperceived as constructing models which had little todo with reality. Sometimes the accusation went sofar as to allege that sociobiologists actually believedin the reality of such things as average gene frequencies,rather than considering these to be mere calculations.And just in the same way in the earlier (andoverlapping) IQ controversy, psychometricians hadbeen accused by largely the same critics of ‘reifying’a mere computation of test scores into a mysteriousentity, general intelligence, g. According to the critics,since this was a mere statistical construct, it‘could not’ exist in anyone’s head. 1 For the critics,real science had to do with research at the molecularlevel (cf. SEGERSTRÅLE 2000, especially chapters 13and 14).The sociobiologists, for their part, felt perfectlyjustified in constructing models, as did the psychometriciansin making their correlations and calculations.For them, these were quite legitimate approachesin their respective fields. They had nodoubts, meanwhile, that their results, obtained by adifferent scientific route, would be compatible withfuture molecular level research.Nowhere is the conflict clearer between a modelingand a ‘realist’ approach than in the case of themeaning of the ‘gene’. For instance, molecular biologistGunther STENT (1977) called DAWKINS’ notionof the selfish gene ‘a heinous terminological sin’,pointing out that the ‘true’ gene was ‘unambiguously’that unit of genetic material which encodedthe amino acid sequence of a particular protein (acistron)—and, in addition, that it was not selfish!Indeed, laboratory geneticists sometimes seemed especiallydispleased with sociobiological models—sometimes one got the feeling that they wanted sociobiologiststo go into the lab and do ‘real’ scientificresearch on genes (e.g., HOWE/LYNE 1992; SEG-ERSTRÅLE 2000, pp386–387).In other words, in these controversies there was aclear dividing line in regard to what was ‘good science’.The critics felt remarkably free to accuse sociobiologistsof scientific ‘error’ for holding viewsand using methods that these scientists themselvesconsidered standard in their own field! Some criticswent as far as using the ‘errors’ of sociobiologists asevidence that the sociobiological agenda ‘must’ bepolitically motivated. According to them, the sociobiologists’scientific quest ‘could not’ be driven byscientific interest, because the only interesting sciencewas done at the molecular level (e.g., LEWONTIN1975; CHOROVER 1979).Now in the case of the PANKSEPPs vs. evolutionarypsychology we seem to have a similar oppositionbetween two views of ‘good science’: a ‘hard data’approach conflicting with a model-happy one. Takethe PANKSEPPs’ criticism of the computational cognitivescience models used by evolutionary psychologists.According to them, real science is done at theanatomical, physical level. PET scans and otherfancy new methods for showing brain activity are atbest correlational, and can easily be misleadingabout the true neural processes. They also point outthat evolutionary psychologists, following the leadof cognitive scientists, use a digital model of mind.But the mind is analog, they protest.In other words, the battle for good science, so evidentin the sociobiology controversy, now continuesin the realm of neuroscience. Modelers again clashwith hardnosed scientific ‘realists’. But if the sociobiologistswere criticized for telling adaptive ‘just-so’stories about hypothetical genes ‘for’ behaviors andspeculating on the basis of very little evidence, thepresent authors go one step further. They accuse theevolutionary psychologists of ignoring a whole bodyof evidence that actually exists: the findings fromcomparative neuroscience! And the consequencesare grave, the PANKSEPPs maintain. Evolutionary psychologistsfeel free to postulate a system of massivemodularity in the neocortex. But from the point ofview of comparative neuroanatomy, such modulescould not have evolved.To rub this in, they cite TOOBY and COSMIDES’(1992) own call for consistency across scientificfields. TOOBY and COSMIDES criticize social scientistsfor what they call the Standard Social ScienceModel, which regards the human mind as a generalpurposemachine for learning. Such a model is anevolutionary impossibility, according to them. ThePANKSEPPs have no direct argument with this.Rather, they point out that when it comes to theirown research, TOOBY and COSMIDES are not applyingtheir own principle: they do not ground themselveson already existing knowledge about the brain.The whole premise of evolutionary psychology iswrong, say the PANKSEPPs. The problem is not theassumption of modularity per se, but evolutionarypsychologists are looking for modules in the wrongplace. Any claim about the structure of the humanmind/brain would have to take into account its evolutionaryhistory. If routinized behaviors of the‘module’ kind exist and confer survival value on humans,these are bound to be of a kind we share withEvolution and Cognition ❘ 7 ❘ 2001, Vol. 7, No. 1

Ullica Segerstråleother mammals. This means we had better look formodules in older, subcortical parts of the brain, particularlythose parts that deal with emotions, thoseimportant motivators for animal survival. (Herethey invoke not only Paul MACLEAN’S famous ‘triunebrain’ but also E. O. WILSON’s opening statementsin Sociobiology). This is why, they argue, comparative,affective neuroscience cannot be ignored.And they have an additional argument. Evolutionarypsychologists have erroneously taken the mostrecent and particular characteristic of the humanbrain—our large neocortex—expecting to find modulesthere. They may believe they have found somemodules in the adult brain, but any such modules arelikely to be a product of development, they point out.In contrast to the evolutionary psychologists, thePANKSEPPs envision the individual as initially workingwith a type of general purpose program; it is onlylater during ontogenesis that more specialized modulesare being formed. So for the PANKSEPPs, cognitivemodules, to the extent they can be found, are actuallyby-products, not primary adaptations.Here we see that the PANKSEPPs adhere to GOULD’sand LEWONTIN’s ‘spandrel’ thinking (GOULD/LEWONTIN 1979), according to which many traitsthat sociobiologists see as direct adaptations mayactually be mere by-products of evolution, just likesome details of architecture—spandrels—are createdaround vaults. This would seem to put themsquarely in the same camp as the ‘anti-adaptationist’critics of sociobiology. And they in fact go a stepfurther, arguing, with GOULD (1991), that humanlanguage, too, is likely to be an evolutionary byproduct.2 And if language is a mere spandrel, thenfocusing on language as the model module is obviouslymisdirected. (This latter point is an unnecessarypiece of overkill, in my view, because the PANK-SEPPs’ argument does not hinge on the nature oflanguage. True, if it were possible to conclusivelyprove that language was a mere spandrel, theywould have a point. But GOULD is operating ratheron the level of principle, while for many followingCHOMSKY’s and PINKER’s work, the idea of a particularlanguage module does appear persuasive. Mightit not be possible to regard language as a modulewithout taking the next step with PINKER (1994,1997a) and argue from the modularity of languageto a view of massive modularity?).From the PANKSEPPs’ point of view, then, in orderto be scientific and ground their own enterprise,evolutionary psychologists ‘ought to’ pay attentionto anatomy and the evolutionary history of themind. They are dismayed that COSMIDES and TOOBYin a recent contribution went so far as to attempt todeal with emotion without making any mention ofaffective neuroscience (COSMIDES/TOOBY 2000).What could be some reasons for this neglect amongthe leading theorists in evolutionary psychology?In the first place, evolutionary psychologists havetheir hands full. They are busy fighting a multi-frontbattle: against cultural anthropologists and mainstreampsychologists on the one hand, and againstsniping critics of ‘adaptationism’ on the other. Inmany regards, they have inherited the mantle of sociobiology,and with this, its critics. 3 Despite theirattempt to emphasize universal features of humansand ‘the psychic unity of mankind’, and thus tryingto fend off accusations of racism and biological determinism,they are labelled ‘ultra-DARWINIANS’ byGOULD and other critics (GOULD 1997a–b; ELDREDGE1995; ROSE 1998). They have to create legitimacy fortheir field at the same time as they have to establisha viable research program.Predictably, too, the reception of evolutionarypsychology among mainstream psychologists hasnot been too hospitable. But there were potentialallies among scholars in other fields which also dealwith the mind, and whose approach is similar or atleast compatible with the methods typically used inevolutionary psychology. Cognitive science seemedthe natural way to go. By associating itself with cognitivescience, evolutionary psychology could rideon the wave of the triumphant cognitive revolution.It could be linked to booming research fieldssuch as artificial intelligence, and to novel methodsfor monitoring cognitive processes. Computationalalgorithms would provide the necessary theoreticalrigor, while new imaging techniques would providethe physical correlates for their claims.The PANKSEPPs are duly respectful of the cognitiverevolution—it is just that they feel that the next revolution,the neuroscientific one, did not have achance to make its mark before the surge of interestin evolutionary explanation. The new kin selectionparadigm swept people off their feet, they complain.And here it becomes clear that, unlike sociobiologistsand evolutionary psychologists, the PANK-SEPPs are interested not only in ultimate answersabout function, but also in the development, evolutionaryhistory, and proximate causes of behaviors.In other words, they want to emphasize the importanceof all of TINBERGEN’s famous ‘four questions’(TINBERGEN 1963). As we see, they are particularlyconcerned about developmental aspects.Interestingly, a very similar complaint about evolutionaryexplanations ‘taking over’ can be foundEvolution and Cognition ❘ 8 ❘ 2001, Vol. 7, No. 1

Minding the Brainamong bona fide ethologists in their comments onthe 1970’s sociobiological revolution. Just like thepresent authors, many ethologists felt that in the1970s their field, too, was on the brink of makinginteresting discoveries and solidifying a new, complex,paradigm for the study of animal behavior (asdemonstrated, e.g., by BATESON/HINDE 1976). Butwhen the exploitable-seeming new sociobiologicalframework arrived, many ethologists chose to focuson only one of TINBERGEN’s four questions, the questionabout function, and this served to drive ethologyinto a narrow direction (BATESON/KLOPFER1989). (Recently, though, the sociobiology ‘goldrush’ appears to be over, and the tide has turnedback toward a more complex ethological approach,e.g., KREBS/DAVIES 1997).But the PANKSEPPs do not only criticize evolutionarypsychology on scientific grounds, they alsowarn the field about potential political consequences.Here it is not clear exactly what they havein mind. The political critics of sociobiology actuallyreasoned in a number of different ways aboutsociobiology. The idea of a genetically based universalhuman nature was anathema for them, becausethey saw this as connected to genetic determinismand other evils. But a further belief was that ‘badscience’ (for the critics, sociobiology and IQ research)would inevitably lead to undesirable socialconsequences. As I have argued (SEGERSTRÅLE 2000),there is no obvious reason why ‘bad’ science (howeverdefined) would be particularly prone to socialabuse; it seems to me that ‘good’ science in thewrong hands would actually be more dangerous. Ifwe apply this to evolutionary psychology, therewould seem to be no inherent connection between,say, belief in modularity of mind and risk of politicalabuse. The PANKSEPPs may simply mean that, justlike sociobiology, evolutionary psychology is goingto be politically attacked as a deterministic, dangerousdoctrine about human nature.If that is what they mean, that has already happened.For instance THORNHILL and PALMER’s (2000)recent book about rape created quite a stir (e.g.,COYNE 2000; SHULEVITZ 2000; for a response seeTOOBY/COSMIDES 2000). And recently there was acollection of critical essays on evolutionary psychology(ROSE/ROSE 2000), which was surprisinglywell received in the popular press at least in theUnited Kingdom. I take this to mean that there is ablossoming market for this kind of moral–cum–scientificcriticism both in the UK and the US—just asthere is a great market for books that explain whywe do what we do. Indeed, a certain symbiosis appearsto prevail between the writers of popular evolutionarypsychology books and their critics—aphenomenon observed already in the sociobiologycontroversy (SEGERSTRÅLE 2000).Of course, evolutionary psychologists have madethe situation rather difficult for political critics. Wellaware of the attacks on sociobiology, they have carefullystated that they are interested in the evolvedhuman mind, not genes, and in a universal humannature, not human differences. They do not employpopulation-genetic models, and do not address geneticdifferences between populations. 4 They explicitlydistance themselves from human behavioral genetics(something WILSON needed for hissociobiological approach). This makes it hard toconstrue evolutionary psychologists as racists—thestandard move in left-wing criticism of biologicaltheories of human nature, and one that was muchemployed in the sociobiology controversy (cf. SEG-ERSTRÅLE 2000, esp. ch. 2, 9, and 15). Finally, they areuninterested in IQ differences, which means theyare not connectable either to the various standardaccusations against psychometricians (cf. SEGER-STRÅLE 2000, esp. ch. 10, 12, 13, 14). Indeed, a recentprimer in evolutionary psychology sounds remarkably‘politically correct,’ repudiating explicitly boththe notion of race and the idea of ‘general intelligence’(COSMIDES/TOOBY 1997).Whatever the evolutionary psychologists say,however, it is clear that the former critics of sociobiologydo not like evolutionary psychology, as little asthey like sociobiology (and they typically criticizethese fields together). In their search for scientificand moral/political counter-arguments the opponentsto evolutionary psychology have generated aplethora of different criticisms, not all necessarilycompatible with one another. This is déjà vu, andone needs only look back at the sociobiology controversy.Still, the tone is now more muted, and the critiquemore scientific and not as personal and nasty.One reason for this more cautious approach may bethat it has become harder to stir up general outragearound claims about the evolutionary basis of humanbehavior. At the turn of the millennium theidea of genes influencing behavior has become moreacceptable to the general public. In fact, some worrythat the situation has totally flip-flopped. They fear anew doctrine of ‘genetic essentialism’ will be holdinggenes increasingly responsible for our behavior(NELKIN/LINDEE 1995; cf. SEGERSTRÅLE 2000, ch. 19).This is why veteran critic of sociobiology, StevenROSE (2000), coauthor of Not In Our Genes(LEWONTIN/ROSE/KAMIN 1984), does not any longerEvolution and Cognition ❘ 9 ❘ 2001, Vol. 7, No. 1

Ullica Segerstråledeclare that nothing is in our genes. In his criticismof evolutionary psychology he now leans on a batteryof other arguments, particularly well-knownanti-adaptationist ones. What is interesting, however,is the remarkable convergence between thecriticism coming from this long-standing critic of sociobiology(whom I have called ‘Britain’s LEWONTIN’)and the PANKSEPPs’ present critique of evolutionarypsychology. One reason for this could be that ROSEis, after all, a neurospsychologist with a ‘realist’ bent.Still, in both ROSE and the PANKSEPPs, I see somethinglike a cluster of similar critical ideas. Just like thePANKSEPPs, ROSE criticizes the evolutionary psychologists’conception of the brain as an information processingmachine. Brains/minds do not only dealwith information, ROSE points out: they deal withmeaning. And meanings involve emotions! ROSE goeson to question the idea that modules have evolved‘quasi independently’ and that they are primary,that is, that they underlie proximal mechanismssuch as motivation. Finally, ROSE points to the survivalvalue of emotion and invokes recent researchin ‘the emotional brain’ by people such as AntonioDAMASIO and Joseph LEDOUX.What I perceive here is an emerging alliance betweenethologists and left-wing critics of sociobiologyand evolutionary psychology. It seems that thecritics of sociobiology, in their resistance to geneselectionism,have grasped for a more ‘holistic’ and‘pluralistic’ paradigm—and voilà, they have foundethology! This means that the critics of sociobiologyand evolutionary psychology have come a long wayfrom their initial position, which seemed to dismisseven the thought of an evolutionary basis for humanbehavior (cf. for instance, ALLEN et al. 1975).And look what more we find! Another element inthe evolving new moral–cum–scientific vision ofthe new opposition appears to be a belief in groupselection, that heretic view so irritating to adherentsof gene selectionism (cf. SEGERSTRÅLE 2000, pp383–384). This belief has certainly been stimulated bythe new study of SOBER/WILSON (1998). So, here wenow have ROSE, always trendy, topping off his critiqueof evolutionary psychology with an argumentfor group-selection. The PANKSEPPs, too, seem to likegroup selection, a way of thinking clearly compatiblewith a (LORENZIAN) ethological model.But criticism of evolutionary psychology doesnot only come from the earlier critics of sociobiology.It comes from a number of directions: developmentallyoriented biologists and psychologists (e.g.,BATESON/MARTIN 2000), biological systems theorists(e.g., OYAMA 2000), a variety of researchers and theoristsof cognitive processes (e.g., VAN GELDER/PORT1995; HUTCHINS 1996), and, finally, even one type ofcognitive scientists: students of developmental cognitiveneuroscience (e.g., JOHNSON 1997). These criticstypically argue that the computational model ofthe mind does not cover many important aspects ofthe mind and living systems, such as internal regulation,coordination, and feed-back mechanisms, orthey protest that human intelligence is of a complextype that simply cannot be translated into machinelanguage.Not surprisingly, researchers on developmenthave found the model used by evolutionary psychologytoo deterministic. Among other thingsthey point out that many supposedly ‘innate’ competenciesactually presuppose learning during anorganism’s lifetime (e.g., BATESON 1987; BATESON/MARTIN 2000). Still, it seems to me that they may besometimes caricaturing the evolutionary psychologists’position in order to drive home their point. Itoften seems unclear just how their own views differ,particularly since TOOBY and COSMIDES already in1992 emphasized that they do not expect modulesto be ‘hardwired’ but only to develop reliably. Thequestion is what this means. Those who study developmentmay want greater explicit recognition ofthe variety of phenotypes that can develop dependingon environmental conditions, and of the defacto unpredictability of animal and human behavior(see SEGERSTRÅLE 2000, pp378–379)—or mightthey perhaps wish to suggest that organisms‘choose’ their environments?An interesting criticism comes from a developmentalpsychologist, Annette KARMILOFF-SMITH,who challenges one of the arguments used to boostnativism and modularity: the so-called ‘double dissociation’found in WILLIAMS syndrome patients.People with this syndrome are said to be good atrecognizing faces, language, and social interactionbut have difficulty with spatial cognition, numbers,and problem-solving. KARMILOFF-SMITH studiedmore closely children and adults with this developmentaldisorder, and concluded that the assumptionthat WILLIAMS syndrome patients were otherwisenormal was incorrect. In fact, WILLIAMSsyndrome patients differed from controls in theirdealing both with faces and with language andgrammar. In regard to faces, the children instead ofadopting holistic, configurational, approach put togetherfaces feature–by–feature. This she interpretedto mean that they in ontogeny had developed anovel, compensatory strategy for dealing with information.They also followed an unusual pathwayEvolution and Cognition ❘ 10 ❘ 2001, Vol. 7, No. 1

Minding the Brainin their language acquisition. According to her, thiswas evidence for the dynamic development of thebrain and the interaction of its different parts in ontogeny(KARMILOFF-SMITH 2000, 1998, 1992).And there is more evidence for flexibility in thedeveloping brain. For instance, brain imaging findingsof face processing (in normal infants) show increasedspecialization and localization of face-processingcircuits during the first year of life.Language, too, shows progressive specialization ofthis sort: it is processed in both hemispheres untilabout the age of six when it (for right-handed people)goes to the left hemisphere (KARMILOFF-SMITH2000; JOHNSON 1997).These kinds of findings would seem to support thePANKSEPPs’ point that the modularity in the adultbrain does not reflect a pre-existing domain specificityof the brain, but is rather a product of epigenesis.Although the PANKSEPPs do not elaborate on this,one would assume that they would concur with thedistinction between two views of development: a‘regulatory’ vision as opposed to a ‘mosaic’ one.While the ‘mosaic’ kind is represented by the evolutionarypsychologists’ vision of ‘the mind as a Swissarmy knife’ and presupposes tight genetic control onepigenesis, the regulatory model involves a probabilisticepigenesis under broad genetic control, butflexible. 5 KARMILOFF-SMITH specifically suggests thatthere exists a number of different evolved learningmechanisms which during ontogeny may “each discoverinputs from the environment that are more orless suited to their form of processing”. This is whyshe calls for a closer study of the Swiss army knife’sontogenesis (KARMILOFF-SMITH 2000).More alarming, perhaps, for evolutionary psychologists—andgiving an indirect boost to thePANKSEPPs—is the recent removal of one of the cornerstonesof the whole modular theory. The very fatherof modularity, Jerry FODOR himself (FODOR1983), appears to have had a change of mind (punintended!), and now seems devoted to attackingthose who describe the mind as a collection of domain-specificspecial processing machines. This isinstantiated by his new The Mind Does Not WorkThat Way (Fodor 2000), a direct critique of StevePINKER’s (1997a) How The Mind Works.Finally, the PANKSEPPs get unexpected supportfrom someone quite sympathetic to sociobiologyand a fierce critic of its critics, anthropologistMelvin KONNER. On the face of it KONNER ‘should’ beenthusiastic about evolutionary psychology, but hetakes a surprisingly critical stance of PINKER’s 1997book (KONNER 1998). One of his main complaints isthat PINKER and evolutionary psychologists in theirmodeling totally ignore the actual anatomical structureof the brain. One cannot make the assumptionof homogenous modularity throughout the brain.In other words, the PANKSEPPs are in good criticalcompany.But our authors are not intending only to be critical.In fact, they say they wish to help evolutionarypsychology. What are, in fact, the possibilities ofbringing cognitive and affective neuroscience together?Is there a middle ground that might bereached? In light of the criticism, it would seem prudentfor evolutionary psychologists to postulate alimited number of basic modules rather than massivemodularity. Plausible candidates for more orless hardwired specialized programs come from animalstudies and research on neonates in humans. Ibelieve there is enough evidence, for instance, tospeak of a ‘face recognition module’, based on infantresearch and the demonstrated existence of‘face neurons’ in monkeys and sheep. Moreover, imitationmay be another such basic capability, as maythe set of basic emotional expressions, as demonstratedby Paul EKMAN and his associates (e.g., EK-MAN/KELTNER 1997). One indication here is that wehave all necessary facial muscles fully developed atbirth, another that neonates—even anencephalicones—react in predictable ways to sweet and soursubstances (SEGERSTRÅLE/MOLNAR 1997, pp8–9). Interestingly,even Gerald EDELMAN, otherwise committedto a neural network, ‘constructivist’ view ofthe mind, believes that there do, in fact, exist a fewbasic inborn capabilities in humans: among theseface recognition, and fundamental affect (SEGER-STRÅLE/MOLNAR 1997, p16).Many of our basic abilities do appear to be connectedto the human face and be of a nonverbal nature.It would seem that the face has important inbuiltcapabilities that will later form the basis for theinfant’s further emotional development and interactionalskills (SEGERSTRÅLE/MOLNAR 1997, p16, seealso Part III on ontogeny). Thus, one more candidatefor relatively hard-wired equipment may becertain fast-moving ‘facial affect programs’ that enableus to competently send and receive nonverbalmessages, identified by Swedish psychologist UlfDIMBERG (DIMBERG 1997). In other words, nonverbalcommunication is an important means whereby‘nature’ is transformed to ‘culture’. Also, recent researchon language emphasizes the importance ofnonverbal communication as a preparatory stage inlanguage acquisition (VELICHKOVSKY/RUMBAUGH1996).Evolution and Cognition ❘ 11 ❘ 2001, Vol. 7, No. 1

Ullica SegerstråleThe PANKSEPPs, with MACLEAN and his triunebrain, and WILSON (in Sociobiology) see emotions asprimary, because of the evolutionary history of thebrain. Still, nonverbal capabilities appear to be atthe same time emotional and cognitive. And thismay eventually help open the door for a synthesisbetween cognitive and affective neuroscience.In general, then, why not relax the modularityidea, concentrate on a few indisputable ones, andpostulate different degrees of hardwiredness for differenthuman capabilities based on the best availableempirical studies across a variety of fields, whileleaving space also for general, undifferentiatedlearning programs? Also, why not keep the questionopen as to where exactly some of these circuits arelocated and how big these modules are. (Perhapseven MACLEAN’s various brains can be seen as modules?)The important thing would seem to be to beable to pool ideas from a variety of fields: cognitiveand affective neuroscience, and a wide spectrum ofinterdisciplinary nonverbal research. That kind ofapproach would seem to go toward TOOBY andCOSMIDES–style consistency across scientific fields,‘consilience’ of the William WHEWELL type (simultaneoussupport for an idea from many fields of science)and may be even E. O. WILSON type ‘consilience’(unity of science) (WILSON 1998).Unfortunately there is a real obstacle to mutuallybeneficial cooperation between scientists from differentdisciplines. Interdisciplinary efforts may givethem no credit—on the contrary, they may be ostracizedby their ‘home’ discipline, particularly if theyare regarded as breaking certain long-standing taboos.But perhaps affective neuroscienceneeds to rethink itscriticism somewhat. Whatevermodules ‘really’ are, andhow many ‘really’ exist, thebasic theoretical contributionsof the evolutionary psychologistsare interesting andAuthor’s addressUllica Segerstråle, Department of SocialSciences, Illinois Institute of Technology,Suite 116, Siegel Hall, 3301 South DearbornStreet, Chicago IL 60616, USA.Email: segerstrale@iit.educan in principle open up new fruitful avenues of researchin other fields—including affective neuroscience.This kind of importation into one field ofideas from another is a well-known device for scientificgrowth—after all, it was physicists rather thangeneticists that created the new field of molecularbiology. What the possibilities are for a field to stimulateanother at a particular time is dependent onthe availability of individuals and their visions andambitions. Fields develop according to concrete opportunitiesperceived and particular initiativestaken—not according to how they ‘ought to’ developfrom some abstract scientific point of view.Finally, we should not forget that evolutionarypsychology is an evolving research program. There arealready signs that the evolutionary psychologists arebacking off somewhat from their initial militancy.For instance, in COSMIDES and TOOBY’s recent primerthe assumption of the Environment of EvolutionaryAdaptation seems to be relaxed, the assumption ofmodularity looks less massive, and epigenesis is moreexplicitly factored in. Emotions, too, are increasinglybrought in, a potential harbinger of the next step ofthe evolving program of evolutionary psychology. Ifthis kind of rapprochement between the evolutionarypsychologists and their critics is taking place, it isonly something that is typical of science. Indeed, inthe future, we may expect both (all?) sides to integratesome of their opponents’ arguments into theirown explanatory frameworks—without necessarilygiving their critics explicit credit. We may even see aconvergence between seemingly totally opposed positions(remember the DARWINISTS and MENDELIANSfighting each other only to beunited in the Modern Synthesis?).But if and when thishappens, we will probably nothave a synthesis with big fanfare.Academia being what itis, both sides are likely toclaim victory and get out.Notes1 While they sharply resisted this kind of research in cognitiveabilities, the critics expressed interest in finding outabout cognitive traits.2 On this topic in 1997 GOULD (1997c) got himself involvedin a furious dispute with Steven PINKER (1997b) on the pagesof The New York Review of Books.3 WILSON thinks evolutionary psychology is “the same” associobiology (personal communication), and so doesDAWKINS.4 Something that WILSON was challenged into doing by criticswho said that he did not address human variation.5 Note, however, that although her arguments supportsthat of our authors, she is actually citing data from imagingprocesses which the PANKSEPPs may consider unreliable!6 KARMILOFF-SMITH herself believes a regulatory approach ismore likely to reach the complexity required by corticalfunctions. She suggests, however, that a deterministic viewof development could be applicable to other parts of thebrain.Evolution and Cognition ❘ 12 ❘ 2001, Vol. 7, No. 1

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Linda Mealey“La plus ça change…”Response to a Critique of Evolutionary PsychologyPANKSEPP & PANKSEPPare not alone in theircriticisms of evolutionarypsychology. In fact, controversiessparked by this‘new’ discipline aboundand have even meritedfront page coverage in arecent issue of the NYTimes’ ‘Science Times’(GOODE 2000). It is clear,however, that most of thecritics, including P&P, areagainst neither evolutionnor psychology—noreven the use of evolutionaryapproaches in psychology.Indeed many,like P&P, are themselvespractitioners of whatmight be called evolutionarypsychology withoutthe capital E capital P(see e.g., MEALEY 1994, in press; MILLER 2000a,2000b; THIESSEN 1998; WILSON 1994). Criticism has,rather, focused on a particular approach taken by avery small but highly visible group of individualswho share a specific stance on a set of related issues.In their target paper, P&P variously refer to this subsetof theorists and their positions, respectively, as“a new breed of evolutionary psychologists” (p3)and “the most visible form of evolutionary psychology”(p30). Speaking as one who would prefer to seemultidisciplinary integration rather than interdisciplinaryderogation, I share P&P’s concerns aboutthe rigidity and doctrinal restrictiveness of “somecurrently fashionable versions of evolutionary psychology”(p6). But this debate is not new, and I believethat as with most controversies in science, itssparks have propelled us more steps forward thanback.AbstractPANKSEPP & PANKSEPP are not alone in their criticismsof Evolutionary Psychology. It is clear, however, thatmost of the critics, including P&P, are against neitherevolution nor psychology—nor even the use of evolutionaryapproaches in psychology. Criticism has,rather, focused on a particular approach taken by avery small but highly visible group of individuals whoshare a specific stance on a set of related issues. Ishare P&P’s concerns about the rigidity and doctrinalrestrictiveness of “some currently fashionable versionsof evolutionary psychology”. But this debate isnot new, and I believe that as with most controversiesin science, its sparks have propelled us more steps forwardthan back.Key wordsCognition, comparative method, evolutionary psychology,modularity, philosophy of science, sciencestudies.The Seven SinsThe environment ofevolutionary adaptedness(EEA)John TOOBY, LedaCOSMIDES, David BUSS,Randy THORNHILL, andothers, describe evolutionarypsychology as akind of ‘reverse engineering’through which wetry to decipher the componentsand workings ofthe human brain/mindby analyzing the types of‘problems’ it has evolvedto ‘solve’ in its ‘environmentof evolutionary adaptation’(e.g., BUSS 1995;COSMIDES/TOOBY 1997;THORNHILL 1997; TOOBY1999; TOOBY/COSMIDES 1990, 1992). Of course, thehuman brain/mind has been evolving ever sincethere was first a neuron, and the problems that ithas evolved to solve have waxed and waned and reversedand conflicted and interacted in countlessways over that immense span of time. The EEA ofthe human brain/mind is thus, not a single definitiveenvironment that we might reconstruct as wemight construct a museum diorama, but rather, amultidimensional hyperspace of the net vectors ofevery selection pressure that ever impinged uponour ancestors (DALY/WILSON 1999; MEALEY 2000).Stated as such, the EEA is a potent concept. Unfortunately,it is a concept that has been trivialized andreduced to a much more simplistic but manageablenotion, i.e., ‘the Pleistocene’, which, as a heuristicfor making novel hypotheses about the brain/mind,is virtually useless. This common substitution of aEvolution and Cognition ❘ 14 ❘ 2001, Vol. 7, No. 1

“La plus ça change…”simple concept for a more complex one needs to bepointed out—much as does the reification of theterm ‘gene’ in the commonly used phrase ‘a gene forx behavior’—but however unfortunate such substitutionmay be, it is a ubiquitous phenomenon in thesocial process that is the communication of scienceand spread of knowledge (see also YOUNG/PERSELL2000). The good evolutionary psychologist mightneed to be more careful, but need not be damned forthis sin.Species-centrismLack of a true comparative foundation is, indeed, aweakness of a modern evolutionary psychology thathas forgotten the word ‘ethology’ and the name‘TINBERGEN’ (see DALY/WILSON 1999). Althoughcomparative psychology and ethology were antagonisticdisciplines through much of their early history(JAYNES 1969), most of their differences were resolvedwith the development and joint use of newtechniques in neuroscience and neuroendocrinology(HINDE 1966); even more recently, progress incomputer modeling and evolutionary taxonomyhave made comparative analyses both easier andmore valuable (HALL 1994; HARVEY/PURVIS 1991;LARSON/LOSOS 1996). Still, comparative psychologyand human ethology seem to be undergoing majordecline—or, if not decline, at least further isolationfrom what is now mainstream psychology: conferencesand journals which once attracted and evenshowcased research on both human and other speciesnow tend to specialize in one or the other. (Seediscussions in HIRSCH 1987 and MEALEY 2001.)A case in point to illustrate the consequences ofthis neglect can be drawn from a highly visible researcharea: language. According to evolutionarypsychologists, language is the prime exemplar of arecently-evolved, complex, but uniquely humanfaculty (PINKER 1994, 1997a, 1997b). Unfortunately,to many this status implies that comparative analysisis useless. Indeed, after the once-popular ape studiesdocumented the species-specific nature of humanlanguage, comparative studies virtuallydisappeared. Yet the notion that our language capacityhas its basis in a set of modular functions is onethat absolutely begs for comparative analysis. For example,for decades psycholinguists cited the phenomenonof ‘categorical perception’ of humanspeech sounds as evidence for the specialness of humanlanguage (EIMAS 1985; EIMAS/MILLER 1992;WERKER/LALONDE 1988; LENNEBERG 1967; LIBERMAN/MATTINGLY 1989; ZATORRE et al. 1992) without botheringto determine whether that particular componentof language was indeed, human-specific. Comparativestudies, however, show that it is not (HAUSER1996, 2000); categorical perception is, rather, a preadaptionfor language that has its basis in some otherfeature or function of the brain. Of course, this insightdoes not negate the argument that human languageis special and requires study of human subjects—butit does highlight P&P’s argument thatevolutionary psychologists, by being too speciescentric,have ignored a tremendous resource for hypothesis-generationand hypothesis-testing, as wellas a huge and valuable extant literature (e.g., HAUSER1996, LIEBERMAN 1977).AdaptationismThe possibility that language emerged from a set ofpre-adaptations, or that language itself is a ‘spandrel’(GOULD 1991; GOULD/LEWONTIN 1979) hasbeen hotly debated (PINKER/BLOOM 1990). I happento think that language is not a spandrel. But I alsohappen to think that others of our most complex attributesmight be: specifically, our intelligence andour capacity for self-reflection, which I believe arelikely to be spandrels of our Theory of Mind.My own opinion about what is or isn’t a spandrelis not particularly relevant here, but the idea thatvery complex seemingly designed functions and processesmight be spandrels is, as P&P claim, an ideathat is not only disregarded by most evolutionarypsychologists, but unfairly attacked and evenmocked (e.g., ALCOCK 1998). If evolved features ofthe human brain/mind are likely to be modular asP&P’s brand of evolutionary psychologist claims,then indeed, the most likely place for us to search forspandrels is amongst those features that seem themost generalized and un-modular… such as the ‘g–factor’ of intelligence and the ‘binding factor’ or‘seamless nature’ of self-awareness.Massive modularitySince the structure of the mind is, in some sense, dependentupon the structure of the brain that housesit, it might be the case that some functional mechanismsof the mind map directly onto certain structuresof the brain. While such direct mapping is nota prerequisite for psychic modularity (FODOR 1983),it is plausible (SHALLICE 1988, 1991), and the factthat different parts of the brain do exhibit functionalspecialization has been known for over ahundred years. Not only are specialized structuresEvolution and Cognition ❘ 15 ❘ 2001, Vol. 7, No. 1

“La plus ça change…”1991), and ‘less evidence’ of cortical modularitydoesn’t mean ‘no evidence’: there is room here forempirical investigation (see ARBIB/ERDI 2000 andPAGE 2000). Cortical localization of function hasbeen documented from many studies of humanbrain damage (e.g., COSMIDES/TOOBY 2000; GRODZIN-SKY 2000; HART/BERNDT/CARAMAZZA 1985; KOLB/WISHAW 1990; SACKS 1985; TRANEL/DAMASIO 1985),as well as in intact, non-human animals (e.g., KEN-DRICK/BALDWIN 1987). Whether these specialized circuitsprovide inputs or outputs or both, or are consideredto be processing and producing affects orcognitions, are still questions that are open to debate.Is face recognition purely cognitive? Is it aninput or is it an output? What about recognition offacial expressions?Anti-organic biasThese questions inevitably lead P&P to reframe bothemotions and cognitions in terms of organic, ratherthan computational representations. I think this isimportant to do. While there are likely to be trulycomputational processes in the brain/mind (e.g.,GALLISTEL 1995), there are also likely to be analogprocesses (e.g., CARMAN/WELCH 1992) and truly organic‘embodied’ representations (see HUMPHREY1992). I suspect that the computational metaphor,though widely used by evolutionary psychologists,is likely to be just metaphor much of the time.But even if the metaphor is literally true more oftenthan not, computation is not sufficient. I suspectthat P&P are correct when they assert that “the foundationsof mind are fundamentally‘embodied’ by organicprocesses that areimpossible to compute exceptin the most superficial way”.Once again, however, this isan old debate which won’t besettled any time soon (see e.g.,FODOR 1981; SEARLE 1981,1990; WELLS 1998).Author’s addressWhat Penance?Linda Mealey, Psychology Department Collegeof St Benedict, St Joseph, MN 56374,USA, and School of Psychology, Univ. ofQueensland Brisbane, Qld 4072 Australia.Email: lmealey@csbsju.eduThe terms ‘environment of evolutionary adaptedness’,‘encapsulated modules’, ‘computations’—and for that matter, ‘representations’—were putforth as short-hand notations for things that we donot yet fully understand and therefore cannot yetadequately describe. Like all metaphors and analogies,they have their limits—and while we do needto take care not to push those limits and still believethat we are speaking ‘truth’, I think that theirworth must be assessed not on their ultimate abilityto ‘reflect’ reality, but on their practical value. Iftheir use encourages inquiry, they are a good thing;if they discourage inquiry, they are a bad thing.On the whole, I think such metaphors have encouragedinquiry and that they are, therefore, useful(see e.g., MEALEY/DAOOD/KRAGE 1996; MURPHY/STICH 2000). DARWIN’s entire book “On the Originof Species by Natural Selection” (1859) was an analogy,and although that analogy led to several misunderstandings—includingHerbert SPENCER’s ‘survivalof the fittest’—we nevertheless, still find valuein DARWIN’s choice of words. The concept of a geneas a ‘particulate entity’ was, too, an analogy—ananalogy that may have led MENDEL to overlook thereality of linkage—but an analogy which we still usetoday.Debates about terminology permeate all disciplines,including other subdisciplines of evolutionarybiology (e.g., GHISELIN 1981) and psychology(e.g., WIDIGER/SANKIS 2000). As elsewhere, the metaphorsof modern evolutionary psychology willsurvive as long as they areuseful. Eventually they willbe weeded out through anotherform of natural selection(SKINNER 1981; VAN PAR-IJS 1981), to be replaced bynew metaphors that are moreuseful in the successive intellectualenvironment. La plusça change…Evolution and Cognition ❘ 17 ❘ 2001, Vol. 7, No. 1

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Carlos StegmannComments on Jaak Panksepp and Jules B.Panksepp’s Paper “The Seven Sins ofEvolutionary Psychology”IHAVE READ WITH GREATinterest Jaak and JulesB. PANKSEPP’s paper “TheSeven Sins of EvolutionaryPsychology”, whosemain thesis meets withmy full agreement, to wit,that evolutionary psychologyshould fully consider,and be consistentwith, the findings of cognitiveneuroscience andbrain research, to “remainon the shores of soundscientific inquiry”.I would like to extendthis thesis: any disciplinemust fully consider, andbe consistent with, other,related disciplines, to producecredible results. Thisapplies especially in therealm of human natureand behavior, which arethe subject of many disciplines,among them sociology,political and legalscience, behavioral biologyand comparative behavioralresearch, moralscience, and epistemology, in addition to the twodisciplines that are the subject of the paper.In a paper which I have submitted to Evolution &Cognition, “The Human Behavior Instinct. How Decisionsfor Action are Reached. An InterdisciplinaryTheory of Human Behavior”, I have posited the followinggeneral postulate to account for the requiredinterdisciplinarity:AbstractThe commented paper states that, “to remain on theshores of sound scientific inquiry” evolutionary psychologyshould be consistent with the findings of cognitiveneuroscience and brain research. Thisargument is extended in the present comments. Thehuman species is an integrated natural entity composedof strata, including the strata of socioculturalgroup behavior, of individual behavior, and of neuralmechanisms. Events or phenomena affecting the entityare one–and–the–same in all and each of thestrata, and thus any factually correct explanation ofhuman nature, including psychology, must be interdisciplinaryin scope. Psychological phenomena mustbe explainable in terms of reductions of laws whichgovern the sociocultural stratum, such as those discoveredby Friedrich HAYEK related to the extended orderof human society, and be reducible in turn to themechanisms that operate in the neurophysiologicalstratum. The comments conclude that only such aninterdisciplinary approach can provide a compass tonavigate in the dark sea of human brain and mindphenomena.Key wordsInterdisciplinarity, psychology, cognition, human behavior,evolution, neuropsychology.“Living beings, includingHomo sapiens, are integratednatural units orentities (systemic wholes)composed of strata, fromthe atomic-particle to thesocio–cultural strata. Stratum-specificnatural lawsapply in each stratum,but events or phenomena,which affect the entity,are one–and–the–same event or phenomenonin all and each of thestrata. Since the event orphenomenon is one–and–the–same, there cannotbe any ‘causation’ betweenstrata, but the naturallaws which apply ineach stratum must be reducibleto the laws whichapply in the hierarchicallylower strata; the ideaof natural law only makessense in this context. Anotherconsequence is thatall the strata of an integratedunit evolve together;there can be nostrata with differing rates of evolution, as of societyand of the human beings which compose it—theevolution of both is one–and–the–same phenomenon.”The commented paper states (p125) that “groupselection” is different from “individual selection”:“Surely, differential survival of groups may lead todifferential survival of brain mechanisms that onlyEvolution and Cognition ❘ 20 ❘ 2001, Vol. 7, No. 1

Comments on Jaak Panksepp and Jules B. Panksepp’s Paper “The Seven Sins of Evolutionary Psychology”operate efficiently in groups. Such effects mayemerge more rapidly via group selection than by individualselection.” (My Italics.)But the groups and the individuals that composethem, and the behavioral phenomena, which affectboth groups and individuals, are one–and–the–samething. However, the natural laws, which govern thebehavior of groups, must be reducible to the naturallaws, which govern the behavior of the individualsthat compose them. This applies to all the integratednatural entities of living beings, but is especially relevantfor the natural entity homo sapiens. For thisentity possesses an upper stratum of socio–culturalphenomena, which the other entities do not, exceptingmay be some rudiments in the societies of greatapes. And here another critique may be added to theobjection that evolutionary psychologists (and sociobiologistsas well) do not sufficiently consider therelevance of brain research in their inquiries. Thephenomena investigated by psychology occur in theindividual behavioral stratum of the human entity,which is interposed between the stratum of neurophysiologicalphenomena, and the stratum of groupbehavior in human society. This stratum has beenthe object of Friedrich A. HAYEK’s work (1982, 1988).HAYEK created the concept of the spontaneous evolutionaryformation of the general laws of conductthat govern the extended order of human society. Psychology,especially evolutionary psychology, shouldconsider the relevance of these findings at the levelof human society, and be able to provide their reductionto individual human behavior. Besides, individualbehavior is also the object of comparative behavioralresearch, (LORENZ 1973, 1978; RIEDL 1988), andof moral science (Immanuel KANT 1904) 1 . It wouldalso be worthwhile to consider, for finding out howthe brain/mind works, the conclusions of epistemology,especially those related to analytic and syntheticcognition (CARNAP 1995). Psychologists, however,seem to be working almost exclusively withinthe limits of their own scientific realm.In the final part of the commented paper (p125)the following question is posed:“How do we incorporate the strikingly relevantevidence culled from our fellow animals into the exceedinglyanthropocentric modes of thought thatcharacterize so much of present day psychology?”I do not think that psychology’s modes of thoughtare “exceedingly anthropocentric”. Psychology is,after all, a human science; its object is the explanationof human behavior. Psychology, to be effective,must clearly incorporate in its research “the underlyingbrain processes shared by all mammals”(p125). But it must also incorporate the findings ofcomparative behavioral research, of moral science,and of epistemology, and be effective in providingexplanations (reductions) of the socio–cultural behaviorof groups. The referred question is followedimmediately by another one:“How shall we accept our animal heritage withoutdemeaning our vast intelligence?”But our “vast intelligence” is included in our “animalheritage”! As conclusively demonstrated by AdolfHESCHL in his book “The Intelligent Genome”(1998), the only possible mechanism of creating informationis the random mutation–and–selectionmechanism of evolution, the genome being its solerepository. All existing information is innate, andthis information includes the structural and operational‘design’ of those vast brain and nerve systemsof our species, where our “vast intelligence” originates.The general overall proposition of the commentedpaper is that explanations of human behaviorshould arise from the interaction between theobjective cognitive abilities which reside in the massivegeneral-purpose human neocortex, and theemotional and motivational systems which reside inthe very old subcortical brain areas, and not, as proposedby evolutionary psychology, from any “finegrainedmolding of special-purpose socio-affectivemechanisms”, or “new and refined emotional modularfunctions that would have emerged in the humanbrain/mind during the past several millionyears of human brain evolution”. I quote several passagesof the paper, related to this proposition, towhich many more may be added:On p108: “…we must remember to be especiallycautious in ascribing discrete special-purpose functionsto brain association areas that appear at birthto be largely general-purpose ‘computational’ devices.On p110: “The organization of the neocortex, althoughstill constrained by many unknown geneticrules, … may be much more of a general-purposecomputational device than modern evolutionarypsychologists have been willing to concede”, and“The possibility is remote … that many unique anddetailed epistemological engravings of sociobiologicalstrategies (i.e., modules) exist within the humanneocortex.”On p113: “It seems to us that much of brain evolutionduring the Pliocene and Pleistocene eras wasbased upon the rapid expansion of general-purposecortico-computational space … rather than on anyfine-grained molding of special-purpose socio-affec-Evolution and Cognition ❘ 21 ❘ 2001, Vol. 7, No. 1

Carlos Stegmanntive mechanisms. Most special-purpose mechanismsin the brain, of which there are many in subcorticalregions, evolved long before humans emerged ascontenders for the top ‘predator’ position in thefeeding hierarchy.”On p122: “The existence of various emotional andmotivational feelings, along with general-purposelearning systems, can provide practically all thetypes of adaptive behavioral strategies that havebeen discussed by evolutionary psychologists.”And on p125: “In our estimation, the type of psychologicalfunctions that evolutionary psychologistsspeak of, arise largely from the utilization ofvery old emotional capacities working in concertwith newly evolved inductive abilities supported bythe vast general-purpose neocortical associationareas.”I fully agree with all those statements. But if the‘modular’ explanation of evolutionary psychology isnot accepted, what other explanation can be proposed?What the paper proposes is a general approach,but it does not suggest a possible concreteexplanation that would substitute for the ‘modules’of evolutionary psychology. As I said at the beginning,a more thorough explanation can only arisefrom a comprehensive interdisciplinary approach,in which all the strata of the integrated natural entityHomo sapiens are considered. The main concepts,which govern such an interdisciplinary approach,may be put as follows:The phylogenetic evolution which culminatedwith the appearance of homo sapiens must haveincluded several instances of the phenomenon‘fulguration’ (LORENZ 1973, 1978), whereby,through the combination of various elements, anew category of existence comes into being, theproperties of which could not have been predictedfrom the properties of the composing elements.The new entity now possesses a new upper stratum,but it is still the integrated whole of all the stratathat compose it, from the atomic-molecular to thespiritual. Certain natural laws act in every stratum,and explanations or reductions must be consideredpossible. The new species is distinguished, chiefly,by the possession of a new cognitive apparatus. Thisapparatus operates with two different mechanismsor processes:1. The mechanism or process of objective cognition,usually called reason, which contains the analyticfaculties which permit to derive synthetic knowledgeabout reality (CARNAP 1995), combined withthe faculty of language and a greatly expandedmemory, and which superseded (but still includes),as a result of the fulguration, the ratiomorphfaculties of the pre-human ancestor;2. The mechanism or process of moral cognition,usually called conscience, which relates objectivelyknown options for action to, also objectivelyknown, values, which in turn are related to certainfeelings which the apparatus generates to make usact (DAMASIO 1994), and which superseded (butstill includes), as a result of the fulguration, theaction-governing instincts of the pre-human ancestor.This agrees with the following statement of thecommented paper (p116):“…the classic distinction between emotional andcognitive processes is sustained by abundant dataindicating that the two can be dissociated functionallyand anatomically…”.At the behavioral psychological level, human decisionmaking is based on beliefs. In objective cognition,a belief is the conviction of the truth of a statement,that is, of the correspondence of the statementwith the objective fact that it intends to describe.The objective fact may refer either to the propertiesof an object, as in scientific discovery, but also, e.g.,in engineering design, in business administration,and in many other practical fields, to the objectivemeans and ways to attain a purpose, in a range thatextends from the many everyday decisions for actionwe take on the spur of a moment, to the elaborateplans based on PERT or CPM methods in largeenterprises. In moral cognition, a belief is the convictionof the rightness of instrumental and end values,i.e., moral general norms of conduct, and institutions,which apply generally to all, like democracyor socialism. These moral convictions constitute ageneral framework for our decisions for action.A belief is actually the feeling of the emotionswhich govern our decisions for action, and which,as said in the commented paper, “arise largely fromthe utilization of very old emotional capacities” inthe ancient subcortical structures of the brain, andis attached, or affixed, to objectively known conceptsin “the vast general-purpose neocortical associationareas”.In objective cognition a scientist is convinced ofthe truth of his or her theory, and an executive of thecorrectness of his or her approach to a practical problem.However, although both hold on firmly to theirbeliefs, they will change them if well-argued fundamentalcriticism makes this necessary. Thus, evolutionarypsychologists may change the approach totheir subject as a result of the PANKSEPP’s critical observations,or an engineer his manual methods forEvolution and Cognition ❘ 22 ❘ 2001, Vol. 7, No. 1

Comments on Jaak Panksepp and Jules B. Panksepp’s Paper “The Seven Sins of Evolutionary Psychology”computer-aided calculations. Objective beliefs arethus always reversible, as objective cognition evolvesby prediction and critique. It can be argued that inclusive-fitnessevolution has developed the reversiblefeature of objective beliefs to govern collaborationbetween individuals and groups, in adaptationto the ‘prediction–and–critique’ feature of this cognitivefield.In moral cognition, however, people hold on adamantlyto their convictions in the rightness of instrumentaland end values. They will not changetheir ways of behavior and institutions, no matterwhat kinds of arguments are employed to this end.This is an historical fact. Early Christians preferredmartyrdom, Giordano BRUNO the stake, SOCRATES thehemlock, so many patriots torture and death, to renouncingtheir beliefs. Moral values are not objectivelydemonstrable. To try to do so is the naturalisticfallacy. This fallacy can be expressed in objectiveterms through the assessment that any change of theinstrumental and end values (general laws of behaviorand institutional aims which govern the extendedorder of society) will provoke so many intricatelyinterwoven direct, indirect, and retroactiveeffects, that their analysis would involve combinatorialnumbers of astronomical dimensions, andthus make a prediction of their overall effects on thestructure of the extended order all but impossible. Itis only ex–post–facto, that is, after the practical applicationof the new value for a period of time, thatits appropriateness or inappropriateness can be demonstrated.Moral beliefs arethus irreversible, as moral cognitionevolves by trial–and–error.It can be argued that inclusive-fitnessevolution hasdeveloped the irreversible featureof moral beliefs to governAuthor’s addressCarlos Stegmann, Rua dos Pessegueiros,101; 05673 – 010 Sao Paulo, Brazil.Email: stegmann@amcham.com.brcompetition between individuals and groups, in adaptationto the ‘trial–and–error’ feature of this cognitivefield.The question now must be asked of how beliefsand the corresponding feelings/emotions areformed. At the behavioral level, beliefs are ‘imprinted’2 in a universal behavioral program, whichcan be called ‘the human behavior instinct’ (it is similarto the universal grammar program, or ‘the languageinstinct” (PINKER 1994), where words are imprinted).The phylogenetically evolved program isreceptive to, and will recognize, or ‘learn’ the culturalinstructions for behavior provided by groupmembers. At the neurophysiological level, the beliefscorrespond to the formation of ‘somatic markers’postulated by Antonio DAMASIO. 3 Peer groups(HARRIS 1998) provide the ‘learning’ of social behavior,and this is one more example of how the behaviorof groups, and of the individuals that composethem, is one–and–the–same phenomenon. It can bepredicted that further neuropsychological investigationswill reveal the existence of two kinds of somaticmarkers: reversible ones, corresponding to objectivecognition (heteronomy) and to KANT’s ‘hypotheticalimperatives’, and irreversible ones, corresponding tomoral cognition (autonomy) and to KANT’s ‘categoricalimperative’.In conclusion, I would like to state again that inmy opinion a more thorough explanation of humanbehavior can only arise from a comprehensive interdisciplinaryapproach, in which all the strata of theintegrated natural entityHomo sapiens are considered.It is my considerate belief thatonly such an approach canprovide a compass to navigatein the dark sea of human brainand mind phenomena.Notes1 KANT’s discovery of the categories of moral behavior – autonomyand heteronomy, is as valid today as in 1786, becausethese categories represent innate characteristics ofthe human behavior apparatus.2 From Rupert RIEDL, “Biology of Knowledge” (1988, p292):“As imprinting we consider that special part of a learningprocess whereby the learning content can be assimilatedonly during a [certain short] phase of the biologic development,and remains irreversibly engraved thereafter. Someorganisms learn by i. the images of their parents or sexualpartners. The basic physiologic mechanism is open for anylearning content, thus facilitating experimental procedureswith organisms which possess this faculty. The mechanismmay likewise be extended to human beings, who may besaid to be imprinted by the conditions of their civilization.”(The “conditions of their civilization” are imprinted as beliefsin the human mind.)3 Antonio DAMASIO, with his theory of “somatic markers”(DAMASIO 1994; DAMASIO et al. 1994), has proposed themechanism of the interaction between objective cognitionand decision-governing feelings/emotions (beliefs). The referredworks are not included in PANKSEPP’s references. Theinterdisciplinary interpretation (reduction) of DAMASIO’sfindings (they were confirmed by neuropsychological tests)follows from the interdisciplinary relations extendedorder → individual behavior → somatic markers.Evolution and Cognition ❘ 23 ❘ 2001, Vol. 7, No. 1

Carlos StegmannReferencesCarnap, R. (1995) An Introduction to the Philosophy of Science.Edited by M. Gardner. Dover Publications Inc: MineolaNY. Originally published in 1966.Damasio, A. (1994) Descartes’ Error. G. P. Putnam’s Sons:New YorkDamasio, H. et al. (1994) The Return of Phineas Gage: Cluesabout the Brain from the Skull of a Famous Patient. Science264: 1102–1105.Harris, J. R. (1998) The Nurture Assumption. The Free Press:New York.Hayek, F. A. v. (1982) Law, Legislation and Liberty. Completeedition. Routledge & Kegan Paul: London. First editions:(1973) Vol. 1, “Rules and Order”. Routledge & Kegan Paul:London. (1976) Vol. 2, “The Mirage of Social Justice”, Universityof Chicago Press: Chicago. (1979) Vol. 3, “The PoliticalOrder of a Free People”. Institute of Economic Affairs:London.Hayek, F. A. v. (1988) The Fatal Conceit—The Errors of Socialism.Routledge: London.Heschl, A. (1998) Das Intelligente Genom. Über die Entstehungdes menschlichen Geistes durch Mutation und Selektion,Springer-Verlag: Berlin, Heidelberg.Kant, I. (1904) Grundlegung zur Metaphysik der Sitten. Basedon the 2d. Edition, revised by Kant, with corrections andgrammatical rectifications by various authors. Verlag PhilippReclam Jr.: Leipzig. Originally appeared in 1786.Lorenz, K. (1973) Die Rückseite des Spiegels. DeutscherTaschenbuch Verlag: München.Lorenz, K. (1978) Vergleichende Verhaltensforschung.Springer-Verlag: Wien, New York.Pinker, S. (1994) The Language Instinct. Harper Collins: NewYork.Riedl, R. (1988) Biologie der Erkenntnis. Die StammesgeschichtlichenGrundlagen der Vernunft. DeutscherTaschenbuch Verlag: München. Originally appeared in1979.Evolution and Cognition ❘ 24 ❘ 2001, Vol. 7, No. 1

Russell Gardner, Jr.Affective Neuroscience,Psychiatry, and SociophysiologyAffectiveNeuroscienceMy reaction to PANKSEPPand PANKSEPP (2000)reflects the viewpoint of aU.S. psychiatric educator,clinician and researcherwho finds their approachhighly compatible with aneeded nascent basic scienceframework of thepsychiatric medical specialty,as well as its alliedhuman clinical servicesdisciplines. This sociophysiologicalframeworkfor understanding thepathogenesis of psychiatricillness (GARDNER 1996,1997) depends in part onthe concept of basic plansfor communicationalbehavior with conspecifics(members of the samespecies) originating inancient brain structureshundreds of millions ofyears ago. The PANKSEPPs document that evolutionarypsychologists (EP) contrastingly focus onrecently evolved modular cognition as the centralbrain advance, a focus encouraged in part by theimpressive cortical specializations seen by SPERRYand GAZZANIGA in people with operatively caused‘split-brains’. No-one denies that the human cerebralcortex is a most impressive organ, but the PANK-SEPPs point out that important human experiences—emotions—reflectthe dominating actionsof subcortical structures, including those originatingin deep time, not just recently expanded adaptationsless than a few hundred-thousand years old.AbstractThe PANKSEPPs laudably emphasize that evolutionarypsychology (EP) has not appreciated the ancient contributionsto the human brain; rather EP conflatessubcortical emotional factors with cognitions amplifiedin recent human evolution. Remarkably, despiteovervaluing the mushroomed cortex, EP pays little attentionto communicative and social facets of humandevelopment although new work shows that a significantproportion of the variance in primate brain sizestems from its correlation with species-group size.Jaak PANKSEPP’s earlier work on emotional communicationsassists the development of a basic science ofpsychiatry, a framework now sadly lacking. Instead,market forces in the absence of a comprehensive scientificrationale have led to what could be summarizedas ‘relationshipless’ values that are contrary tothe ally-formation particularly facilitated by humanbrains. In concert with PANKSEPPs’ argument, psychiatryrequires a basic science framework titled sociophysiologyentailing comparisons (similarities) aswell as contrasts of humans to other animals.Key wordsEmotion, communication, sociophysiology, psychiatry.Evolution and Cognition ❘ 25 ❘ 2001, Vol. 7, No. 1Emotional expressionsreflect communicationalbasic plans so that thePANKSEPP focus complementsthe sociophysiologicalframework.They highlight theproblems by suggestingproblems in EP thinkingare “sins”. Making definitivecomments but usingmildly humorous imageryholds no surprise becauseJaak PANKSEPP’smost recent striking researchfindings show playand laughter to be muchmore primitive mammaliancommunicationsthan heretofore suspected.With BURGDORF,he used the infant tickleresponse in the rat pup asthe means of exploringthis communicative interactionwell known tohumans (PANKSEPP/BURG-DORF 2000). They collecteddata and drew conclusions with great care,paying attention to all the alternative hypothesesthat they, colleagues and critics could generate. Thiscautious methodology lends additional power to theother arguments Jaak PANKSEPP makes.Elsewhere he emphasized the subcortical locationof neural systems that mediate seven emotion-systems;he investigated, consolidated and documentedthese in his extraordinary book, AffectiveNeuroscience (PANKSEPP 1998). Armed with originalresearch done mostly with rat subjects, he has persuasivelyargued for the existence of emotions innon-human animals and also for the relevance of

Russell Gardner, Jr.these subcortical systems to human consciousnessand other facets of thought (PANKSEPP 2000). I feelthe PANKSEPP approach fosters integration betweenthe behavioral–mental–experiential (BME) and theorgan–cellular–molecular (OCM) levels of analysis.His work exemplifies scientific “docking” betweenthe levels.By contrast, the EP thinkers fundamentally restricttheir thinking to the BME level. They use theword biology to suggest otherwise, but many ofthese thinkers refer little if at all to actual brain research.Rather, they focus primarily on parametersof behavior that they imagine solidified in a PleistoceneEra of Evolutionary Adaptedness resultingfrom genome changes that led to humans becomingdistinct from other animals, with other functionsheld in common such as eating and defense takenfor granted and less interesting. Philip LIEBERMAN(2000), who has benefited from Paul MACLEAN’s formulations(MACLEAN 1990), argued in another extraordinaryrecent book that even the human specialty,verbal language, stems in large part fromsubcortical mechanisms, rather than reflecting a bureaucracyimplicitly limited to the cerebral cortex aspopularized, by EP linguist Steven PINKER (1994), forinstance. LIEBERMAN notes EP modular theorists usecore metaphors from World War II electronics thatinfer direct sequential connections from module tomodule. But feed-forward and feedback connectionsfound in all cortical areas directly counter the presupposedsequential connections.Not only LIEBERMAN, but PANKSEPP too benefitedfrom MACLEAN’s emphasis on evolutionary originsof present day brains that focused on complex behaviorsinvolving other conspecifics. PANKSEPPworked directly with John Paul SCOTT who died in2000 and to whom the PANKSEPPs dedicate theirseven sins–paper. SCOTT pioneered the genetics ofbehavior, focusing in landmark studies on behavioraldifferences in dog breeds (SCOTT/FULLER 1965).Significantly SCOTT (1989) wrote on the evolution ofhuman social systems.Conceptually linking the BME and the OCE levelsof analysis has commanded great interest but alsoincurred great problems over the two and a half millenniasince the school of HIPPOCRATES defined thebrain as the body’s control center. Typically, as withthe EP workers focusing only on the BME level, eitherone or the other level has been featured in isolation,with the other revealed as trivial with onlyminimum information needed. For a recent exampleof a contrasting focus restricted to the OCM level,KANDEL and SQUIRE (2000) published an article withan engaging title that included, “breaking down scientificbarriers to the study of the brain and mind”.But in fact they considered only cognition and memoryon the upper level and suggested that new advancesin the dramatic breakthroughs in OCM researchwill illumine the BME level. Their toneshowed little appreciation for the need of a two-waystreet —that conceptualizations at the BME level arerequired for docking between the two levels. Yet suchare required for full meaning to such disciplines aspsychiatry and related clinical human care-givingdisciplines. Emergent properties at the more complexlevel—such as story-production—determinesmuch behavior not predictable from features of theless complex level. Beyond its cognitive and memoryaspects alone, conspecific communication representsa brain-driven activity that requires analysis.Contrasting to the KANDEL-SQUIRE approach, linguistPIATTELLI-PALMARINI (2000) opined recently in Naturethat “The brain scientists will have to know exactlywhat they are expected to find the neural basesof”.The PANKSEPPs’ “emotional” approach (pun intended)curiously returns us to what the HIPPOCRATICSchool suggested when as the brain emerged as themajor controlling center of the body during the 5 thcentury B.C.: “Men ought to know that from nothingelse but the brain comes the joys, delights, laughterand sports, and sorrows, griefs, despondency, andlamentations. And by this, in an especial manner, weacquire wisdom and knowledge, and see and hearand know what are foul and what are fair, what arebad and what are good, what are sweet and what areunsavory” (FINGER 2000). Note in this HIPPOCRATICthinking, wisdom and knowledge take second placeto affectively charged communications and feelings.I intended the above pun because I suspect that oneof the barriers to considering fully the integrationbetween the BME and OCM levels stems from strongfeelings on the part of senior neuroscientists thatemotions and other communicational topics interferewith proper science, extending to the idea thatthose who pay attention to them are necessarily unreliable,involved with fluff somehow, or willing totolerate dubious information. MACLEAN, SCOTT andJaak PANKSEPP represent a handful of pioneers whojuxtapose emotions and other conspecific communicationsas meaningful topics that integrate the twolevels. But KANDEL and SQUIRE in their listing of 20 thcentury notables omit neuroscientists MACLEAN andPANKSEPP, though both made highly significant contributions,at least if BME–OCM integrative issueswere valued.Evolution and Cognition ❘ 26 ❘ 2001, Vol. 7, No. 1

Affective Neuroscience, Psychiatry, and SociophysiologyMost psychiatrists have lived either on the BMElevel or the OCM. But the two levels should be juxtaposed,at least as an ideal. However, selective allegiancesat one or the other conceptual level tell thetumultuous 20 th century story of the medical specialty,which I will summarize briefly before movingon to how a group of psychiatrists and other scholarsthat I here represent have worked to remedy the situation.To preview these developments, the Across-Species Comparisons and Psychopathology (ASCAP)Society brainstormed for a decade, attempting appropriateconceptual docking between the BME andOCM levels. How can we rationally formulate psychiatricconditions and social distresses using theother medical specialties and their functional relationsto their basic sciences as models? That is, apathological condition is best diagnosed and treatedwhen understood as a variant of the normal conditionlike a stroke from a damaged blood vessel canbe diagnosed and treated best with knowledge ofnormal brain circulation. Sociophysiology representsthe brain physiology of social and communicativeprocesses—especially among conspecifics.The organ of interest surely cannot be limited to thecerebral cortex but involves all parts concerned withdetection, analysis and actions of other people,though the cortex of course can hardly be ignored.The sociophysiological approach to such mattersboth compares (examines similarities between) humansand other animals and also contrasts them.The large cortex, we presume, results from its storyusingcapacity (DONALD 1991), which provides greatresourcefulness for therapy, whether this primarilyrepresents pharmacotherapy, psychotherapy, both,or group and family variants.20 th Century PsychiatryFreud, a sophisticated 19 th century neurologistwhen he founded psychoanalysis, felt too little wasknown of the nervous system so he focused only onBME with the expectation that his work wouldeventually link fruitfully with the OCM (SULLOWAY1992). He relied in his mind-models on machineimageryof the 19 th century (RABINBACH 1990). Hismost important therapeutic tactic, however,involved transference, the idea that the patientbehaves with the doctor in ways that reflect previousrelationships. After the World War II exodusfrom Europe of distinguished psychoanalysts, thisarena of thought and clinical activity became theU.S. mid-century academic standard although, notbeing evidentially based in the body, the approachdid not exhibit parallelism with other medical specialties.But in recent decades, the pendulum swung to theother extreme. A major shift in psychiatry stemmedin part from the anti-analytic movement led by biochemist–psychiatristEli ROBINS after he experienceddamage from his too-authoritarian psychoanalyst.ROBINS and his articulate prolific followers fosteredan increasingly data-based form of clinical activitywith curious de-emphasis of pathogenesis. Psychoanalysishad featured pathogenesis with metaphorsindependent of brain-data; perhaps in response, theROBINS-led psychiatrists eschewed pathogenesis aspremature and to be done eventually by biochemists(an OCM level of analysis). They propounded thataccurate description of disorders and their correlatesand treatments should come first, including a freshemphasis on the major disorders such as bipolar andschizophrenic conditions. They demanded that anyclinical treatment should possess research validation.The movement omitted relevancy of normalbehavior, focusing only on the pathological.However, the American Psychiatric Association’sDiagnostic and Statistical Manual newly emerged in1980 with codified diagnoses featuring operationaldefinitions. This potentially helped BME–OCM integrationbecause it fostered naturalistic descriptions.Moreover, psychopharmacology has helped manypatients with serious maladies, and, as well, providesformulations of how the brain works with referenceto anxious and affective states as well as clarifyingbehaviorally and thought-disordered states. TheOCM level of analysis greatly benefited patients. Butpsychiatry struggles.From outside the field, distinguished physiologist–anthropologist,Jared DIAMOND, commented recentlyon academic psychiatry in a “think tank” columnof the New York Times (DIAMOND 2001).Echoing my own impressions and those of manypsychiatrist colleagues, DIAMOND states, “[A] toonarrowfocus on biological psychiatry … hurts patients.I fully accept the importance of biologicalpsychiatry, having devoted some of my own researchto problems in that area … But now the problemhas swung to the opposite extreme: psychiatrydepartments have become bastions of molecularpsychiatry, at which more time is devoted to studyingand teaching psychopharmacology than to whatare called talk therapies”. DIAMOND goes on to concludeabout transference that “Understanding [thepatient–doctor] unique relationship was one ofthe… insights that put Freud right up there withDARWIN”.Evolution and Cognition ❘ 27 ❘ 2001, Vol. 7, No. 1

Russell Gardner, Jr.The enormous swing in psychiatric practice augmentedby major changes in payment schemes producedpowerful distressing repercussions, documentedby anthropologist Tanya LUHRMAN in herbook, “Of Two Minds: The Growing Disorder inAmerican Psychiatry” (2000). In the course of thinktank deliberations in the Research Committee at theGroup for the Advancement of Psychiatry (GAP), academicchild psychiatrist Karen Dineen WAGNERsuggested that the ideal for psychopharmacologyand managed care has become “relationshipless psychiatry”.Examiners for Board Certification at theAmerican Board of Psychiatry and Neurology feelthat interpersonal skills are seriously lacking inmany observed candidates. In conclusion, therenow exist increasingly high-level awarenesses bothin and out of the field that psychiatry needs a coremodel for how the brain at its center operates. Theabove-mentioned Research Committee of the GAPsuggested that such a model would be “the socialbrain” (BAKKER et al. 2001).SociophysiologyI turn now to my compatible but different intellectualroute from that of Jaak PANKSEPP. My backgroundas an academic psychiatrist includedresearch on the ethology of sleep movement (GARD-NER/GROSSMAN 1975) and evolved to a role as educator/administrator.In the course of this, anassignment as chairman of a medical school’s curriculumcommittee affected my thinking. Theresponsibility occurred during a major curricularrevolution and involved a charge from the dean toincrease hours for educating medical students in“behavioral science,” a national trend. I of coursesupported these changes because behavioral scienceissues and interviewing skills provided foundationsfor all future physicians, not psychiatrists only. Buteven as I fostered the major changes successfully, Ialso felt cognitive dissonance because I had beenparticularly interested in medical pathogenesissince medical school and saw no evidence my felloweducators showed understanding of this for psychiatry,nor, for that matter, did my psychiatristcolleagues, even those interested in education. Notradition existed for understanding psychiatricsymptoms or illnesses as deviations from normalbrain processes, as symptoms of congestive heartfailure represent the heart muscle’s more limitedability. I realized that psychiatry’s basic scienceneeded articulation and started with the alpha communicationstypical of a manic patient (GARDNER1982). PRICE (1967) well before me had suggestedthat ancient biology of social rank hierarchy playssignificant roles in affective illness.I found with Joan and Carl GUSTAVSON that operationallydefined “manic” communications did notdiffer from those of charismatic leaders (GARDNER etal. 1985). This resonated with the idea that an alphabasic plan likely underlay the behavior, and givensimilarities of these communications to those ofdominant animals, represented factors transmittedvia ancient genes determining brain structure. Withthe concept of propensity states antedating languagein communication (psalic), I proposed that communicationand sociality foster important ancient brainstates retained in the present (GARDNER 1988,1998a). Signaling the “planful” attributes of livingmatter emphasized by MAYR (1982), psalic also refersto programmed spacings and linkages in conspecifics,fundamental aims of communication. Particularpsalics take definition from existing in (1) normalhumans, (2) psychiatrically disturbed humans, and(3) animals. Two psalics include alpha psalic, ofcourse, seen in mania, normal leadership, and animaldominance, and the related audience psalic(state of receptivity to conspecifics as in cult membership,normal audiences, and animal subordination).Sample others include mating, nurturant andnurturance-eliciting psalics.Molecular biologist C. U. M. SMITH (1993) wrotean editorial supportive of the psalic concept for theBritish Journal of Psychiatry. Support for basic planapproaches stem now from recent neurobiologicalresearch. KANDEL and SQUIRE’s neuroscience review(2000) cited the following data-based advances:“molecular machinery and electrical signaling propertiesof neurons are widely conserved across animalspecies,” that “what distinguishes one species fromanother, with respect to their cognitive abilities, isthe number of neurons and the details of their connectivity,”and that “remarkable principles of evolutionaryconservatism are emerging from the study ofnerve cells”.In 1987, I commenced publication of The Across-Species Comparisons and Psychopathology (ASCAP)Newsletter that represented a brainstorming operation.It ran monthly for twelve years with an internationalcirculation and now continues on a quarterlybasis as The ASCAP Bulletin. The ASCAPpublication also now additionally serves as the communicationalorgan for the Psychotherapy Sectionof the World Psychiatric Association. The ASCAP Societycommenced in 1991, its first president ethologistMichael CHANCE, and meets at least yearly. AS-Evolution and Cognition ❘ 28 ❘ 2001, Vol. 7, No. 1

Affective Neuroscience, Psychiatry, and SociophysiologyCAP activities include conferring the Aaron T. BeckAward to the most deserving essay on evolutionarybiology and psychopathology. Membership disciplinesand roles include psychiatry, ethology, psychology,sociology, social work, literature, philosophy,anthropology, animal behavior, veterinaryscience, sociobiology, psychotherapy, and psychiatriceducation, specifically, residency directorshipand department chairmanship. Formerly, a numberof evolutionary psychologists also counted amongstthe ranks (indeed some served as president), butsome have dropped away, sensing, I believe, that basicplans conceptions remain valued by the coremembers of the society. Thus, from its beginnings asthe organ of the ASCAP Society, the mission statementhas read as follows:“The ASCAP Society represents a group of peoplewho view forms of psychopathology in the contextof evolutionary biology and who wish to mobilizemembers and resources of various disciplines so as toenhance the further investigation and study of theconceptual and research questions involved. Thisscientific society is concerned with the basic plans ofbehavior that have evolved over millions of yearsand that have resulted in psychopathologically relatedstates. We are interested in the integration ofvarious methods of study ranging from cellular processesto individuals in groups.”Publications in refereed journals that do not reflectan EP slant have resulted from the newsletter’sbrain-storming efforts. Examples include a report onPRICE’s hierarchical theory of affective illnesses publishedinitially in the British Journal of Psychiatry(PRICE et al. 1994) and reprinted in an edited book.Another often cited report was published in theAmerican Journal of Psychotherapy (SLOMAN et al.1994). Textbooks on evolutionary or DARWINIAN psychiatryhave now emerged, written by ASCAP members(STEVENS/PRICE 1996; MCGUIRE/TROISI 1998). Abook entitled Genes on the Couch (GILBERT/BAILEY2000) stemmed from the pages of The ASCAP Newsletterand from the 1997 AS-CAP Society annual conference.To illustrate how sociophysiologyworks in clinicalpractice, I regularly explain topatients about evolutionaryAuthor’s addressRussell Gardner, Jr., 214 Du Rose Terrace,Madison, Wisconsin 53705-3323 USAEmail: rgj999@yahoo.commatters by making the following points: peopleboth contrast and compare to other animals. Wecompare (are similar) to them in that we possessmany of the same needs and instincts, as say, one’sdog. Other animals have emotions (pet ownersknow PANKSEPP to be correct). Moreover, we shiverwith cold as do other animals and experience otherautomatic reactions that stem from lower brain levels.However, we also contrast in having a largerbrain, one indeed three times heavier than thechimpanzee brain despite nearly identical genomes—1.6%difference is commonly cited—andthe human cerebral cortex flattened would cover anarea four times greater than a chimp cortex would(CALVIN 1990). What is that big brain for? FollowingDUNBAR (1996), we concluded that allies, thinkingand planning (ATP) are key helpful attributes, withallies chief among these (contrary to EP thinking). Ifone has shiver-like responses that cause trouble (impulsivity,rage, anxiety), one should look for allies,who then aid thinking and planning. Indeed we seeallies as not only physicians and therapists but also12-step programs, anger management groups,school counselors and innumerable other culturalor societal institutions. Humans rely on strangers aswell as kin. I have proposed that we possess a peoplefocused“story-using brain” (probably dependenton cortex) that also facilitates such conspecific benefits(GARDNER 1998b).I believe that these represent understandable, unfanciful,common-sensical and data-based explanations.Patients find them helpful. They do not supplantother psychotherapies, but rather augmentthem. Other psychotherapies, may, however, benefitfrom sociophysiological analysis (GARDNER 2001).In summary, I agree that EP has underemphasizedthe ancient contributions to the human brain andconflated emotional factors with cognitions augmentedby recent human evolution. We welcomeJaak PANKSEPP’s work on emotional communicationsas complementary to sociophysiology’s development.We hope that underliningboth comparisons andcontrasts of humans to otheranimals will develop a wideningbasic science frameworkthat calms the field of psychiatryand aids its needy patients.Evolution and Cognition ❘ 29 ❘ 2001, Vol. 7, No. 1

Russell Gardner, Jr.ReferencesBakker, C./Gardner, R./Koliatsos, V./Kerbeshian, J./Looney, J.G./Sutton, B./Swann, A./Verhulst, J./Wagner, K. D./Wamboldt, F./Wilson, D. R. (2000) The Social Brain: AUnifying Foundation for Psychiatry. A manuscript submittedfor publication composed by the Research Committeeof the Group for the Advancement of Psychiatry (GAP).Calvin, W. H. (1990) The Ascent of Mind: Ice Age Climatesand the Evolution of Intelligence. Bantam Books: NewYork.Diamond, J. (2001) Freud, influential yet unloved: of science’sbig 2, only Darwin gets respect these days. The NewYork Times, Feb 17, 2001: 21.Donald, M. (1991) Origins of the Modern Mind: Three Stagesin the Evolution of Culture and Cognition. Harvard UniversityPress: Cambridge MA.Dunbar R. I. M. (1996) Grooming, Gossip, and The Evolutionof Language. Harvard University Press: Cambridge MA.Finger, S. (2000) Minds Behind the Brain: A History of the Pioneersand Their Discoveries. Oxford University Press:New York.Gardner, R. (1982) Mechanisms in manic–depressive disorder:An evolutionary model. Archives of General Psychiatry39: 1436–1441.Gardner, R. (1988) Psychiatric Syndromes as Infrastructurefor Intraspecific Communication. In: Chance, M. R. A. (ed)Social Fabrics of the Mind. Lawrence Erlbaum: Hove (EastSussex), pp 197–226.Gardner, R. (1996) Psychiatry needs a basic science titled sociophysiology.Biological Psychiatry 39: 833–834Gardner, R. (1997) Sociophysiology as the basic science ofpsychiatry. Theoretical Medicine 18: 335–356Gardner, R. (1998a) The brain and communication are basicfor clinical human sciences. British Journal of Medical Psychology71: 493–508.Gardner, R. (1998b) Language and stories: Thinking of dinosaursin Mexico. The Across-Species Comparisons and Psychopathology(ASCAP) Newsletter 11(2): 20–27.Gardner, R. (2001) Exegesis of R. Michels’ “Thinking WhileListening”. The Across-Species Comparisons and Psychopathology(ASCAP) Bulletin 2(1): 10–15Gardner, R. and Grossman, W. I. (1975) Normal Motor Patternsin Sleep in Man. Chapter 3 in: Weitzman, E. D. (ed)Advances in Sleep Research, Volume II. Spectrum Publications:New York, pp. 67–107.Gardner, R./Gustavson, J. C./Gustavson, C. R. (1985) Alphabehavior in manics as a model communicational state. InvitedPresentation at Conference entitled “The Ethology ofPsychiatric Populations”, co-sponsored by The Animal BehaviorSociety & The International Society of HumanEthology, Raleigh, North Carolina.Gilbert, P./Bailey, K. G. (eds) (2000) Genes on the Couch.Taylor & Francis: Philadelphia PA.Kandel, E. R./Squires, L. R. (2000) Neuroscience: Breakingdown scientific barriers to the study of the brain and mind.Science 290: 1113–1120.Lieberman, P. (2000) Human Language and Our ReptilianBrain: The Subcortical Bases of Speech, Syntax, andThought. Harvard University Press: Cambridge MA.Luhrman, T. M. (2000) Of Two Minds: The Growing Disorderin American Psychiatry. Alfred A. Knopf: New York.MacLean, P. D. (1990) The Triune Brain in Evolution: Role ofPaleocerebral Function. Plenum: New York NY.Mayr, E. (1982) The Growth of Biological Thought: Diversity,Evolution, and Inheritance. Harvard University Press:Cambridge MA.McGuire, M. T. & Troisi, A. (1998) Darwinian Psychiatry. OxfordUniversity Press: New York.Panksepp, J. (1998) Affective Neuroscience: The Foundationsof Human and Animal Emotions. Oxford University Press:New York.Panksepp, J./Burgdorf, J. (2000) 50-kHz chirping (laughter?)in response to conditioned and unconditioned tickle-inducedreward in rats: Effects of social housing and geneticvariables. Behavioural Brain Research 115: 25–38.Panksepp, J./Panksepp, J. B. (2000) The seven sins of evolutionarypsychology. Evolution and Cognition 6(2): 108–131.Piattelli-Palmarini, M. (2000) Speaking in too many tongues.Book review of Lieberman, P. (2000) Human Language andOur Reptilian Brain. Harvard University Press: Cambridge.Nature 408: 403–404.Pinker, S. (1994) The Language Instinct: How the Mind CreatesLanguage. William Morrow: New York.Price, J. S. (1967) Hypothesis: the dominance hierarchy andthe evolution of mental illness. Lancet 2: 243–246.Price, J./Sloman, L./Gardner, R. Jr./Gilbert, P./Rohde, P.(1994) The social competition hypothesis of depression.British Journal of Psychiatry 164: 309–315. Reprinted in:Baron-Cohen, S. (ed) (1997) The Maladapted Mind: ClassicReadings in Evolutionary Psychopathology. PsychologyPress: Hove (East Sussex).Rabinbach, A. (1990) The Human Motor: Energy, Fatigue,and the Origins of Modernity. University of CaliforniaPress: Berkeley.Scott, J. P./Fuller, J. L. (1965) Dog Behavior: The Genetic Basis.University of Chicago Press: Chicago.Scott, J. P. (1989) The Evolution of Social Systems. Gordonand Breach Science Publishers: New York.Sloman, L./Price, J. S./Gilbert, P./Gardner, R. (1994) Adaptivefunction of depression: Psychotherapeutic implications.American Journal of Psychotherapy 48: 401–416.Smith, C. U. M. (1993) Evolutionary biology and psychiatry.British Journal of Psychiatry 162: 149–153.Stevens, A./Price, J. S. (1996) Evolutionary Psychiatry: A NewBeginning. London: Routledge.Sulloway, F. J. (1992) Freud: Biologist of the Mind. Beyondthe Psychoanalytic Legend. Harvard University Press:Cambridge.Evolution and Cognition ❘ 30 ❘ 2001, Vol. 7, No. 1

Gerhard MeisenbergDegrees of ModularityTHE TARGET ARTICLE BYPanksepp and Panksepp(P&P) is about theholy grail of the psychologicalsciences: to generatea description of thehuman mind that can bemapped on the descriptionof the neural circuitrystudied by neuroscientists.P&P’s questionis whether evolutionarypsychology, in the formthat emerged during the1990s, is a good startingpoint for this grand synthesisand whether it is atall compatible with whatwe already know aboutthe brain.The variety of evolutionarypsychology criticizedby P&P is based inlarge part on the writingsof Leda COSMIDES andJohn TOOBY. Its core assumptionis the principleof massive modularity,also known as the SwissArmy Knife model. It postulatesdozens to hundredsof special-purposemechanisms, each ‘designed’ by evolution to solvean adaptive problem (KENRICK/SADALLA/KEEFE 1998;PINKER 1994; TOOBY/COSMIDES 1992). According toFODOR (1983), a module is a functionally specializedprocessing device. It is informationally encapsulated,working only on a narrow set of inputs specifiedby its input connections; it produces a specialized,stereotyped, ‘shallow’ output; it is hard-wired,developing during ontogenesis without the need foreffortful learning; and its inner workings are not accessibleto introspection: it is cognitively impenetrable.AbstractIt is argued that the distinction between cognitive andmodular functions as originally proposed by J. FODOR(1983) is valid, but that two types of modular functionsshould be distinguished: (1) hard-wired ‘primary’modules that are mainly but not exclusivelysubcortical and are mainly concerned with motivation(e.g. fear conditioning); and (2) ‘modularized’functions that are mainly cortical, are shaped by recurrentsensory and/or cognitive input, and are concernedmainly with sensory analysis and motorcontrol (e.g. language). A strict distinction betweenhard-wired subcortical circuits and non-modular corticalfunctions, as proposed by Panksepp and Panksepp,is rejected. Also the ‘Swiss army knife model’ ofevolutionary psychology is held to be simplistic. Insteadit is proposed that many brain mechanisms, includingthe cognitive system, are multi-purpose.These ‘guns for hire’ are recruited by functionally specializedmodules to produce adaptive behavior. It isargued that the most useful conceptual framework isan expanded ethological model in which specializedbrain systems may use either sensory or cognitive inputand may produce either behavioral or cognitiveoutput.Key wordsEvolutionary psychology, modularity, cognitive system,exaptation.The special-purposedevices of COSMIDES andTOOBY have the essentialproperties of FODOR’smodules, with the additionalspecification thatthey evolved as adaptiveresponses to challengesthat were repeatedly encounteredin ancestral environments.However, forFODOR modularity waslimited to the sensory andmotor periphery of themind while the cognitivesystem proper was nonmodular.COSMIDES andTOOBY have dropped thisdistinction by claimingthat also cognition itself isdomain-specific, specialpurpose,and effectivelymodular. The prominentexamples for this view areCHOMSKY’s theory of universalgrammar, recentlypopularized by StevenPINKER (PINKER 1994), andthe finding that deonticreasoning routines—thatis, reasoning about socialinteractions—differ fromreasoning in other domains in adaptively meaningfulways (COSMIDES/TOOBY 1992).FODOR drew his examples of modularity from thesensory and motor systems which are in large partcortical. The modular systems that P&P are interestedin are special-purpose subcortical circuits thatare concerned with motivation and emotion. P&Pinsist that the closest we have to evolved specialpurposedevices for dealing with adaptive challengesare the subcortical mechanisms that generate actiontendencies, emotions, and affective states. They offerwhat looks like a dichotomy between subcorticalEvolution and Cognition ❘ 31 ❘ 2001, Vol. 7, No. 1

Gerhard Meisenbergmodular systems and the developmentally malleable‘association cortex’.This dichotomy is too simple to be useful. Thereare various types of functional organization in thevast expanses of association cortex. First, there is thecognitive system proper that FODOR opposes to themodules of the sensory–motor periphery. The coreconstituent of the cognitive system is working memory,with short-term memory buffers and executivecontrol systems to manipulate the information thatis held in the buffers (BADDELEY/LOGIE 1999). Presumably,the neuronal populations that form thebuffers are able to respond with sustained activitychanges to external stimuli (GOLDMAN-RAKIC 1995).The information held in the buffers at any one timecan be described as a mental representation, a mentalmodel, or a cognitive map, depending on the researchtradition and the theorist’s preferences. Thisinformation can be entered into and retrieved fromlong-term declarative memory through the medialtemporal lobe system. Presumably the mental representationsin the buffers of working memory are theonly elements of the mind that are consciously experiencedand accessible to introspection (BADDELEY/ANDRADE 1998). Modules cannot be introspectedbecause they keep no records of their activities. Intelligenceis best defined as the ability to manipulatethe information that is held in the buffers of workingmemory (KYLLONEN/CHRISTAL 1990).Association cortex is also involved in rapid, nonconscioussensory–motor processing. The elaborationof visually-guided movements in areas of theparietal cortex is the most thoroughly studied example(BATISTA et al. 1999; COLBY/DUHAMEL 1996).These functions are not ‘primary’ modules of thekind that FODOR and the evolutionary psychologistshave in mind because they are not strictly hardwired.They are learned by a neural substrate that isstrategically located and structurally pre-designedfor this kind of learning. We can call them modularized,as opposed to truly modular functions, or wecan call them ‘secondary’ modules if we like. Exceptfor their learned rather than hard-wired nature, secondarymodules have most or all of the propertiesthat FODOR uses to define (primary) modules.Modularized functions may have been learnedwith conscious effort before becoming modularized.This is certainly the case for car-driving routines.Studies on eye movement conditioning in the parietalcortex of monkeys have also produced evidencefor reinforcement systems that may possibly be usedto guide learning in the absence of cognitive input(PLATT/GLIMCHER 1999). I agree with P&P that someof the modular functions that are commonly citedby evolutionary psychologists, including the classicalexample of language, are modularized ratherthan truly modular.Even many of the instinctive behaviors studied byethologists depend on learning. Thus, in classicalimprinting the sensory template on which the keystimulus is fitted can mature only with proper sensoryinput. The goslings that became imprinted onKonrad LORENZ (LORENZ 1935) were born with acrude sensory template that specified no more thana large moving object that makes noises. What exactlymother goose looks like has to be learned. Inthis case the motor output is quite rigid, but the sensorytemplate has to be fleshed out through learning.In the case of language, a rather pliable neuralsubstrate is strategically located between the auditorycortex and the motor outputs to the vocal apparatus.This neural substrate seems to be pre-formattedfor making categorical distinctions betweensounds, words, and word meanings, and for recognizinghierarchical relationships. In deaf-mutes thissame substrate imposes categorical distinctions andhierarchical relationships on visual inputs duringthe learning of sign language (NEVILLE et al. 1998;NISHIMURA et al. 1999), in apparent violation ofFODOR’s definition that modules are informationallyencapsulated. That we are dealing with fairly generalproperties of the neural substrate, rather than theunique features of a language module, is also suggestedby the observation that compared to righthemispherethinking, left-hemisphere thinking ingeneral depends on categorical distinctions,bounded entities, and hierarchical relationships betweenthese entities (CORBALLIS 1991; DEGLIN/KINS-BOURNE 1996). This is the reason why the left hemispheresthat write target articles and commentariestry so hard to define categorical distinctions betweensubcortical circuits and association cortex, or betweenmodular and modularized systems althoughin reality we are most likely dealing with a continuum.There may be few if any brain functions thatare 100% hard-wired or 100% arbitrarily learned.It is possible that there is a tendency for undifferentiatedcortex to evolve into hard-wired modules.The primate visual system has a complex hard-wiringwith at least 30 ‘centers’ that are involved in specializedfunctions such as motion detection, colorvision, shape recognition, face recognition, and therecognition of biological movements (VAN ESSEN/DEYOE 1995). We may speculate that this vast system,which is now largely hard-wired, evolved from apoorly-differentiated substrate that originally per-Evolution and Cognition ❘ 32 ❘ 2001, Vol. 7, No. 1

Degrees of Modularityformed visual analysis in a modularized, rather thanmodular fashion.P&P claim that not a single sociobiological modulehas ever been demonstrated at the cortical level.However, there are face-selective cells in the ventralvisual stream of the temporal neocortex and relatedbrain areas that develop within months after birth(RODMAN/SCALAIDHE/GROSS 1993), probably from acrudely pre-formed circuitry that is fine-tuned by imprinting-likelearning. Even newborns prefer lookingat faces as compared to other equally complexstimuli (GOREN/SARTY/WU 1975; MAURER 1985). Theface cell circuitry and its behavioral manifestationsappear too early in development to be produced by‘nothing but’ recurrent sensory input and reinforcementhistory. There are also systems for smiling andother emotional expressions in the cingulate cortex(DAMASIO 1994, pp139–143; FRIED et al. 1998). Babiesstart smiling at faces by about 3 months of age. It isreasonable to hypothesize that this smile reflex involvesthe face cells of the ventral temporal lobe, thesmile circuitry in the cingulate cortex, and hypotheticalprojections from the face cells to the ‘smile center’.The three-month delay after birth for the onsetof smiling is not necessarily due to the need for learningby an undifferentiated neural substrate. It mayreflect the gradual maturation of one or anothercomponent in an essentially hard-wired system.Hard-wired brain circuitry need not be present at thetime of birth. It may mature at any time.It is likely that the baby’s smile reflex evolved inresponse to an adaptive challenge: the need to preventchild neglect and infanticide. The preference ofadults for smiling faces that is exploited by the infantmay well have evolved in contexts other than parentalcare. We know of other examples where signalsevolved to exploit a pre-existing bias in the receiver(RYAN 1998). Since smiling at faces is an effectiveresponse to an important adaptive challenge, the infant’ssmile reflex qualifies as a special-purpose’sociobiological’module. Apes have facial emotionalexpressions related to human smiling (GOODALL1988, pp273–276), but true human-style smilingtook shape only during the past 5 million years,since the human–chimpanzee split, most likely byjumbling up and rearranging elements from olderfacial-expression circuitry. It remains to be seenwhether infant apes direct smiling-related facial expressionsat their mothers. P&P seem to imply thatthe neocortex is too recent to have permitted theevolution of specialized hard-wired circuitry. However,neocortex of sorts has been around for morethan 100 million years. If the smile reflex couldevolve in a mere 5 million years, then there has beenplenty of time for the evolution of hard-wired circuitsin the neocortex.The example of the smile reflex suggests that thebest functional models for behavior are frequentlythose that have been developed by classical ethologyover the past century. According to ethologists, thesmile reflex is an instinctive response that is triggeredby a key stimulus (a face) that activates an internaltemplate (the face cell circuitry) which in turnis connected to a circuitry that patterns the motoroutput (the ‘smile center’ in the cingulate cortex).Since ‘instinct’ is out of fashion, we may well call theoverall circuitry a ‘module’; or we can describe theface cell circuitry and the smile center as two modulesarranged in series. In the ethological model theinput to the hard-wired circuitry is a sensory stimulusand the output is an observable behavior. In orderto accommodate conscious perception, willed actionand reasoning, we have to extend this frameworkby acknowledging that either the output of thecircuitry or its input or both may consist of the elementsof a cognitive representation. For the time being,this expanded ethological model is more usefulthan the lofty models of cognitive architecture derivedfrom research in artificial intelligence.The existence of hard-wired motivational systemsat the cortical level is indeed uncertain. Patients withdamage to the orbitofrontal cortex show behavioraldisinhibition, poor decision making and sociopathy,especially if the damage is acquired early in life(BLAIR/CIPOLOTTI 2000; DAMASIO 1994; DUFFY/CAMP-BELL 1994; PRICE et al. 1990). These patients have normalintelligence, and their working memory seemsto be intact. We can speculate that the orbitofrontalcortex analyzes input from the cognitive system andsends inhibitory signals to subcortical emotionaland motor systems. The cognitive input would consistof mental representations of intentions, emotionsand interpersonal relations in those constellationsthat would cause damage to self or others if theintentions were acted out. The experience with neurologicalpatients seems to show that proper socialconduct and even what appears to be foresight areproduced by ‘morality modules’ in the orbitofrontalcortex. These modules may also feed back into thecognitive system as part of the ‘central executive’, todirect the processing of cognitions in working memory.Other modules in the same brain area may functionas a ‘parliament of instincts’, coordinating signalsfrom subcortical motivational systems to feedinto the central executive of working memory. Otherprojections may go from the subcortical systemsEvolution and Cognition ❘ 33 ❘ 2001, Vol. 7, No. 1

Gerhard Meisenbergstraight into the buffers of working memory to produceconscious feelings. If all this seems too messyto be true, ask a neuroanatomist to describe the fiberconnections in this brain region (PRICE/CARMICHAEL/DREVETS 1996)! Of course, the extent to which orbitofrontalfunctions are truly modular or merelymodularized, and how modularization is achieved,are wide open questions.And what about the subcortical systems that P&Pchampion as the closest approximation to sociobiologicalmodules? My first objection to their descriptionis that it exaggerates the achievements of subcorticalneuroscience. True, there is an abundantliterature about the brain systems that underlie autonomicregulation, arousal, and even some specificemotional responses, but we still don’t know how itall fits together. For example, electrical self-stimulationof subcortical brain sites has been studied in ratsfor nearly half a century, since 1953. And yet, we stillhave no good description of the underlying circuitry.We don’t even know whether self-stimulation sitesin places such as the midbrain, lateral hypothalamusand septal area belong to the same reinforcementcircuitry or whether they represent functionally independentsystems. We don’t even know the exactrole of the mesolimbic dopamine system in this circuitryalthough we have drugs such as amphetaminethat can produce euphoria by stimulating this system(KALIVAS/NAKAMURA 1999; ROBBINS/EVERITT1996; SPANAGEL/WEISS 1999).There can be no reasonable doubt that the subcorticalsystems mentioned by P&P are hard-wired. Andstill, some of them are not good examples for ‘sociobiological’modules because they did not evolve intheir present form in response to a specifiable adaptivechallenge. Thus, the cells of the mesolimbicdopamine system become active when the animaleither experiences or expects reward, and they reducetheir activity when the expectation of rewardis frustrated (SCHULTZ 1998). Together with the observationthat psychostimulant drugs activate thissystem, this points to a specific involvement in positivereinforcement and possibly even the consciousexperience of pleasure. However, the same neuronsare also activated during stress, for example when arat expects a painful electrical shock or is immobilizedon the laboratory bench with duct tape (D’AN-GIO/SERRATO/SCATTON 1990; IMPERATO et al. 1992).Unless laboratory rats are masochists, the dopaminecells are unlikely to function as mediators of pleasureor positive reinforcement in these situations. We canstill save the function-specific model by claimingthat mesolimbic dopamine cells belong to an incentivesystem that is recruited whenever an active behavioralresponse to an environmental stimulus isrequired. Still, the ‘meaning’ of mesolimbic dopamineactivity in different situations may range all theway from ecstasy to panic. It all depends on activitiesthat are going on in other parts of the brain at thesame time.The serotonin system is another example. Accordingto P&P, animals appear to be “relaxed, satisfiedand confident” when serotonin is high, but they alsomention that knockout mice with markedly reducedserotonin release are less fearful in situations whereanimals normally exhibit heightened fear responses.The situation is confusing because serotonin releaseis affected in many other situations as well. It participatesin the regulation of REM sleep (MONTI/MONTI2000), and the effects of fear-inducing and stressfulstimuli on the serotonin system are so variable andsituation-dependent that any speculation about aunitary function for this neurotransmitter seemshopeless (CHAOULOFF 2000). Serotonin seems to haveso many functions that we cannot assign it to a ‘fearmodule’, a ‘sociality module’, a ‘relaxation module’,or a ‘sleep module’.Perhaps the closest subcortical approximation toa function-specific module that evolved in responseto a specific environmental challenge is the systemof conditioned fear described by LEDOUX (LEDOUX1996). In this case the input is an association betweena sensory stimulus and an unpleasant outcome,and there are well-defined, hard-wired outputsinto the somatomotor, autonomic, endocrine,and possibly cognitive systems. Presumably this systemevolved as a solution to a recurrent adaptiveproblem: to avoid situations in which unpleasantnesshas been experienced in the past. Although itaccepts cognitive as well as sensory input, it operatesautomatically and is not easily controlled by the cognitivesystem, as evidenced by the difficulty of treatingphobic patients.Taken together, the most reasonable model is thatthe brain does have adaptively meaningful hardwiredconnections that function in a modular fashion:reflex-like and instinct-like. In typical cases,such stimulus–response arcs have distinct circuitriesfor sensory analysis, such as the ‘face cells’ in theventral temporal lobe, and motor circuitries such asthe ‘smile center’ in the cingulate cortex. Hard-wiredcircuitries that mature with minimal need for specificsensory inputs—although they may depend ontrophic influences from their afferents—are moretypical for subcortical areas but do occur in corticalregions as well, especially in and around the sensoryEvolution and Cognition ❘ 34 ❘ 2001, Vol. 7, No. 1

Degrees of Modularityand motor cortices. We also have domain-generalsystems that can be recruited by the function-specificmodules. These ‘guns for hire’ include, amongothers, the serotonin system and perhaps the mesolimbicdopamine system at the subcortical level,and the cognitive system at the cortical level.The relationship between special-purpose modularsystems and the cognitive system becomes of paramountimportance when the output is not a behaviorbut a reasoning routine. Thus COSMIDES andTOOBY note that social contract reasoning does notsimply reflect a facilitation of the same type of logicalreasoning that is applied to other situations(COSMIDES/TOOBY 1992, pp187–193). They proposethe existence of two parallel reasoning systems, onefor the physical world and the other for social contractsand cheaters. However, rather than postulatingtwo entirely independent systems, it would bemore parsimonious to postulate a single cognitivesystem in which social as well as physical relationshipscan be represented. One place to look for thedifference between deontic and indicative reasoningare the inputs of working memory, which consistentirely of the pre-chewed products of modular systems:sensory cortex for information about the externalworld, and subcortical circuitry for informationabout emotional valence and relevance forbehavior. If the sensory systems can analyze theirinputs for color, face identity, and biological motion,why not for conformity to the stereotype (or archetype,since it presumably develops from pre-wiredcircuitry in all normal humans) of a social-exchangesituation? And if subcortical emotion systems canhighlight important sensory stimuli or important elementsof cognitive representations—such as thesmell of a fast food shack when you are hungry—why should they be unable to highlight the cheaterpart in the mental representation of a social interaction?Another place to look for specialized deontic circuitryis in the ‘central’ executive of working memorywhich is, most likely, rather decentralized. Wecan hypothesize that the executive system containsreasoning modules which use the cognitive representationof the social-contract situation as their inputand send their own outputs back into the cognitivesystem, for example to generate alternativescenarios about possible cheating. The empirical evidencethat deontic reasoning differs from reasoningabout the physical world in kind and not only in thedegree to which general-purpose logic is recruitedcould simply mean that some components of thecentral executive respond only to social-contract situations.This would be sufficient to provide a “guidancesystem” (TOOBY/COSMIDES 1992) for adaptivebehavior.Evolutionary psychologists may question theevolvability of brain systems that are not specificallydevoted to the solution of a single adaptive problem.However, guns for hire can evolve from functionspecificsystems when elements of these systems areco-opted for new behavioral contexts, either with orwithout the loss of their original function. Thus, thebrainstem reticular formation most likely evolved asa motor center in the first vertebrates for the purposeof coordinating swimming movements and mediatingthe impact of sensory stimuli on these movements(MASINO 1992). It lost most of its motor functionswhen motor control shifted to higher centersin midbrain, cerebellum, basal ganglia, and cerebralcortex. Coarse adjustments of muscle tone and theactivation of higher brain areas including the cerebralcortex are now its main functions, and it getsrecruited whenever these functions have to be adjusted,from sleep–wake regulation to pain responses,orientation to noises, responses to danger,and driving on New York City highways.The cognitive system, with working memory andits long-term memory system in the hippocampusand adjacent areas of the medial temporal lobe, mayhave evolved from a short-term memory system thatenabled the animal to associate stimuli that wereperceived with a time delay. A brain without shorttermmemory would not be able to associate stimulifor classical conditioning unless they are received atexactly the same time (CLARK/SQUIRE 1999). Later on,this system was elaborated as a cognitive mappingdevice to construct a mental model of the environment,for use during foraging and other activities(including taxi driving, MAGUIRE et al. 1997) that requirednavigation in the environment. Finally, it acquiredever more complex devices to manipulate themental models in working memory, thus creatingthe ability to construct models of what could berather than what is or has been.These are merely evolutionary just-so stories, butthey show that we can build evolutionary scenariosfor domain-general multi-function brain systems.The theoretical framework of contemporary evolutionarypsychology, with its emphasis on domainspecific,function-specific mechanisms, is too narrow.We have to be aware of the adaptive challengesand selective pressures that shaped our brain, but wealso have to be aware of older adaptations that werealready in place when a new challenge was encountered.We have to realize that ancient adaptationsEvolution and Cognition ❘ 35 ❘ 2001, Vol. 7, No. 1

Gerhard Meisenbergcan assume new functions and that new functionalsystems can be created by re-wiring componentsfrom older ones.Evolutionary psychologists will have to accept theexistence of multi-purpose ‘guns for hire’ in thebrain. They will also have to realize that the constructsof classical ethology are likely to prove moreproductive as guides for future research than the information-processingmetaphors derived from AI. Finally,they have to widen their outlook by dwellingnot only on adaptations that evolved, but also onthose that could have evolved but didn’t. For example,according to HAMILTON’s rule (HAMILTON 1964)you should love your brother half as much as yourselfand your half-brother one quarter as much. And yet,you will never know how much you should love yourbrother unless you send DNA from yourself, yourbrother and your putative father to the paternity lab.We never evolved a dependable, hard-wired systemfor kin recognition. Evolutionary psychologistsshould point out that this deficiency is central to theevolution of human sociality because it enables us todeploy kin-selected forms of altruism promiscuously.There are constraints on the number and complexityof genetically hard-wired brain circuits. Accordingto the latest counts, we have only a littlemore than 30,000 genes in our genome (VENTER etal. 2001). Perhaps we cannot have any more becausethe rate of detrimental mutations is so high (EYRE-WALKER/KEIGHTLEY 1999; GIANNELLI/GREEN 2000)that we wouldn’t be able to maintain a larger genome.Most of our genes are either housekeepinggenes that are expressed in most or all cells, or theysupport specialized functions of terminally differentiatedcells, or they are involved in specialized developmentalprocesses other than brain developmentsuch as building the heart, the eye, and theimmune system. There can be only a few thousandgenes at most that are devoted mainly to the constructionof specialized brain circuits, and few if anyof them may play roles in the development of oneand only one ‘module’. Because of this genetic poverty,combined with the intrinsic mindlessness ofthe evolutionary process, behavioral adaptationsmust by necessity be crudeand imperfect. Because of thisimperfection, adaptationistreasoning is only moderatelyuseful for the prediction ofcognitive–behavioral mechanismsalthough it is essentialas a reality check for psychologicaltheorizing.Author’s addressGerhard Meisenberg, Ross University Schoolof Medicine, Portsmouth Campus, Dominica(Eastern Caribbean).Email: gmeisenberg@rossmed.edu.dmEven if evolutionary psychologists widen theiroutlook along these lines, I doubt that a separatediscipline named evolutionary psychology can persist.Sociobiology and evolutionary psychology wereresponses to the biology-neglect of the mainstreampsychology and social science traditions of the 20 thcentury. As soon as scholars in the psychological andsocial sciences incorporate evolutionary thinking asa reality check for their own theories, a separate disciplineof evolutionary psychology will no longer beneeded. Its historical mission will be fulfilled.And what can neuroscientists do to approach themystic unity between their own science and the behavioralsciences? Overspecialization is a far morepressing problem than evolution-neglect. One veryspecific bug that P&P illustrate only too well, possiblywithout noticing, is the division of neuroscientistsinto anthropocentric and ratocentric camps.Most of the neurochemical research about biogenicamines, neuropeptides, and other neurotransmittersand modulators has been done in rats. Also researchon subcortical emotional systems is generally donein rats. For lack of human volunteers, anthropocentricneuroscientists do most of their work with monkeysalthough the advent of functional neuroimaginghas permitted complementary studies inhumans. Anthropocentric research concentrates onthe cortical mechanisms of sensation, motor control,memory and cognition. Subcortical mechanismsare neglected, and the study of emotion inprimates is lagging far behind the study of otherhigher brain functions.P&P stress, correctly I presume, that species differencestend to be smaller for early-evolved subcorticalsystems than late-evolved cortical systems. And yet,species differences in social bonding exist even betweenthe closely-related prairie voles and meadowvoles that they cite as examples of neurochemicallyspecific social bonding. Monogamous habits inthese species are affected by oxytocin, but even if ahomologous circuitry exists in humans, chances areit is modulated not by oxytocin but by other neurochemicals.Sure enough, many of the early-evolvedsystems, such as those concerned with fear responsesand reinforcement learning,are likely to be very similareven in rats and humans, butwe need primate studies toprove the point. The brainmechanisms for more complexemotional responses,such as guilt, shame, compassion,or romantic love, cannotEvolution and Cognition ❘ 36 ❘ 2001, Vol. 7, No. 1

Degrees of Modularityeasily be studied even in chimpanzees, let alone rats.Even if the subcortical emotion circuits turn out tobe similar in rodents and primates, the importantinterface between cognition and emotion has to bestudied in primates. Contrary to what P&P seem tobelieve, emotions are in all likelihood not experiencedat subcortical sites. The feelings that we candescribe and remember are messages that have beensent into the cortical working memory system by thesubcortical emotion circuits.ReferencesBaddeley, A. D./Andrade, J. J. (1998) Working memory andconsciousness: An empirical approach. In: Conway, M. A./Gathercole, S. E./Cornoldi, C. (eds) Theories of Memory,Vol. 2. Psychology Press: Hove (East Sussex), pp. 1–19.Baddeley, A. D./Logie, R. H. (1999) Working memory. Themultiple-component model. In: Miyake, A./Shah, P. (eds)Models of Working Memory. Mechanisms of Active Maintenanceand Executive Control. Cambridge UniversityPress: Cambridge, New York, pp. 28–61.Batista, A. P./Buneo, C. A./Snyder, L. H./Andersen, R. A.(1999) Reach plans in eye-centered coordinates. Science285: 257–260.Blair, R. J. R./Cipolotti, L. (2000) Impaired social response reversal.A case of “acquired sociopathy”. Brain 123: 1122–1141.Chaouloff, F. (2000) Serotonin, stress and corticoids. Journalof Psychopharmacology 14: 139–151.Clark, R. E./Squire, L. R. (1999) Human eyeblink classicalconditioning: effects of manipulating awareness of thestimulus contingencies. Psychological Science 10: 14–18.Colby, C. L./Duhamel, J.-R. (1996) Spatial representations foraction in parietal cortex. Cognitive Brain Research 5: 105–115.Corballis, M. C. (1991) The Lopsided Ape. Evolution of theGenerative Mind. Oxford University Press: New York, Oxford.Cosmides, L./Tooby, J. (1992) Cognitive adaptations for socialexchange. In: Barkow, J. H./Cosmides, L./Tooby, J.(eds) The Adapted Mind. Evolutionary Psychology and theGeneration of Culture. Oxford University Press: New York,Oxford, pp. 163–228.Damasio, A. (1994) Descartes’ Error. Emotion, Reason, andthe Human Brain. G. P. Putnam’s Sons: New York.D’Angio, M. B./Serrato, A./Scatton, B. (1990) Mesocorticolimbicdopaminergic systems and emotional stress. Journalof Neuroscience Methods 34: 135–142.Deglin, V. L./Kinsbourne, M. (1996) Divergent thinkingstyles of the hemispheres: how syllogisms are solved duringtransitory hemisphere suppression. Brain & Cognition31: 285–307.Duffy, J. D./Campbell, J. J. (1994) The regional prefrontal syndromes:a theoretical and clinical overview. Journal ofNeuropsychiatry and Clinical Neurosciences 6: 379–387.van Essen, D. C./Deyoe, E. A. (1995) Concurrent processingin the primate visual cortex. In: Gazzaniga, M. S. (ed) TheCognitive Neurosciences. MIT Press: Cambridge MA, London,pp. 383–400.Eyre-Walker, A. P./Keightley, D. (1999) High genomic deleteriousmutation rates in hominids. Nature 397: 344–347.Fodor, J. (1983) The Modularity of Mind. MIT Press: CambridgeMA.Fried, I./Wilson, C. I./MacDonald, K. A./Behnke, E. J. (1998)Electric current stimulates laughter. Nature 391: 650.Giannelli, F./Green, P. M. (2000) The X chromosome and therate of deleterious mutations in humans. American Journalof Human Genetics 67: 515–517.Goldman-Rakic, P. S. (1995) Cellular basis of working memory.Neuron 14: 477–485.Goodall, J. (1988). In the Shadow of Man. Houghton Mifflin:Boston. Originally published in 1971.Goren, C. C./Sarty, M./Wu, P. Y. K. (1975) Visual followingand pattern discrimination of face-like stimuli by newborninfants. Pediatrics 59: 544–549.Hamilton, W. D. (1964) The genetical evolution of social behaviour.Journal of Theoretical Biolology 7: 1–52.Imperato, A./Angelucci, L./Casolini, P./Zocchi, A./Puglisi-Allegra,S. (1992) Repeated stressful experiences differentlyaffect limbic dopamine release during and following stress.Brain Research 577, 194–199.LeDoux, J. (1996) The Emotional Brain. Touchstone: NewYork.Kalivas, P. W./Nakamura, M. (1999) Neural systems for behavioralactivation and reward. Current Opinion in Neurobiology9: 223–227.Kenrick, D. T./Sadalla, E. K./Keefe, R. C. (1998) Evolutionarycognitive psychology: The missing heart of modern cognitivescience. In: Crawford, C./Krebs, D. L. (eds) Handbookof Evolutionary Psychology. Ideas, Issues, and Applications.Lawrence Erlbaum Ass.: Mahwah NJ, pp. 485–514.Kyllonen, P. C./Christal, R. E. (1990) Reasoning ability is (littlemore than) working-memory capacity?! Intelligence14: 389–433.Lorenz, K. (1935) Der Kumpan in der Umwelt des Vogels[Companionship in bird life]. Journal für Ornithologie 83:137–213.Maguire, E. A./Frackowiak, R. S. J./Frith, C. D. (1997) Recallingroutes around London: Activation of the right hippocampusin taxi drivers. Journal of Neuroscience 17: 7103–7110.Masino, T. (1992) The spinal motor system in early vertebratesand some of its evolutionary changes. Brain Behaviorand Evolution 40: 82–97.Maurer, D. (1985) Infants’ perception of facedness. 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Gerhard MeisenbergNishimura, H./Hashikawa, K./Doi, K./Iwaki, T./Watanabe, Y./Kusuoka, H./Nishimura, T./Kubo, T. (1999) Sign language“heard” in the auditory cortex. Nature 397: 116.Pinker, S. (1994) The Language Instinct. Morrow: New York.Platt, M. L./Glimcher, P. W. (1999) Neural correlates of decisionvariables in parietal cortex. Nature 400: 233–238.Price, B. H./Daffner, K. R./Stowe, R. M./Mesulam, M. M.(1990) The comportmental learning disabilities of earlyfrontal lobe damage. Brain 113: 1383–1393.Price, J. L./Carmichael, S. T./Drevets, W. C. (1996) Networksrelated to the orbital and medial prefrontal cortex; a substratefor emotional behavior? Progress in Brain Research107: 523–536.Robbins, T. W./Everitt, B. J. (1996) Neurobehavioural mechanismsof reward and motivation. Current Opinion in Neurobiology6: 228–236.Rodman, H. R./Scalaidhe, S. P. O./Gross, C. G. (1993) Responseproperties of neurons in temporal cortical visual areas ofinfant monkeys. Journal of Neurophysiology 70: 1115–1136.Ryan, M. J. (1998) Sexual selection, receiver biases, and theevolution of sex differences. Science 281: 1999–2003.Schultz, W. (1998) Predictive reward signal of dopamineneurons. American Journal of Physiology 80: 1–27.Spanagel, R./Weiss, F. (1999) The dopamine hypothesis of reward:Past and current status. Trends in Neurosciences 22:521–527.Tooby, J./Cosmides, L. (1992) The psychological foundationsof culture. In: Barkow, J. H./Cosmides, L./Tooby, J. (eds):The Adapted Mind. Evolutionary Psychology and the Generationof Culture. Oxford University Press: New York, Oxford,pp. 19–136.Venter, J. C. and 263 coauthors (2001) The sequence of thehuman genome. Science 291: 1304–1351.Evolution and Cognition ❘ 38 ❘ 2001, Vol. 7, No. 1

Ian PitchfordNo Evolution. No CognitionJAAK & JULES PANKSEPPpostulate a striking dichotomybetween ‘geneticallydedicated circuits’for emotions and generalpurposecomputationalspace. The former arephylogenetically ancientsubcortical structures, orneurochemical operatingsystems, which have homologiesin many species,and reflect fitness concerns;the latter is subservedby plastic neocortex.The research programof neuroevolutionarypsychobiology aims toelucidate the way inwhich human abilitiesemerge from developmentalinteractions betweenthese two mechanisms.The subcorticalemotional and motivationalsystems constitute“the essential character ofthe human mind”; theymay have distinct variantsin “different lines ofthe human family”, may owe their existence togroup selection, and can “regulate the constructionof personality differences, as well as social systems”.Evolutionary psychologists are exhorted to adoptthis model and thereby enter the ranks of the ‘dataconstrained’ instead of pursuing the ‘tradition’ ofCOSMIDES and TOOBY, which is described as a “potentiallyvirulent strain of neo-DARWINIAN thinking”.Although the PANKSEPPs describe ‘seven sins’ theseembody four complaints: evolutionary psychologistsa). search for adaptive modules where there isonly general-purpose computational space; b). areanthropocentric; c). conflate emotions and cognitions,and d). have an anti-organic bias. However,AbstractAlthough scholars in the natural and human scienceswill generally disavow any belief in distinct materialand immaterial substances contemporary debates arephrased largely in terms that would have been familiarto the Greek philosophers, and which still dividehuman characteristics into ‘divine’ or ‘transcendent’attributes—in modern terminology the surrogateterms are rational, cognitive, discursive, autonomousand creative—and ‘animal’ or ‘corporeal’ attributes—the surrogates being emotional, instinctive, determined,immutable, and bounded. This essential dualismpreserves the three key dichotomies of mind/body, cognition/emotion and nature/nurture found inmany, if not most, discussions of human nature. Neuroevolutionarypsychobiology’s concern to divide thebrain into determined affective components and unboundedplastic neocortex by employing such conceptsas ‘exaptations’, ‘spandrels’, and ‘emergence’resides firmly within this quasi-theological Westernphilosophical tradition. This dualistic approach providesno coherent foundation for the critique of evolutionarypsychology.Key wordsModularity, developmental systems theory, dualism,spandrel, exaptation, brain evolution.the PANKSEPPs readily acknowledgeevidence forthe existence of ten ormore modules in the neocortex,and evidence forthe functional integrationof cognition andemotion demonstratedby the work of DAMASIO,LEDOUX, and ROLLS. ThePANKSEPPs’ rejection ofevolutionary psychologyemerges from their beliefthat neocortical adaptationsshould be innate,discrete, and detachedfrom subcortical structures,which in turn restson theoretical misconceptionsand their mischaracterizationof theneurofoundational issues.DevelopmentalSystems Theory andInnatenessThe PANKSEPPs claim tohave assimilated the prescriptionsof developmentalsystems theory, as embodied in the work ofOYAMA (2000b) and GRIFFITHS (1997), but throughouttheir paper they refer to “genetically dedicatedcircuits” and “genetically dictated adaptations”,which suggests that they are not aware that this theoryexplicitly opposes the notion of genes as privilegedcausal entities. Indeed, OYAMA has said that“the idea that traits are ‘transmitted’ in heredity,rests on notions of genetic programming that areultimately quite preformationist” (OYAMA 2000a,p21). Following LEHRMAN’s (1953) lead, developmentalsystems theorists argue that nature selectsfor outcomes. Any of the inherited components ofthe developmental system, which includes genes,Evolution and Cognition ❘ 39 ❘ 2001, Vol. 7, No. 1

Ian Pitchfordother biological resources, patterns of maternalcare, play, and so forth can ‘mutate’ producingnovel phenotypic characteristics. In contrast to theperspective encouraged by the ‘genetic blueprint’ or‘genetic program’ metaphors, this approach allowsfor phenotypic variability, and for constant, stableoutcomes, provided that all of the resources requiredby the developmental system are available(GRIFFITHS 1997, p186). This theory is applicable tothe whole of evolutionary biology, and not only tothe neocortex as the PANKSEPPs seem to think.Perhaps the most insidious consequence of thegenetic blueprint idea is the expectation that phenotypiccharacteristics are innate, meaning ‘hereditarilydetermined’, or arising independently of environmentor experience (LEHRMAN 1953). Describinga trait as ‘innate’ confuses at least four propertiesthat can vary independently: “(1) that it is found inan individual because of their ancestry rather thantheir current environment; (2) that its growth doesnot depend on that environment for anything butbasic sustenance; (3) that it is present at birth or earlyin development; and (4) that it is part of the ‘nature’of the species… The result of this mismatch betweenconcept and reality is that when theorists discoverthat one element of the innateness concept appliesto a trait, they are liable to assume that the otherelements must also apply” (GRIFFITHS 1997, p104).Our faculties are the product of developmental systems,and consequently they are neither innate norstructured by the environment. The PANKSEPPs formulationperpetuates the very ‘nature versus nurture’dichotomy that developmental systems theoryaims to transcend.Adaptations, Exaptations, and SpandrelsThe PANKSEPPs are surely correct that we should beconfident where convergent lines of evidence pointin the same direction, which makes their assertionthat recent brain evolution is characterised by “therapid expansion of general-purpose cortico-computationalspace (which permitted the emergence offoresight, hindsight and language)” appear strikinglyincongruous. The obfuscations of those wholike to characterise themselves as opposed to ‘DAR-WINIAN fundamentalism’ (GOULD 1997) play a rolehere. The PANKSEPPs tell us that “since the emergenceof massive, general-purpose cortical space, exaptationsand spandrels have arisen everywhere welook”. Their mentor GOULD (1984) does not disagreewith biology’s emphasis on natural selection but believes“that we have become overzealous about thepower and range of selection by trying to attributeevery significant form and behavior to its direct action”.Obviously, we should not be interested in ‘attributing’anything at all to natural selection. Weneed to look at the evidence that “a function isserved with sufficient precision, economy, efficiency,etc. to rule out pure chance as an explanation”(WILLIAMS 1996, p10).In their original paper on ‘spandrels’ GOULD/LEWONTIN (1979) make entirely prosaic observationsconcerning the ubiquity of phyletic constraints, andargue that the evidence for Aztec cannibalism, thechin, and papillary ridges as adaptations is notstrong. In a second paper cited by the PANKSEPPsGOULD (1991) observes that ‘exaptations’ can be definedas characters not selected for their currentfunction which may or may not have arisen originallyby the direct action of natural selection. Usefulcharacters that did not arise by the action of naturalselection are a type of exaptation, or coopted nonaptation,called a spandrel. GOULD mentions weight,the delayed ossification of skull bones, and languageas probable exaptations.It seems clear from these examples that (with theexception of language, which appears to share littlein common with the other examples) spandrels aretrivial features that would not be identified as adaptationsby anyone using the conventional criteria. Itis equally clear that we have no good reason to restrictthe term ‘adaptation’ to only those charactersthat perform the function for which they were firstselected. As GRIFFITHS and STERELNY point out“GOULD and VRBA think that a trait is an adaptationonly for the purpose for which it was first selected.But what justifies this special status for the first ofmany selection pressures? The importance of theconcept of adaptation in biology is that it explainsthe existence of many traits of the organisms we seearound us. This explanation is not just a matter ofhow traits first arose, but of why they persisted andwhy they are still here today” (STERELNY/GRIFFITHS1999, p219). The only complex functional characteristicclaimed as an exaptation is language, andthis is done by argument from authority—the authorityin question being CHOMSKY, who is said tohave “long advocated a position corresponding tothe claim that language is an exaptation of brainstructure” (GOULD 1991, p61). However, CHOMSKYhas not “expressed views on the lack of a role fornatural selection in… the origin of language”. Onthe contrary he believes “that natural selection isoperative in this case” (personal communication1999).Evolution and Cognition ❘ 40 ❘ 2001, Vol. 7, No. 1

No Evolution. No CognitionThe PANKSEPPs are ambivalent here and acknowledgethat “there is substantial neuro-evolutionaryevidence for the emergence of special abilities suchas language”. Their rejection of it as an adaptationseems to be based in part on the misconception acquiredfrom the work of GOULD and colleagues thatonly something arising from “de novo genetic shaping”can be characterised as such. It is notable thatconnectionist models most in keeping with the general-purposearchitecture that the PANKSEPPs proposehave produced results incompatible with what weknow of language processing, acquisition, and pathology.CHOMSKY observed recently that “in thecase of language, the evidence for connectionistmodels is… about zero” (quoted in SMITH 1999,p135). The case of Christopher, a severely retardedautistic savant who remains unable to take care ofhimself, but who has mastered over twenty languages,seems particularly problematic for themodel proposed by the PANKSEPPs. Despite his abilitiesChristopher was unable to master the artificiallanguage Epun, which violates the rules of naturallanguage. Surprisingly, a control group of bright undergraduatesfared no better (SMITH/TSIMPLI 1995).Even if we were to ignore arbitrarily the other evidencefor the modularity and evolution of languageit makes no sense to interpret these findings in termsof the properties of plastic neocortex. The claim that“language emerged within a spandrel of evolvingmultimodal brain complexities” is mere sophistry.Instead of relying on the quaint notion that theirhypothesis should be taken seriously because it hasnot been falsified (just like most other hypotheses)the PANKSEPPs should provide what they require ofevolutionary psychologists: a plausible account ofthe distal and proximal mechanisms responsible forgenerating the complex functionality and patternsof acquisition and dissociation that we observe. Tocall on ‘learning’ will not suffice. Induction and explanationin science require reference to causal homeostaticmechanisms (GRIFFITHS 1997, pp187–201), not appeals to folk psychological concepts thatdo not represent projectable categories (see GRIF-FITHS 1997, pp175–176, and TOOBY/COSMIDES 1992,pp111–118, for useful discussions).Evolutionary Psychology, Selfishness,and Inclusive FitnessThe PANKSEPPs regard the theory of inclusive fitnessto be “most clearly applicable to sub-human species”,and their antipathy to its application to Homosapiens (or at least to the neocortex) seems to stemfrom their belief that selfish gene theory implies thatall human behaviour is genetically constrained to beselfish. They tell us that “perhaps all too many individualsutilize those capacities largely for their ownselfish ends, but it is important to emphasize that thenature of the higher regulatory systems in humansdoes permit many alternative courses of action”.This misconception appears to underpin their endorsementof group selection. I wont discuss thissimple misinterpretation here as it has been dealtwith many times in the literature, not least by DAWK-INS (1981, 1989). The improbability (though not impossibility)of group selection as an explanation forevolved traits is a standard text book matter (MA-JERUS/AMOS/HURST 1996, pp110–115), and JohnMAYNARD SMITH (1998) has described much of thecontroversy as “largely semantic”.Unlike sociobiologists, evolutionary psychologistsare skeptical of the extent to which evolved behaviouraltendencies cause human behaviour to assumethe form that maximizes inclusive fitness(SYMONS 1989). BARKOW (1990) has described threeways in which sociocultural traits can affect the fitnessof their participants: “a) they can enhance fitnessbecause the cultural trait is a direct reflection ofan evolved psychological mechanism; b) they canlower fitness or be neutral for it, because culturalprocesses are semi-independent of biological evolution;or c) they can enhance fitness epiphenomenally,that is, in a manner having little or no connectionwith past genetic selection”. Because ourcontemporary environments (and hence our developmentalsystems) incorporate such novel aspects asmass media, drugs, processed foods, pollutants,large group sizes, reduced interaction with kin, andmany other phenomena not typical of the hunter–gatherer environment, it is highly probable that currentbehaviour is unlikely to be a reliable guide toancestral behaviour, and evolutionary psychology’scommitment to the study of evolved psychologicalmechanisms should take precedence over (thoughnot entirely replace) the study of current fitness.Neuro-Foundational IssuesFor the PANKSEPPs the neurofoundational issues weneed to take into consideration are the existence ofevolutionary layers in the brain; the preservation ofancient motivational mechanisms; the evidence for‘mind’ in the variability of behaviour related to intrinsicrepresentations rather than external stimuli;the supervenience of mind on the brain; the irreducibilityof mind to the functions of the brain, and anEvolution and Cognition ❘ 41 ❘ 2001, Vol. 7, No. 1

Ian Pitchfordappreciation that the foundations of human naturewere ‘solidified’ way before the Pleistocene era. Iwould like to suggest a broader perspective based onempirical and theoretical considerations.In 1935 William KING GREGORY suggested thatreplication of body parts due to genetic mutationhad been a major mechanism in evolution. In 1971this idea was extended by John ALLMAN’s and JonKAAS’ proposal that the evolution of the brain wascharacterised by the replication of cortical areas (AL-LMAN 1999, p40). RILLING’s and INSEL’s comparativeMRI study of the primate neocortex confirms thefinding that the human brain is slightly over threetimes larger than would be expected for a primate ofthe same body size (1999b). However, the data indicatea striking discrepancy between human andpongid brains in the extensive gyrification in theprefrontal cortex of the former, an important findinggiven the role of this region in complex problem-solving(KOECHLIN et al. 1999), and social intelligence(ROWE et al. 2001; SHALLICE 2001; STUSS/GALLUP/ALEXANDER 2001). As RILLING/INSEL (1999b)conclude this departure from allometric trends“suggests selection for increased gyrification in theprefrontal cortex throughout hominid evolution”.The other area noted for significantly more gyrificationthan expected is the seventh coronal slice, aregion incorporating WERNICKE’s area, long implicatedin the production and comprehension of language(1999b). RILLING and INSEL also note that theincrease in human neocortical gray matter is notproportional with the increase in the volume of therest of the brain and that, although the increase inwhite matter outpaces that in grey, this increasefalls well short of that necessary to retain the samelevel of interconnectivity between neurons. Anotherscan of 11 primate species concentrating onthe corpus callosum and anterior commissure demonstratesthat the increase in primate brain size hasresulted in increasingly independent hemispheres(RILLING/INSEL 1999a). Through their work on theinsular cortex of bottlenose dolphins MANGER et al.(1998) have found that although brain sizes varydramatically across animal species, the range ofmodule size is restricted, though the number of corticalareas across species is highly variable (KAAS1993; KAAS/REINER 1999). For additional evidenceon mosaic brain evolution see BARTON/HARVEY(2000) and DE WINTER/OXNARD (2001). These findingsconfirm that brain evolution is characterisedby the independent evolution of brain structureswith anatomical and functional links. Indeed, oneof the most distinctive features of the neocortex isits modular organization (JONES 2000; MOUNTCASTLE1997; ROCKLAND 1998).The existence of a neuronal type found only inthe brains of pongids and hominids is also likely tobe of importance. Using samples of the anterior cingulatecortex (BRODMANN’s area 24) of 28 primatespecies NIMCHINSKY et al. (1999) found a spindleshapedcell in layer Vb specific to humans and greatapes. The anterior cingulate is known to be involvedin response selection (AWH/GEHRING 1999; TURKEN/SWICK 1999), and performance monitoring (CARTERet al. 1998), but also appears to have a number ofdiscrete, functional regions subserving importantaspects of cognition, emotion, and notably vocalization(BUSH/LUU/POSNER 2000). NIMCHINSKY et al.(1999) note that “the emergence of this unique neuronaltype in a neocortical area involved in vocalizationin primates coincides with the evolution as adefinable anatomic structure of the planum temporale,a region that is important for language comprehension.In view of the language comprehensionabilities of great apes, it is therefore possible that severalcortical structures involved in the production ofspecific vocalizations and in communicative skillssustained simultaneous, considerable, adaptivemodifications during brain evolution in hominoids”.In considering neuroevolutionary matters weshould always keep the issue of sexual dimorphismin mind. There are two types of human brain, maleand female, and it is reasonable to expect that thesehave been subject to different selection pressures.For example, women have a higher proportion ofgrey matter to cranial volume, whereas men have ahigher proportion of white matter and cerebrospinalfluid to cranial volume. Women also have a relativelylarger corpus callosum than men. GUR et al.(1999) found that of the top ten performers in a spatialtask, nine were men, and seven of these men hadgreater white matter volumes than any of thewomen in the study. Our large brains may not simplyprovide an excess of plastic neurons capable ofsubserving any function, but may be a solution tothe problem of retaining adequate functioning overa prolonged life span (HUMPHREY 1999), somethingthat could be of particular importance to caregivers.ALLMAN has found that there is a significant correlationbetween brain weight and maximum life-spanin haplorhine primates (ALLMAN/MCLAUGHLIN/HAKEEM 1993), and that the maximum human lifespanis close to what would be expected for a primateof our relative brain size (ALLMAN 1999, p172). ALL-MAN et al. (1998) have also discovered in a variety ofEvolution and Cognition ❘ 42 ❘ 2001, Vol. 7, No. 1

No Evolution. No Cognitionspecies that caregivers live longer, whether male orfemale, and “that there is no difference in survivalbetween the sexes in species in which both parentsparticipate about equally in infant care”. The factthat human females are the primary caregivers, andthat human grandmothers are able to enhance theirfitness post-menopausally by assisting the reproductivesuccess of their daughters may also help to explainthe structural and functional differences betweenthe brains of men and women (HAWKES et al.1998; O’CONNELL et al. 1999).All adaptations have costs and benefits, and it iscertain that psychological mechanisms are not costfreebecause the rate of DNA damage in mammaliancells is extremely high, amounting to tens of thousandsof DNA damages per day. This implies an enormousmetabolic cost in maintenance and repair (DU-KAS 1999). Also, as many of the processes within thebrain are mediated by the same neurochemicals,functional systems must have the capacity to ensurethat the correct information is elicited as required.One benefit derived from the piecemeal addition ofoverlapping systems is explained by DUKAS in hisanalysis of the costs of memory: redundancy helpsto reduce the amount of error and noise in the system,and therefore “probably plays a key role in ensuringa high level of accuracy” (DUKAS 1999). Thecost of redundancy is in terms of increased brainmass and energetic expenditure on maintenance, repairand replication. As HENINGER (1999, pp93–94)explains: “One of the main features of the nervoussystem is the mutually dependent, diffuse, and oftenredundant biologic processes that subserve functions.In contrast to the relative specificity of sensoryand motor systems, the systems subserving sleep–wakefulness, arousal-motivation, emotional reactivity,memory, and higher order behavioural functionsare more widely distributed anatomically. Thesystems demonstrate extremely complex nonlinearresponse characteristics so that there is not a simpleone–to–one correspondence between measures ofneuronal function and the behaviours studied. Inaddition, there is a great deal of plasticity so thatremaining systems can compensate for deficits”.The PANKSEPPs are surely right when they claimthat ancient phylogenetic mechanisms are conserved,and that consequently homologies can teachus much about the construction of the humanmind, but I can think of no reason for believing thatthe neocortex has been moulded by processes substantiallydifferent to those responsible for the subcorticalstructures involved in the basic emotions.The PANKSEPPs vision of a brain consisting of horizontallayers, derived from MACLEAN’s model of thetriune brain, is deeply misleading. Instead of horizontallayers our model should postulate integratedvertical modules built from co-evolving structuresdistributed throughout the brain, the different subcomponentsof which may be traced to differentevolutionary eras, and explained by the influence ofdifferent selection pressures.Finally, we should be aware that the distinctivecultural traits of human beings appear to haveemerged (or grown in significance) during a periodin which brain sizes have decreased. It appears thatsince the Late Pleistocene (around 30,000 years ago)human brain size has decreased by approximatelyten per cent with this decrease being paralleled by adecrease in body size. HENNEBERG (1998) notes “itmay be concluded that the gross anatomy of thehominid brain is not related to its functional capabilities.The large human brain:body size ratio maybe a result of the structural reduction of the size ofthe gastrointestinal tract and, consequently, its musculoskeletalsupports. It is related to richer, meatbaseddiets and extra-oral food processing ratherthan the exceptional increase in the size of the cerebrum.The exceptional mental abilities of humansmay be a result of functional rather than anatomicalevolution”.All of the foregoing theoretical and empirical considerationsindicate that the sudden emergence of ahighly plastic general-purpose neocortex responsiblefor multimodal functioning is a distinctly implausibleevolutionary event, and that evolutionarypsychology’s commitment to modularity is sound.ConclusionEvolutionary psychologists, including the PANK-SEPPs’ exemplars COSMIDES and TOOBY, advocate theassimilation of findings from evolutionary biologyand its related disciplines (TOOBY/COSMIDES 1998),including neurobiology (TOOBY/COSMIDES 1995), ina vertically integrated hierarchy of mutually consistentscientific disciplines that does not require or call“for reductionism nor for the conquest and assimilationof one field by another” (COSMIDES/TOOBY/BARKOW 1992, p12). Evolutionary psychology’sadoption of the concept of modularity from neurologycreates a natural affinity with the neurosciences,whilst the emphasis on psychological mechanismsand the mismatch between ancestral and contemporaryenvironments provides a bridge to psychologyand the social sciences. This richly interdisciplinaryfield has already generated an abundance of researchEvolution and Cognition ❘ 43 ❘ 2001, Vol. 7, No. 1

Ian Pitchfordin developmental psychology, neurology, psychiatry,and primatology, particularly on theory ofmind, which has produced compelling new theoriesof autism and schizophrenia. This multi- and interdisciplinaryresearch exhibits the consilience, orconfluence of different lines of evidence, so typicalof robust scientific work (seeCORCORAN 2000 and BARON-COHEN 2000 for a summary). Itcompares extremely favourablywith that based on theneurochemical individuationof traits and disorders admiredby the PANKSEPPs, which hasAuthor’s addressIan Pitchford, Centre for PsychotherapeuticStudies, University of Sheffield, 16 ClaremontCrescent, Sheffield, S10 2TA, UK.Email ian.pitchford@scientist.comproduced so many unsatisfactory theories and treatments(VALENSTEIN 1998).Neuroevolutionary psychobiology’s reassignmentof the tabula rasa to the neocortex revivifiesCARTESIAN dualism and represents a desperate strategylikely to be extremely deleterious to scientificinvestigation. 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Scott AtranThe Case for Modularity: Sin or Salvation?IntroductionThe central thesis ofPANKSEPP and PANKSEPP(2001, henceforth, P&P)is that there are no speciallyevolved, higher-orderdomain-specificcognitive mechanisms tospeak of. Apart from thebasic DARWINIAN emotions(fear, surprise, anger,sadness, joy, disgust),so-called cognitive ‘modules’are more likely “theproduct of dubious humanambition ratherthan sound scientific reasoning”.What evolutionarypsychologists (as wellas many cognitive anddevelopmental psychologists)take to be modularstructures are actually theepigenetic products ofthe ancestral emotionalfunctions of the brain(rooted in what was oncecalled “the limbic system,”MACLEAN 1990)and more recent “general-purposebrain mechanisms”.In more than a dozen places, P&P repeat asmantra that empirical evidence indicates the modernhuman mind was created through the dualfunctioning of subcortical mammalian emotionsand a neocortical general-purpose computationaldevice. Furthermore, this highly flexible, all-purposeintelligence probably emerged “more rapidlyvia group selection than by individual selection”.There is much to commend the cautionary talethat P&P tell regarding possible excesses of sociobiologicalspeculation, including what are arguably‘just-so’ stories about modularized adaptations forAbstractThe case for evolved cognitive modules rests on severalconverging lines of evidence: Functional design(analogy), ethology (homology), universality, precocityof acquisition, independence from perceptual experience(poverty of stimulus), selective pathology(cerebral impairment), resistance to inhibition (hyperactivity),ease of cultural transmission. No factor maybe necessary but evidence for all or some is compelling,if not conclusive, for domains like folkmechanics,folkbiology, folkpsychology. By constrast, noempirical evidence supports PANKSEPP and PANKSEPP’scentral thesis: that what evolutionary psychologists(and many cognitive and developmental psychologists)consider modular structures are actually epigeneticproducts of subcortical mammalian emotionsand neocortical general-purpose computations. Arguably,no significant empirical discovery about languageor other higher-order cognitive structures yetowes to inquiries about evolutionary origins and functions.Nevertheless, adopting evolutionary psychology’srequirement that candidate exaptations andspandrels be described, as far as possible, in connectionwith evolved adaptations, opens new avenues forexploring and testing modular designs.Key wordsModularity, domain-specificity, language, folkpsychology,folkbiology.Evolution and Cognition ❘ 46 ❘ 2001, Vol. 7, No. 1rape (THORNHILL/PALMER2000), homicide (BUSS1999) and emotional disorderssuch as depression(NESSE/LLOYD 1992).P&P’s position on the derivativeand epigeneticcharacter of ‘secondary’or ‘social’ emotions (guilt,love, empathy, etc.), isalso defensible (e.g., DAM-ASIO 1994), although seriouslydebatable (cf.LEDOUX 1996; GRIFFITHS1997). Finally, P&P’s takeon the current, overlyfundedfad for neuroimaging(driven more bytechnological innovationand the industry it supportsthan by any theoreticalinsight) is credible.There can no more be atheory directly derivedfrom observations of neuroimagingthan there canbe a theory derived directlyfrom observationsof meter readings; it isonly in the service ofsome prior abstract theorythat such observationscan make sense. Nevertheless, there is no empiricalevidence whatever to indicate that P&P’scentral thesis is true, and much to suggest it is false.Evolutionary Psychology:No Explanatory Value forHigher-Order Cognition (Yet)To-date, evolutionary psychology has not predictedor discovered any significant or surprising aspect oflanguage or higher-order cognitive devices of thesort I describe below. But this does not mean that all

The Case for Modularity: Sin or Salvation?evolutionary psychology accounts are ‘just-so’ stories.The best account of language as an evolutionaryadaptation centers on the claim that “languageshows signs of design for the communication ofpropositional structures over a serial channel”(PINKER/BLOOM 1990). PINKER and BLOOM describehow specific syntactic structures conform to thislanguage-specific design in ways that provide functionaladvantage: for example, through certainstructure-dependent rules (phrase structure) andprinciples of embedding (recursion) that allow theformulation and expression of infinitely many discreteideas by finite and few means. This enables themultiple thoughts of multiple individuals to becombined, tested in imagination and consequentlyincluded or excluded from having a role in some futureaction. A population whose individuals couldcontemplate alternative scenarios in any sequenceand at any rate, benefit from the cognitive travailsof others, and let conjectures die instead of themselves,would surely have had an evolutionary advantageover a population that couldn’t.PINKER and BLOOM offer a well-reasoned and oftennonobvious analysis of evolutionary tradeoffs involvedin opting for linear communication throughan auditory medium: visual displays better communicatehighly complex topological relationships (“apicture is worth a thousand words”), gestures betterconvey emotion, and language is inefficient transmittinginformation about taste or smell. For all itsinsight and plausibility, though, PINKER and BLOOM’saccount remains wholly backward-looking: nostructural discovery, novel prediction or theoreticalbreakthrough ensues within the CHOMSKYEAN frameworkthat the authors adopt.According to Jerry FODOR (1998), whose reasoningis close to CHOMSKY’s (1988, p170), it is not likely thatnatural selection gradually produced an adaptivemutation for a language instinct. Rather, selectionpressures unrelated to language simply made humanbrains a little larger and more complex. Unlike gradualand incremental adaptations, a little added complexitycan go a long way fast to produce multiplenovel structures:“Make the giraffe’s neck just a little bit longer andyou correspondingly increase, just by a little, the animal’scapacity to reach the fruit at the top of thetree… But make an ape’s brain just a little bigger (ordenser, or more folded, or, who knows, grayer) andit’s anybody’s guess what happens to the creature’sbehavioural repertoire”. (FODOR 1998).Much ink has spilled and acrimony vetted in thedebate between evolutionary psychologists and thescientists who criticize them. In the case of language,however, no empirical issue has yet to turn on thedebate. Despite very different evolutionary stories,CHOMSKY, FODOR, PINKER and BLOOM fundamentallyagree on the specific computational structures thatcharacterize language, on its innateness, on itshighly specialized mode of operation (modularity)and on the fact that all of this is uniquely the productof evolution, whether adaptation or by-product.The Language ModuleFrom a purely logical standpoint, if a mind is able totake fragmentary instances of experience (relative tothe richness and complexity of the whole data set)and spontaneously predict (project, generalize) theextension of those scattered cases to an indefinitelylarge class of intricately related cases, then the inferentialstructure responsible for this prediction cannotpossibly derive from the experience. As Humenoted, the structure must be prior to experience, justlike the cranes and architects used for constructingbuildings must exist prior to initial construction: abuilding does not just build itself. There is no otherpossibility. What, then, are the physical possibilitiesfor a mental structure to be “prior to experience”?One could hold with SOCRATES or BUDDHA thatpeople are born with a past-life ‘memory’ for thestructure of experience they will encounter later inlife; or, one could hold with Thomas AQUINAS or ImmanuelKANT that God put structure in people’sminds. If, however, one accepts humans as biologicalcreatures whose species attributes emergedthrough the same evolutionary processes that governthe emergence of all other species, then thereappears to be no alternative to a priori mental structuresbeing evolved biological structures (whether asadaptations or by-products of adaptations). To sayan evolved biological structure is ‘innate’ is not tosay that every important aspect of its phenotypicexpression is ‘genetically determined’. Biologicallypoised structures channel development, but do notdetermine it—like mountains that channel scatteredrain into the same mountain–valley river basin(WADDINGTON 1959).P&P might grant all this, but argue that an allpurpose,domain-general computational device sufficesto give the mind/brain inferential power beyondthe information given. As an empirical claim,this entails that one should be able to deduce thestructural principles and parameters governing anyspecific domain from general purpose-structures actingunder the constraints of experience in the rele-Evolution and Cognition ❘ 47 ❘ 2001, Vol. 7, No. 1

Scott Atranvant domain. For example, P&P offer that language“may be closely linked to anterior cingulate and adjacentfrontal lobe tissues which appear to mediatecertain types of pain, feelings of separation distressand thereby social sensitivities” (cf. MACLEAN 1990).But there is nothing in what we know of generalintelligence (e.g., conditioning, association, etc.),pain, distress, social sensitivity, or the structures ofanterior cingulate and adjacent frontal lobe tissues,to even remotely hint at the highly structured, andstructurally peculiar, principles and parameters ofhuman language. This includes: anaphora (the structuralconstraints on how even widely separated partsof an expression co-refer), quantification (the structuralconstraints that ‘who,’ ‘many,’ few,’ etc. imposeon the syntactic behavior of subjects and objects),negation (it is literally impossible to learn negationthrough perceived experience or ‘interaction,’ as onecannot perceive or interact with something thatisn’t), word order (all languages have a specified linearorder linking arguments to predicates), and so forth(CHOMSKY 1982).Briefly, the central tenet of CHOMSKY’s (2000) approachis that there is a language system, LS, of thehuman brain. LS reflexively discriminates and categorizesparts of the flux of human experience as ‘language,’and develops complex abilities to infer andinterpret this highly structured, and structurally peculiar,type of human production. In a general sense,there is nothing intrinsically different about LS—interms of innateness, evolution or universality—thanthe visual system (VS), immune system (IS), respiratorysystem (RS), or any other complex biologicalsystem. Learning syntactic structure through ‘socialinteraction’ is no more plausible an alternative thanlearning by ‘osmosis’. If a child is initially deprivedof linguistic input, the child may never come to acquire,or ‘know,’ much in the way of language, justas a kitten initially deprived of normal exposure tothe visual world may never come to ‘know’ much byway of object recognition (HUBEL 1988). Still, emotionand social interaction no more ‘create’ languagein the child’s mind than lighting and object patterning‘create’ vision in the kitten.Interacting ModulesLS is no more (or less) ‘autonomous’ from the surroundingsocial environment, or from other mentalsystems, than VS is detachable from surroundinglight and object patterning or from other physicalsystems (including, in humans, linguistic and othercognitive systems of meaning, MARR 1982). LS andVS do not exist, and cannot develop, in isolation,but only as subsystems of even more intricate structures.Thus, claims about the biological ‘autonomy’or ‘modularity’ of LS or VS refer only to a specifiablelevel of systemic functioning within a system hierarchy.A difficult empirical issue concerns the extentto which other cognitive ‘performance’ systems arethemselves specifically adapted for language. Thereis little doubt that the sound system is highly structuredfor access (psycholinguistic experiments withneonates and even fetuses, dichotic listening experiments,comparative phonology, and so forth arecompelling). The interface between syntactic andsemantic systems is much more obscure. Speculationis rampant, debate is furious, and critical experimentsare few, as might be expected from arelatively new and dynamically changing science.But progress is being made, slowly and laboriously,as intense research by teams of well-trained investigatorscontinues.Cognitive and developmental psychologists andpsycholinguistics have identified a number of structuralprinciples in human cognitive systems that relateto the interface between LS and these other systems.Among the principles discovered is “thewhole–object constraint” (CAREY 1985). Children,whatever their culture or language, apparently assumeas a default that nouns in general apply towhole objects (e.g., a rabbit) and not to parts of theobject (e.g., a piece of a leg, a patch of fur, or spatiallyseparated patches of fur and pieces of leg), or theobject–and–its–environment. Children actuallyhave to learn that this is not always the case (just aschildren have to learn that the shadow of an objectis not itself an object, SPELKE et al. 1995). This apparently‘innate bias’ helps to solve the problem of radicalindeterminacy of translation. Consider an anthropologistwho visits an exotic tribe for the firsttime and sees a member of that tribe pointing tosomething that the anthropologist immediatelyidentifies as a running deer. The anthropologist isfairly safe in assuming that the informant alsothought of pointing to a running deer (and not amoving piece of deer, a shifting pattern of deer furand grass, etc.). This is so even if the informant alsobelieves that the deer is some other person’s ancestor.If this were not the case—if radical indeterminacywere an omnipresent possibility—then anthropologywould be impossible and ethnography couldbe only fiction.Before learning to talk, children first learn to followgaze, engage in joint attention, contingently interactwith others to achieve goals, and so forth.Evolution and Cognition ❘ 48 ❘ 2001, Vol. 7, No. 1

The Case for Modularity: Sin or Salvation?Those who argue that language is acquired throughsocial interaction might be tempted to claim thatthese facts undermine the modular model of language(SHANKER 2001). Yet, most of the work in thisarea—called the child’s ‘theory of mind’ or ‘folkpsychology’–focuseson many of the same sorts of questionsand approaches that generative grammariansdo: How is the child able to reliably infer such richmental structures about other minds from a few gesturesand without mastery of language? And how isit that children also infer that people’s mental structures(intentions) can cause others to act a distance(without any physical contact)? The emerging consensusin the field is that children are biologicallyendowed with a ‘theory of mind’ (ToM) that maturesin predictable ways over the first three years of life(LESLIE 1994; BARON-COHEN 1995; SPELKE et al 1995).Some features of developing ToM interface with maturingfeatures of developing LS in systematic waysthat are only now being experimentally studied. Thisis one of the most exciting and productive researchareas in cognitive and developmental psychology (tojudge from journal publications and grant funding),and also one of the most thoroughly ‘modular’.Interactions between modular cognitive systemsare complex and difficult to tease apart, perhapsmore so than interactions among various bodily systems.For example, belief in supernatural agents,which characterizes all religions in all societies, involvesa host of modular expectations and interfacing:folkmechanics, folkpsychology, folkbiology,primary and secondary emotions, predator–preyschema, and so forth (BOYER 1994; BARRETT/NYHOFF2001; ATRAN in press). Nevertheless, predictive theoriesare being proposed and empirically tested.Are Modules Adaptations orEvolutionary By-Products of Adaptations?Leading evolutionary biologists, such as StephenGOULD and Richard LEWONTIN (1979) describe theproducts of higher-order human cognition as evolutionary‘by-products’ or ‘spandrels’. This is a stanceP&P adopt. A spandrel is an architectural term forthe structural form or space that arises as a necessaryconcomitant to another decision in design, and isnot designed to have any direct utility in itself. Forexample, the space beneath a flight of stairs is a byproductof constructing an inclined stairway ratherthan a vertical ladder. The fact that people mightsubsequently use this ‘leftover’ space for storagedoes not entail that the space was designed to be astorage space.For GOULD and LEWONTIN, higher-order humancognitive structures originated as functionless spandrelsthat have been subsequently modified undercultural selection rather than natural selection. Biologicallyfunctionless, or nearly functionless, spandrelssupposedly include: religion, writing, art, science,commerce, war and play. These evolutionaryby-products are cultural ‘mountains’ to the biologically‘adaptive molehill’ (GOULD 1991, pp58–59;WILLIAMS 1992, pp77–79). On this account, evolutionarypsychology would have little to reveal aboutthe emergence and structure of such culturally-elaboratedspandrels: “The number and complexity ofthese spandrels should increase with the intricacy ofthe organism under consideration. In some regionwithin a spectrum of rising complexity, the numberand importance of useable and significant spandrelswill probably exceed the evolutionary import of theprimary adaptation” (GOULD 1997, pp10754–10755;cf. FODOR 1998).GOULD’s account of the emergence of distinctlyhuman cognitions as spandrels of a big brain ishardly convincing. The very notion of a ‘big’ or‘large’ or ‘complex’ brain is too vague to empiricallyconstrain the evolutionary story about how it mighthave evolved. The big brain is taken as an adaptationfrom which all cognitive spandrels arise. But an adaptationto what? A design for what? What are itsevolutionary-relevant computational structures?Big brain stories tend to be even broader in scope andhandwaving than language–evolution stories. Humanbrains supposedly broke away from ape brainsunder selection pressures that run the gamut fromrunaway social competition (ALEXANDER 1989), togut reduction (AIELLO/WHEELER 1995), to huntinglarge game (HILL 1999), to niche construction (LA-LAND et al. 2000), to runaway sexual selection(MILLER 2000), and so on. Alternatively, the big brainprimarily evolved as a conduit for culture (HARRIS1975), as a vehicle for language (JERISON 1976), or asa host for the independently evolving ideas, or‘memes,’ that compete to colonize it (DENNETT 1995;BLACKMORE 1999). Finally, the big brain may haveevolved under any number ‘positive feedback’ pathwaysinvolving some or all of the factors mentioned.But even if true, and even if we knew the reasons(selection pressures responsible for) why, we are unlikelyto learn anything of particular interest from allthis about how the mind works. The notion of a bigbrain is as uninformative about cognitive structuresand functions cognition as the notion of big body isabout bodily structures and functions. Never mindexaptations, such as chins, or spandrels, such asEvolution and Cognition ❘ 49 ❘ 2001, Vol. 7, No. 1

Scott Atranpalm lines. From the fact of a bigger body, whatcould one possibly deduce about hearts, livers, kidneys,hands, faces, placentas and so on? And nevermind exaptations, such as language, or spandrels,such as religion. From the fact of a bigger (or denser,or more folded, or grayer) brain, what could one possiblydeduce about perception, emotion, categorization,inference or any of the other capabilities humansshare with apes but in more vastly elaboratedform? Probably nothing at all.It may well be true that little insight is to be gainedinto higher-order human cognitive structures byconsidering possible evolutionary origins and functions.Adopting the hypothesis of GOULD andLEWONTIN or P&P, which assumes this truth, is practicallyguaranteed to block insight, whether or not itis true. By contrast, adopting evolutionary psychology’srequirement that candidates for exaptationsand spandrels be described, as far as possible, withreference to evolved adaptations, then it might bepossible to find out if the hypothesis is true or not.If it is true, then evolutionary psychology wouldhave provided the empirical evidence that shows itto be a significant and surprising scientific insight,and not one that depends entirely on intuition, analogy,eloquence or wishful thinking. If it is not true,then evolutionary psychology will have helped todiscover something new about human nature.Three Examples of Modularity: NaiveMechanics, Theory of Mind, FolkbiologyEver since CHOMSKY jump-started the ‘cognitive revolution,’successors to the behaviorists whobelieved in an all-powerful general thinking devicehave tried to reconcile CHOMSKY’s insights with faithin flexible intelligence by reluctantly granting somespecificity to language, and language alone. But cognitivepsychology today concentrates more on discoveryand exploration of domain-specificmechanisms than on general-purpose computation.Each such device has a particular ‘content-bias’ inthat it targets some particular domain of stimuli inthe world (‘set of inputs’): for example, the edgesand trajectories of rigid three-dimensional bodiesthat move by physical contact between them(mechanics), the contingent motion a self-propelledactors that can coordinate interactions withouthaving physical contact (agency), or thebehaviors and appearances of nonhuman livingcreatures (species relations). The particular inferentialstructure of each domain-specific processor thentakes the isolated exemplars (or relatively poor samples)of the stimulus-set actually encountered in aperson’s life, and spontaneously projects these relativelyfragmentary instances onto richly-structuredcategories (‘classes of output’) of general relevanceto our species: for example, the objects and kinds offolkphysics (naive mechanics), folkpsychology(ToM) and folkbiology. Much work on domainspecificityhas developed, and now develops, independentlyof sociobiology or evolutionary psychology(ATRAN 1989; HIRSCHFELD 1996; KEIL 1989; LESLIE1994; CAREY/SPELKE 1994; SPERBER 1985).Within the emerging paradigm of cognitive domain-specificity,there is much speculation and controversy—again,as might be expected in any newlyemerging science. For example, there are competingaccounts of how human beings acquire basic knowledgeof the everyday biological world, including thecategorical limits of the biological domain and thecausal nature of its fundamental constituents. Oneinfluential view of conceptual development in folkbiologyhas been articulated by Susan CAREY and hercollaborators (CAREY 1985; CAREY/SPELKE 1994;JOHNSON/CAREY 1998). On this view, young children’sunderstanding of living things is embedded in afolkpsychological, rather than folkbiological, explanatoryframework. Only by age 7 do children begin toelaborate a specifically biological framework of theliving world, and only by age 10 does an autonomoustheory of biological causality emerge that is not basedon children’s understanding of how humans thinkand behave. A competing view is that folkbiology andfolkpsychology emerge early in childhood as largelyindependent domains of cognition that are clearlyevident by ages 4 or 5, and which may be innatelydifferentiated (ATRAN 1987; KEIL 1989; GELMAN/WELL-MAN 1991; HATANO/INAGAKI 1999).To address this issue, my colleagues and I carriedout a series of cross-cultural experiments (LÓPEZ et al1997; MEDIN et al. 1997; ATRAN et al. 2001). One setof experiments shows that by the age of 4–5 years(the earliest age tested in this regard) urban Americanand Yukatek Maya children employ a concept ofinnate species potential, or underlying essence, as aninferential framework for understanding the affiliationof an organism to a biological species, and forprojecting known and unknown biological propertiesto organisms in the face of uncertainty. Anotherset of experiments shows that the youngest Mayachildren do not have an anthropocentric understandingof the biological world. Children do notinitially need to reason about nonhuman livingkinds by analogy to human kinds. The fact thatAmerican children show anthropocentric bias ap-Evolution and Cognition ❘ 50 ❘ 2001, Vol. 7, No. 1

The Case for Modularity: Sin or Salvation?pears to owe more to a difference in cultural exposureto nonhuman biological kinds than to a basiccausal understanding of folkbiology per se. Together,the first two sets of experiments suggest that folkpsychologycan’t be the initial source of folkbiology.They also indicate that to master biological science,people must learn to inhibit activation of universaldispositions to view species essentialistically and tosee humans as inherently different from other animals(ATRAN 1990, 1998).A third set of results show that the same taxonomicrank is cognitively preferred for biological inductionin two diverse populations: people raised inthe Midwestern USA and Itza’ Maya of the LowlandMesoamerican rainforest (ATRAN et al. 1997; COLEYet al. 1997). This is the generic species—the level ofoak and robin. These findings cannot be explained bydomain-general models of similarity because suchmodels cannot account for why both cultures preferspecies-like groups in making inferences about thebiological world, although Americans have relativelylittle actual knowledge or experience at thislevel. In fact, general relations of perceptual similarityand expectations derived from experience producea ‘basic level’ of recognition and recall for manyAmericans that corresponds to the superordinatelife-form level of folkbiological taxonomy—the levelof tree and bird (ROSCH et al. 1976). Still, Americansprefer generic species for making inductions aboutthe distribution of biological properties among organisms,and for predicting the nature of the biologicalworld in the face of uncertainty. This supportsthe idea of the generic-species level as a partitioningof the ontological domains of plant and animal intomutually exclusive essences that are assumed (butnot necessarily known) to have unique underlyingcausal natures. The implication from these experimentsis that folkbiology may well represent an evolutionarydesign: universal taxonomic structures,centered on essence-based generic species, are arguablyroutine products of our ‘habits of mind’, whichmay be in part naturally selected to grasp relevantand recurrent ‘habits of the world’.Evidence for ModularityThe evolutionary argument for a naturally-selectedcognitive disposition, such as folkbiology, involvesconverging evidence from a number of venues:Functional design (analogy), ethology (homology),universality, precocity of acquisition, independencefrom perceptual experience (poverty of stimulus),selective pathology (cerebral impairment), resistanceto inhibition (hyperactivity), and culturaltransmission. None of these criteria may be necessary,but the presence of all or some is compelling, ifnot conclusive. Group selection is not a factor.1. Functional Design. All organisms must functionto procure energy to survive, and they also must procure(genetic) information for recombination andreproduction (ELDREDGE 1986). The first requirementis primarily satisfied by other species, and an indiscriminateuse of any individual of the other species(e.g., energy-wise, it does not generally matter whichchicken or apple you eat). The second requirementis usually only satisfied by genetic informationunique to individual conspecifics (e.g., genetically,it matters who is chosen as a mate and who is consideredkin). On the one hand, humans recognizeother humans by individuating them with the aid ofspecies-specific triggering algorithms that ‘automatically’coordinate perceptual cues (e.g., facial recognitionschemata, gaze) with conceptual assumptions(e.g., intentions) (BARON-COHEN 1995). On the otherhand, people do not spontaneously individuate themembers of other species in this way, but as examplarsof the (generic) species that identifies them ascausally belonging to only one essential kind.Natural selection basically accounts only for theappearance of complexly well-structured biologicaltraits that are designed to perform important functionaltasks of adaptive benefit to organisms. In general,naturally selected adaptations are structuresfunctionally “perfected for any given habit” (DAR-WIN 1883, p140), having “very much the appearanceof design by an intelligent designer… on which thewellbeing and very existence of the organism depends”(WALLACE 1901, p138). Plausibly, the universalappreciation of generic species as the causal foundationfor the taxonomic arrangement ofbiodiversity, and for taxonomic inference about thedistribution of causally-related properties that underliebiodiversity, is one such functional evolutionaryadaptation. But a good story is not enough. 12. Ethology. One hallmark of adaptation is a phylogenetichistory that extends beyond the species inwhich the adaptation is perfected: for example, ducklingscrouching in the presence of hawks, but notother kinds of birds, suggests dedicated mechanismsfor something like species recognition. Some nonhumanspecies can clearly distinguish several differentanimal or plant species (CERELLA 1979; LORENZ 1966;HERRNSTEIN 1984). Vervet monkeys even have distinctalarm calls for different predator species orEvolution and Cognition ❘ 51 ❘ 2001, Vol. 7, No. 1

Scott Atrangroups of species: snake, leopard and cheetah, hawkeagle, and so forth (HAUSER 2000). Chimpanzees mayhave rudimentary hierarchical groupings of biologicalgroups within groups (BROWN/BOYSEN in press).To be sure, the world itself is neither chaos nor flux:species are often locally self-structuring entities thatare reproductively and ecologically isolated fromother species through natural selection. But there isno a priori reason for the mind to always focus oncategorizing and relating species qua species, unlessdoing so served some adaptive function (e.g., itmakes little difference which tiger could eat a personor which mango a person could eat). And the adaptivefunctions of organisms rarely, if ever, evolve or operatein nature as all-purpose mechanisms.3. Universality. Ever since the pioneering work ofBERLIN and his colleagues, evidence from ethnobiologyand experimental psychology has been accumulatingthat all human societies have similar folkbiologicalstructures (BERLIN et al. 1973; BERLIN 1992;ATRAN 1990, 1999). These striking cross-cultural similaritiessuggest that a small number of organizingprinciples universally define folkbiological systems.Basic aspects of folkbiological structure (e.g., taxonomicranking, primacy of generic-species) seem tovary little across cultures as a function of theories orbelief systems.4. Ease of Acquisition. Acquisition studies indicatea precocious emergence of essentialist folkbiologicalprinciples in early childhood that are not applied toother domains (GELMAN/WELLMAN 1991; KEIL 1994;HATANO/INAGAKI 1999; ATRAN et al. 2001).5. Independence from Perceptual Experience.Experiments on inferential processing show thatthat humans do not make biological inductions primarilyon the basis of perceptual experience or anygeneral similarity-based metric, but on the basis ofimperceptible causal expectations of a peculiar, essentialistnature (ATRAN et al. 1997; COLEY et al.1997).6. Pathology. Cerebral impairments (WILLIAMS syndrome,brain lesions caused by certain types of herpesvirus, etc.) suggest selective retention or loss offolkbiological taxonomies or of particular taxonomicranks. Neuropsychological studies have reporteda pathological performance in recognition atthe life-form and generic-species levels (e.g., recognizingan item as an animal but not as a bird orrobin), and dissociation at the life-form level (e.g.,not recognizing items as trees). Existing studies,however, do not say anything about the generic-speciesrank as the preferred level of representation forreasoning, perhaps because of methodology (linkedto averaging over items and failure to include sets ofgeneric species) (WARRINGTON/SHALLICE 1984; SAR-TORI/JOB 1988; JOB/SURIAN 1998).7. Inhibition and Hyperactivity. One characteristicof an evolved cognitive disposition is evident difficultyin inhibiting its operation (HAUSER 2000).Consider beliefs in biological essences. Such beliefsgreatly help people explore the world by proddingthem to look for regularities and to seek explanationsof variation in terms of underlying patterns.This strategy may help bring order to ordinary circumstances,including those relevant to human survival.But in other circumstances, such as wanting toknow what is correct or true for the cosmos at large,such intuitively ingrained concepts and beliefs mayhinder more than help. For example, the essentialistbias to understand variation in terms of deviance isundoubtedly a hindrance to evolutionary thinking.In some everyday matters, the tendency to essentializeor explain variation in terms of deviation fromsome essential ideal or norm (e.g., people as mentalor biological ‘deviants’) can be an effortlessly ‘natural’but wrong way to think.Because intuitive notions come to us so naturallythey may be difficult to unlearn and transcend. Evenstudents and philosophers of biology often find itdifficult to abandon commonsense notions of speciesas classes, essences or natural kinds in favor ofthe concept of species as a logical individual—a genealogicalbranch whose endpoints are somewhat arbitrarilydefined in the phyletic tree and whose statusdoes not differ in principle for that of other smaller(variety) and larger (genus) branches. Similarly, racism—theprojection of biological essences onto socialgroups—seems to be a cognitively facile and culturallyuniversal tendency (HIRSCHFELD 1996).Although science teaches that race is biologically incoherent,racial or ethnic essentialism is as notoriouslydifficult to suppress as it is easy to incite (GIL-WHITE 2001).8. Cultural Transmission. Human cultures favor arapid selection and stable distribution of those ideasthat: 1) readily help to solve relevant and recurrentenvironmental problems, 2) are easily memorizedand processed by the human brain, and 3) facilitatethe retention and understanding of ideas that aremore variable (e.g ., religion) or difficult to learnEvolution and Cognition ❘ 52 ❘ 2001, Vol. 7, No. 1

The Case for Modularity: Sin or Salvation?(e.g., science) but contingently useful or important.Folkbiological taxonomy readily aids humans everywherein orienting themselves and surviving in thenatural world. Its content tends to be stable withincultures (high interinformant agreement, substantialhistorical continuity) and its structure isomorphicacross cultures (BERLIN et al. 1973; LÓPEZ et al.1997). Folkbiological taxonomy also serves as a principledbasis for transmission and acquisition of morevariable and extended forms of cultural knowledge,such as certain forms of religious and scientific belief(ATRAN 1990, 1998).Consider, for example, the spontaneous emergenceof totemism—the correspondence of socialgroups with generic species—at different times andin different parts of the world. Why, as LÉVI-STRAUSS(1963) aptly noted, are totems so “good to think”?In part, totemism uses representations of genericspecies to represent groups of people; however, thispervasive metarepresentational inclination arguablyowes its recurrence to its ability to ride piggybackon folkbiological taxonomy. Generic speciesand groups of generic species are inherently wellstructured,attention-arresting, memorable andreadily transmissible across minds. As a result, theyreadily provide effective pegs on which to attachknowledge and behavior of less intrinsically welldeterminedsocial groups. In this way totemic groupscan also become memorable, attention-arrestingand transmissible across minds. These are the conditionsfor any idea to become culturally viable (seeSPERBER 1996 for a general view of culture along thelines of an ‘epidemiology of representations’). A significantfeature of totemism that enhances bothmemorability and its capacity to grab attention isthat it violates the general behavior of biological species:members of a totem, unlike members of a genericspecies, generally do not interbreed, but onlymate with members of other totems in order to createa system of social exchange. Notice that this violationof core knowledge is far from arbitrary. In fact,it is such a pointed violation of human beings’ intuitiveontology that it readily mobilizes most of theassumptions people ordinarilymake about biology in order tobetter help build societies aroundthe world (ATRAN/SPERBER 1991).In brief, modularized structures—suchas those which producefolkmechanical, folkpsychologicaland folkbiologicalconcepts—are special players incultural evolution. Their nativestability derivatively attaches to more variable anddifficult–to–learn representational forms, thus enhancingthe latter’s prospects for regularity and recurrencein transmission within and across cultures.This includes knowledge that cumulativelyenriches (e.g., to produce folk expertise), overrides(e.g., to produce religious belief) or otherwise transcends(e.g., to produce science) the everyday ontologyprescribed by our evolved cognitive modules.Conclusion: Neither Sin nor Salvation,but a Promissory NoteDespite the initial independence of work in domainspecificity,there is now increasing convergence inthe ways cognitive anthropologists and psychologists,and evolutionary biologists and psychologists,think about related issues. The general consensus isthat domain-specific mechanisms likely evolvedover millions of years of biological and cognitiveevolution to deal with specific sorts of relevant andrecurrent problems in ancestral environments (‘taskdemands’), such as recognizing inert objects (e.g.,rocks), reducing biodiversity to causally manageableproportions (e.g., species), or anticipating agents(e.g., the intentions of potential predators or prey).In the computer lingo that now dominates the field,the ‘computational mind’ consists of a variety of distinct,task-specific information-processing devices.Nothing from work on subcortical emotions or neocorticalprinciples of general association has provided,or hints at providing, the slightest insightinto the cognitive structures and processes discoveredso far in these emerging fields.Evolutionary psychology is still in its infancyand is not yet the new scientific paradigm somewould love and others hate. Some find the idea of‘Stone Age Minds for a Space Age World’ bold andirreverent, many find it false and demeaning, othersfind it ridiculous. As the field stands now, all maygarner uncertain support for their position. Thisnew field surely will not solve all of the problemsits fervent supporters sayAuthor’s addressScott Atran, CNRS–Institut Jean Nicod, 1bis, avenue de Lowendal, F-75007 Paris,France, and Institute for Social Research,The University of Michigan, 426 THompsonSt., Ann Arbor, Michigan 48106-1248,USA. Email: satran@umich.eduit will. But neither will itface the massive roadblocks to understandingthat its unrelenting detractorssee at every turn.At present, the field is apromissory note, much asDARWIN’s theory was at itsbeginning.Evolution and Cognition ❘ 53 ❘ 2001, Vol. 7, No. 1

Scott AtranLittle by little, biologists were able to deliver onDARWIN’s promises. This process has speeded up considerably—almostirreversibly—thanks to access atthe molecular level. There is still a long way to go.Through recent advances in cognitive science, evolutionarypsychology has gained entrance to mentalstructure, and so potentially to the brain’s evolvedneural architecture. It has an even longer way ahead:much less is currently known about how the mind/brain works than how body cells function. Perhaps,in the end, evolutionary psychology’s interpretationsof complex mental designs as telltale signs ofancient environments will prove no truer than phrenology’sreadings of bumps and other conformationsof the skull as indications of mental facultiesand character (phrenology was a very serious andhotly debated discipline a century ago). Then againperhaps not, which makes the effort worthwhile.Note1 Although the adaptive relationship of structure to functionis often manifest, as with the giraffe’s neck or the rhinocerus’shorns, often it is not. In such cases, evolutionary theoristsadopt a strategy of ‘reverse engineering’. Reverseengineering is what military analysts do when a weaponfrom an enemy or competitor in the arms market falls intotheir hands and they try to figure out exactly how it wasput together and what it can do. Reverse engineering iseasiest, of course, if the structure contains some signatureof its function, like trying to figure out what a toaster doesgiven the telltale sign of toasted bread crumbs left inside.But in many cases recognizing the appropriate signs alreadyrequires some prior notion of what function the structuremay have served. 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Jaak Panksepp and Jules B. PankseppA Continuing Critique ofEvolutionary PsychologySeven Sins for Seven Sinners, Plus or Minus Two 1IntroductionAs the astute reader of ourpaper hopefully noticed,our overall goal was tohelp guide evolutionarypsychology toward amore productive conceptualand experimentalpath than has been characteristicof the disciplineduring its earlier stages ofdevelopment. The ‘moralistic’stance we adoptedwas intended to be an attention-grabbingtactic(as is the current title,which is optimally directedto at least five nonparticipantsmentionedin footnote 1, and wecould easily add or subtracta few additional sinsor sinners, including ourselves).We also chose tobe confrontational due toour belief that progress inthe Evolutionary Psychology2 project should beconstrained by the evidential standard common toall of the natural sciences. Perhaps from the perspectiveof the standard social science model, EvolutionaryPsychology has attained a credible paradigm, butthis is not the case from a cross-species, neuroethologicalperspective. There is abundant room for improvementin our quest for a reasonably accurateand integrated view of human mind/brain evolution.AbstractOf the many issues raised by commentators, twostand out: A debate about modularity and the evolutionof language. A neocortically focussed developmentalsystems perspective, coupled with anunderstanding of the ingrained emotional and motivationalsystems we share with other mammals, willprovide more insights into what types of intrinsicfunctions evolution actually constructed in the humanmind/brain than will any proposed variant ofcortical modularity. For instance, it is unlikely thatour higher cerebral expansions could develop languageabilities without the ancient pre-existing subcorticalsystems that generate socio-emotional urgesand non-verbal, gestural pragmatics. Although humansmay well have ‘communicative instincts’ andeventually ‘proclivities towards language acquisition’,there is little evidence they are born with ‘languagemodules’. It is imperative for evolutionary psychologiststo be constrained by the neuroscience evidencegarnered from all mammalian species.Key wordsEvolutionary psychology, human nature, sociobiology,affective neuroscience, modulartity, language,group selection, social selection, natural selection.Unfortunately, when itcomes to mind/brain issues,no one can yet unambiguouslyspecify theoptimal course of investigation,but if obvious mistakesare not corrected atthe outset, they will haveincreasingly deleteriouseffects as time passes. Forinstance, the erroneousconcept that the wholebrain is a general-purpose,associative learningmachine, guided simplyby reinforcement processes(i.e., the phlogistonof 20 th century behavioralpsychology), had a conceptuallydeleterious effecton mainstream psychology.The notion thathuman cortical evolutionhas yielded a mind/brainconstructed of massivecognitive modules seemsto be a mistaken perspectiveof the opposite extreme(FODOR 2000). Amore realistic view lies somewhere in between. Bothspecific and comparatively non-specific processes residewithin the human brain, and once they interactduring development, especially in higher regions ofthe brain (e.g., neocortex), the epigeneticallymolded neural realities cannot be disentangled ifone sides with either polar view. All evolutionarypsychologists should appreciate that incorporatingdevelopmental systems theory as well as data fromEvolution and Cognition ❘ 56 ❘ 2001, Vol. 7, No. 1

A Continuing Critique of Evolutionary Psychologyrelated species will be absolutely essential for all conclusions—apoint acknowledged by several of ourcommentators (most prominently PITCHFORD).Most importantly, in the target article we arguedthat it is counterproductive to generate claims thatdo not take all the available evidence into account.Non-biologically based accounts for the evolutionaryhistory of the human mind/brain cannot work if assertionsare not consistent with existing neuroscientificevidence. Thus, our guiding principle was thatit is much easier to steer a corrective course duringthe early phases of any intellectual journey ratherthan when the passage is already well on its way. Wethink most scholars will agree that current revolutionaryvariants of Evolutionary Psychology, in theirenthusiasm to modify the prevailing and highlyflawed standard social science model, have also beenmaking a variety of obvious mistakes that now needto be rectified. To the extent that the discipline canachieve corrective action, it will flourish. To the extentthat it cannot, it will increasingly become a scientificcult.One of the greatest risks to future progress is prematureclosure on conceptual issues that are not yetempirically resolved. It is probably fair to say thatwould include essentially everything evolutionarypsychologists have suggested about the mind/brain(including most prominently the neuro-evolutionarynature of language). This is not as obviously thecase in the cognitive, behavioral and affective neurosciences,where the abundance of converging humanand cross-species findings gives greater confidencethat lasting knowledge bases have beenachieved. Evolutionary psychologists should find itextremely beneficial to pay attention to these findings!We were disappointed that there were few commentson the sampler of ‘solutions’ that we sharedin the target article, but we were delighted that RusselGARDNER emphasized the importance of such evidence.It will certainly behoove evolutionary psychologiststo assimilate the facts that have beenaccumulated from ‘lower’ species, even though everyfinding remains open to various alternative neuropsychologicaland neurobehavioral explanations.Since most individuals in neuroscience are receptiveto evolutionary views, it should be informative forevolutionary psychologists to pay close attention totheir interpretation of the neuronal evidence. Indeed,this passage is being achieved on some cognitiveissues (DUCHAINE/COSMIDES/TOOBY 2001; GALLIS-TEL/GIBBON 2001), but not, to the best of ourknowledge, for the abundant knowledge concerningour basic emotional and motivational systems. Suchneglect resembles the disturbing bias evident in thestandard social science model, where evidence derivedfrom our fellow species has traditionally beenmarginalized. Also, since most investigators of humanbehavior do not have causal manipulative accessto the underlying brain systems, they tend toassiduously ignore the relevant neural evidenceculled from other species. However, since we aresuch a contentious and self-centered tribal species,it is hard to coax those interested in more etherealhuman issues to pay attention to such fundamentallyimportant infra-human evidence.Since the publication of our target paper, we haveall experienced the impressive publications of the HumanGenome Projects (LANDER et al. 2001; VENTER etal. 2001) and many have been humbled by the findingthat the best estimate of the total number of genesin the human species was more than halved. Others,like ourselves, have been delighted with the evidencethat at a genetic level, we are not all that differentfrom our fellow mammals—a conclusion that haslong been obvious to many. Thus, our vast intellectual/cognitive abilities cannot be explained by a massivelymore sophisticated genome, but rather by our massivelyexpanded cortex, the mushrooming of which probablyemerged through comparatively modest genetic changesrather than any new and detailed genetic constructions.One should repeat this as another mantra wheneverone ponders what evolution did or did not constructin the highest cortico–cognitive reaches of the humanbrain/mind. Although one might fall back onthe emergence of new gene–gene interactions andheterochronic neural progressions in the human species,we again submit that the simplest explanation isthe flexibility of mind that general-purpose corticalmushrooming provided. The many ancient attentional,emotional and motivational read–only–memory(ROM) type systems we share with other animalsnow have access to more abundant general-purpose,random–access–type memory (RAM) spaces that allowcomplex re-symbolizations of our basic urges.Due to our massive ignorance of how the cortexreally works to generate conscious perceptions,memories, plans and intentions, taken in conjunctionwith the similarities in the local circuits of sixlayeredneocortex in all mammals, we believe thisRAM-type metaphor should be the default assumptionconcerning the function of our heteromodalcortices. Although there is no question that corticalareas become specialized for a variety of adaptiveand even maladaptive functions during development,to assume that specialized, sociobiology-typemodules evolved in the higher cerebral areas via ge-Evolution and Cognition ❘ 57 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. Pankseppnetic selection (which we believe is essential to entertainany hard form of modularity) is so patentlyspeculative and at odds with hosts of facts that itcannot be deemed a very plausible neuropsychologicalviewpoint. It is also a societally dangerous perspective,for it does not leave much room for culturaltraditions in the molding of very basic behavior patterns(DUGATKIN 2000; DE WAAL 2001). In otherwords, the biggest danger with Evolutionary Psychologyis its failure to acknowledge the massiveflexibility and plasticity of the expansive humanneocortex, and its concurrent failure to deal seriouslywith the many functionally dedicated systems(heavily influenced by genetic factors) that do existsubcortically. In saying this, we also recognize theremarkable importance of genetic imprinting inguiding the construction of such brain systems(JOHN/SURANI 1996; KEVERNE et al. 1996).All too often, extreme evolutionary viewpointsfail to acknowledge our capacity for well-reasoned,well-intended actions within the many complexitiesof our social worlds. In any event, it should be clearto everyone who has kept up with modern neurosciencethat the genetic tethers on our basic attentional,emotional, motivational and memorial processesare as great as those of other mammals, but atthe same time our capacity for selfless action is asvast as the epigenetic/cognitive imaginings of ourhighest cerebral spaces. The essential role of subtlesocial and cultural processes in human evolution hasbeen massively underestimated by those who merelyfocus on selfish gene concepts as opposed to individualsand dyads as the true vehicles of evolution.We will respond to our commentators in twoways. First we will address each individual commentary,with considerable repetition of the most salientpoints many commentators chose to focus on, sothat each section can be read independently. Thenwe will move on to several key topics that need to behighlighted and re-emphasized in more detail.Again, we undertook this exercise with the goal ofachieving the most accurate evolutionary picture ofour animal and human nature that is currently possible,and we shall proceed in that vein.Ullica Segerstråle:A Historical PerspectiveWe appreciate the excellent framing for this debatethat SEGERSTRÅLE provided. Her comments help putthe controversy in an appropriate historical context.Her superb recent coverage of the sociobiologywars (SEGERSTRÅLE, 2000) has nicely set the stage fora better appreciation of the present skirmishes. Wewholeheartedly agree with most of the themes sheshares, but would quibble with some of the particulars.For instance, we are not adverse to attempts to“find out about the functioning of the mind/brainby asking questions about adaptations and design”.Our objection is only if such questions are askedwithout considering the understanding that hasalready been attained, as it has in abundance withregard to our basic attentional, emotional, motivationaland memorial abilities. Moreover, we arehappy that SEGERSTRÅLE recognizes the need toincorporate ethology into the foundation of psychologicalthinking. We disagree that ‘real science’is done at the ‘anatomical, physical level’ andgreatly admire theoretical perspectives. Indeed,technologies such as fMRI and PET imaging onlyyield correlations, not causal relationships. Causationcan only be assigned within the framework ofwell-controlled experiments where underlying variablesare manipulated by experimenters.. Andcausal forces are what evolutionary psychologistsseek to discover. As SEGERSTRÅLE recognizes in mostof her comments, our battle is against a scientific‘tribalism’ that ignores broad swaths of the relevantevidence that comes from ‘nonverbal communication’of our fellow animals. She is correct in recognizingthat the study of such shared behaviors,especially their neural substrates, will provide theneeded bridge between the discussion of such issuesin humans and the detailed analysis of the underlyingneural causes in other animals.The case of human facial and bodily expressionsis pregnant with historical antecedents (DARWIN1998) and we simply note that such processes obviouslyhave dedicated neural circuits that evolutionprovided as tools for negotiating social interactions.The nature of these systems, especially in terms ofoverall bodily controls, is not all that different inhumans and other animals (although we typicallyhave much richer facial and vocal expressions).However, while most other animals only have aninstinctual route to such expressions, we humansalso have learned volitional, cortico–cognitiveroutes that are not directly associated with our geneticheritage. Thus, even when damage to thosehigher areas leads to an incapacity in the ability ‘tochoose’ emotional expressions, the arousal of subcorticalemotions still generates overtly expressiveemotional tendencies on both face and body (RINN1984; BOROD 2000). Cortically-mediated facial expressionsare probably not controlled by geneticallydictated‘modules’, although there may be intrinsic,Evolution and Cognition ❘ 58 ❘ 2001, Vol. 7, No. 1

A Continuing Critique of Evolutionary Psychologyperceptual/motor proclivities enabling the cortex tomirror perceived actions (GALLESE/GOLDMAN 1998).SEGERSTRÅLE correctly points out that we do “accusethe evolutionary psychologists of ignoring awhole body of evidence that actually exists”. SEGER-STRÅLE also suggests possible reasons for why evolutionarypsychologists have failed to include datafrom comparative neuroscience in their recent syntheses(COSMIDES/TOOBY 2000). While we agree thatnurturing a new scientific field is taxing, we feel thatfailure to include the large body of neuroscientificdata in attempting to explain the ultimate biologicalcausation of human behavior cannot be attributedto the business of a ‘multi-front battle’—it is simplyneglect (probably arising largely from the traditionalsocial science bias which tends to intentionally ignorefacts derived from other animals). We also appreciatethat in doing this, evolutionary psychologistsare no different than many other cognitivepsychologists who developed new and dubious anthropocentric,intellectual traditions after they becamefed up with the all too many abundant sins ofmany 20 th century behaviorists (for an overview, seePANKSEPP 1990).We sympathize with the multi-front battle thatEvolutionary Psychology presently has to wage withmany other disciplines. We who believe that studyinganimal mind/ brains will tell us a great deal abouthuman nature have an even larger challenge to overcome.Our battle is against the prevailing anthropocentricperspective of evolutionary psychology, notto mention psychology in general. Once it is acceptedthat many of the sociobiology-type systemsthat humans possess reside in homologous subcorticalareas we share with other animals, the temptationto find ‘modules’ in the higher reaches of thehuman mind/brain should diminish. This is theneurodevelopmental issue (and blessing) that liberatesus from any rigid determinism. Again, we haveno doubt that there is a great deal of sensory andmotor specialization in the cortex, including face-,odor-, posture-, touch-, and voice-recognition dispositions,but we see no evidence that these functionsare sociobiological in any deep sense of the word,nor are they uniquely human. Yes, language comesclosest to a cortical, sociobiological ‘module’, but letus appraise the evidence cautiously. As SEGERSTRÅLE(2000) understands as well as anyone, much of theconfusion in this area has arisen from our politicalurges and convictions (often unstated) rather thanfrom our willingness to confront all the evidence inunbiased ways. As she says, it is indeed time to “relaxthe modularity idea”.Linda Mealey: What’s in a Word?Linda MEALEY provides a useful and balanced overviewof the general arguments, as well as variousparticulars. She gracefully navigates the pragmaticmiddle course, highlighting important empiricaland conceptual landmarks along the way. We willonly reinforce her views by elaborating on a coupleof related topics that concern us most. We also haveconsiderable concern with her advocacy of the traditional‘modular’ point of view that, in our estimation,adds nothing substantive to an evolutionaryanalysis of brain functions (even though it mayhave seemed like a good semantic way to start in themidst of massive ignorance).Of all the ‘sins’, the species-centric perspective isforemost on our minds. It is simply unconscionable,considering all that we presently know about thegenetic and epigenetic construction of the brain,that throughout the past three decades practically allvariants of psychology have ignored the evidencegained from our fellow creatures (at least as a functionof published words). The failure of academicpsychology to assimilate information from the neurosciencerevolution has been documented severaltimes via citation analyses, and many of us believethat psychology simply cannot be a solid basic scienceunless it begins to incorporate such issues intoits framework.This complaint should be especially poignant forevolutionary psychology, for many of our mind/brain abilities are shared with other animals. Eventhough we clearly outdistance most other species inthe complexity and refinement of our cognitiveabilities, To the best of our knowledge, the mushroomingof the requisite human neocortex occurredin response to fairly minimal genomic changes. Inother words, our cortical expansions are the resultof a relatively modest number of heterochronic genetic/regulatorychanges rather than any type ofmajor re-engineering that accommodated new anddistinct cerebral skills (even though such skills mostassuredly do emerge from the epigenetic interactionsof our massive brains with our bodily abilities).In short, the highly specialized functions of the oldmammalianbrain, situated largely subcortically,took most of evolutionary time to be constructed. Ifwe truly want to scientifically understand the foundationof human nature, a proportionate amount ofresearch effort should be devoted, in psychology, toelucidating these functions. Instead, within psychologysuch efforts have been a drop in the bucket.Only a few investigators gathered around the deepEvolution and Cognition ❘ 59 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. Pankseppwell of psychological ignorance (see FODOR 2000)are paying attention to how rapidly other relevantneuroscientific wells are filling with substantiveknowledge for understanding the human condition.All of this important mind/brain work has beenconducted without the use of terms such as modularity.Although MEALEY is correct that our view doesaccept the importance of brain systems, networksand circuits as being of foundational importance forunderstanding mind, for many of us, the term modularityis a pretentious new-comer that adds nothingto any ongoing analysis. Indeed, it gives usfaulty images of ‘encapsulated’ information processingsystems that greatly oversimplify highly interactivemind/brain functions that are most rightfullyanalyzed in neurobiological terms. A similarmistake was made in early neuroscience when ‘centers’were used as common concepts, but that wassoon dropped (except among simplifiers and popularizers)when it was realized that all psychologicalprocesses emerged from distributed circuits that interactedwith many other functional systems. Sinceevolutionary psychologists need to come to termswith the organic processes of the brain, they wouldbe wise to discard terms that serve no useful empiricalor didactic purposes.Even though we should certainly not ‘speak ofnature versus nurture’ or ‘emotion versus cognition’,we also should not forget that such distinctions serveas heuristics for certain research endeavors, especiallythose that dip below the surface of psychologicalexperience, to the neurobiological substratesfrom which such processes arise. Although genes obviouslycannot act independently of environments,genes can guide the creation of an enormous varietyof creatures and a diversity of mind/brains within awide range of supportive environments. The distinctionbetween emotion and cognition helps those ofus working at the level of the brain to focus on thenature of affective brain processes that desperatelyneed more empirical attention (PANKSEPP 1990,2000a, 2001a). However, if we only study the psychological/behavioralresponses of organisms, thensuch distinctions seem difficult to justify and investigatorsmay prematurely discard them, althoughthey are still useful in other (e.g., neuro-evolutionary)contexts.MEALEY eloquently argues that the worth of ourpsychological concepts ultimately hinges “not ontheir ultimate ability to ‘reflect’ reality, but theirpractical value”. She asserts that “if their use encouragesinquiry, they are a good thing; if they discourageinquiry, they are a bad thing”. Being pragmatists,we agree with that, but would demure a bitwhen we consider the neurobiological groundingthat is absolutely essential, each and every moment,for our status as creatures of the world. Psychologistsare not accustomed to recognizing that the neuralanalysis provides a vast, new world of ‘reality’ testing.Once we begin to consider all aspects of thesystems in which we are interested, we will needmany new concepts and perspectives to parse ‘reality’and we will have to discard others. In neuroscience,the concept of modularity is a non-starter,for it provides no new and useful distinctions. Atbest, it seeks to replace more informative terms thathave been used for decades (e.g., circuits, systems,faculties), while bringing along a great deal of extraand unneeded conceptual baggage. On the otherhand, concepts that have long been discarded, suchas ‘drives’ may need to be resurrected, for it seemsclear from the neuroscience evidence that as we manipulatebasic motivations, synthesis of variousneuropeptides is genetically induced in the brain tofacilitate the energization of specific behavioral tendencies(PANKSEPP 1993, 1998a). We certainly supportpluralism in psychological research, but not ofthe variety that neglects (and too often shuns) themain organ of mind and behavior.Carlos Stegmann:A Philosophical PerspectiveFrom a philosophical/humanistic perspective wesympathize with the views advanced by STEGMANN.From a practical scientific point of view, we do not,for such holistic views often frustrate any systematicexperimental analyses of brain functions. A satisfyingholism that postulates different levels withinhierarchical systems reflect ‘one–and–the–samething’ does not advance the analytical discriminationsthat are essential for the scientific enterprise.Also, we note that there is massive aversion withinthe psychological community for accepting the ideathat within the historically layered and blendedcomplexities of the evolved nature of the mind/brain “each stratum must be reducible to the lawswhich apply in the hierarchically lower strata”.Although we do believe there are bound to be strongrelations among such strata within the brain, weprefer to see them through the modern philosophicallens of supervenience—whereby lawful relationshipsat one hierarchical level will be systematicallylinked, but not completely reduced, to lawful relationshipsat lower levels (KIM 1993).Evolution and Cognition ❘ 60 ❘ 2001, Vol. 7, No. 1

A Continuing Critique of Evolutionary PsychologyWe also acknowledge STEGMANN’s concerns withour advocacy of ‘group selection’, for we do believethat pro-social processes must ultimately be instantiatedat an individual (and perhaps dyadic) selectionlevel. We raised this issue partly as an enticementto radical selfish-gene selectionists, assumingthey are still out there, rather than from any deepconviction that group selection, as traditionally understood,is a robust vector in animal evolution (see“Group Selection” section below)Our advocacy is based on the recognition that inthe evolution of social processes, brain mechanismsmay have emerged which only operate optimallywithin certain social contexts and some may be sufficientlygeneral that they may provide a competitiveedge even in the context of outright altruisticbehaviors. For instance, since our social bondingmechanisms appear to be designed in such a way asto be based more on learning than intrinsic kin-recognition,it is easy to envision why humans mayadopt the children of strangers, even though that isnot to their direct genetic advantage. It may simplybe to their emotional advantage, even though itmay be easy to theoretically (and in our estimationarbitrarily) postulate that it may provide some potentialgenetic advantage because of the massivelyshared genetic heritage that all humans share. Inany event, we believe that affective feelings are themain psychological currency which guides the behaviorof individuals in genetically advantageous(i.e., survival and reproduction promoting) directions.We look forward to reading STEGMANN’s morecomprehensive description of his position, and wewill be especially interested in seeing how his ideasmight be subjected to empirical tests. Without clearpredictions, all views, no matter how holistically attractivethey may seem at semantic levels, will remainscientifically sterile. We trust that STEGMANNhas taken his interesting analysis to the predictivelevel. Explanation and synthesis without the precisionof new predictions can all too easily appeal toour intellectual–aesthetic senses while not contributingmuch to the rigor of our analytical endeavors.Indeed, that is one of the ‘siren-song’ appeals of EvolutionaryPsychology—it provides such a simple andoutwardly credible conceptual umbrella, that oftenthe reductionistic strategies we must develop to scientificallyunderstand the true nature of the underlyingprocesses become of secondary importance.The goal of our article was largely to bring variousneglected scientific issues to the foreground of discussion.Russel Gardner:A Neuropsychiatric PerspectiveIn his commentary, GARDNER highlights the desperateneed to fully integrate perspectives for the studyof matters concerning the brain and mind. Typicallythey are still considered separately, and one of themajor challenges he poses is to build bridges thatlink those disparate realms in ways that are subjectto scientific evaluation. Of course, that has been themain goal of affective neuroscience (PANKSEPP1998a). Indeed, the cardinal scientific motivation ofthe senior author has been to develop credible scientificstructures that may help highlight the deepnature of psychiatric/emotional disorders in thehuman species. This project, at its outset, recognizedthat there seems to be no other robust scientific pathto understanding the underlying emotional andmotivational dynamics of the human brain thanthrough the detailed analysis of homologous processesin related animals. The details of the humanbrain are simply too inaccessible, but faith in theexistence of deep evolutionary homologies hasalready paid off handsomely in biological psychiatry(CHARNEY/NESTLER/BUNNEY 1999), and it will continueto do so at an ever increasing pace as investigatorslearn how to effectively modulate neuropeptidesystems (PANKSEPP 1993, 1998).The supposition that we can understand the essentialoutlines of our own evolved emotional processesby studying the brains of other animals has not traditionallybeen well received by the psychologicalcommunity. For many, it remains too much of a theoreticalleap to believe that other animals experienceemotions. However, on the basis of evolutionary considerationsas well as massive anatomical and neurochemicalhomologies, this possibility is now a solidhypothesis. It is also one that can now be experimentallytested because the neurochemical discoveries inother animals can lead to pharmacological interventionsthat generate predictions and can be evaluatedin humans. For instance, the abundant work on corticotrophin-releasingfactor implicating it in anxietydisorders and depression, has led to concordant findingsin humans (NESTLER 1998). There are many otherexamples (CHARNEY et al. 1999).GARDNER also provides a didactic service by highlightingthe history of such approaches in the psychiatriccommunity, noting how ‘selective allegiances’among powerful academic circles have probably retardedthe bridging work that needs to be done. Theemergence of evolutionary psychiatry, partly underthe umbrella of the Across-Species Comparisons andEvolution and Cognition ❘ 61 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. PankseppPsychopathology (ASCAP) group, has been a clearforce in providing new perspectives for future developmentsin biological psychiatry. While DSM-IV diagnosticshave stabilized into a practical and economicallyuseful way to deal with mental disordersin our society, it should not be seen as anything closeto the final scientific word. The goal now should beto link many emotional disorders to the activity ofspecific brain systems. The conceptualization of thebasic emotional systems of the brain (e.g., PANKSEPP1998a, 2000b, 2001b) should be an essential componentof future understanding in this area.GARDNER also highlights several higher-order emotionalissues that should be fascinating to evolutionarypsychologists. The ‘alpha communication’ stylethat is evident in manic patients is also evident inmany scientific discussions. Given the choice, practicallyall scientists and scholars are more eager to talkabout their own work and ideas than to listen to thestories of others. This urge to talk rather than listento the next fellow is a very prominent feature of ourspecies and it needs to be explained by the evolvedfeatures of our communicative apparatus. We suggestthat such urges are much more dependent on subcorticalemotional systems (e.g., dominance mediatingones) than intrinsic ‘drives’ of cortico–cognitivebrain systems that epigenetically specialize in languageskills. We note that modern neuroscience hasdemonstrated, in abundance, the extent to whichseemingly pre-ordained cortical functions can betaken over by other functions, as MEISENBERG asserts,in a ‘guns for hire’ fashion.In any event, there will be many interesting clinicaland evolutionary psychological predictions to bemade once we fully understand the primitive sourcesof our emotionality. It is noteworthy that groups suchas ASCAP have emerged to provide “understandable,unfanciful, common-sensical and data-based explanations”of human behavior. In this vein, we also notethat the recent emergence of a neuro-psychoanalyticmovement that has deep respect for the primitiveemotional systems that all mammals share, is anotherharbinger of a new age of reason in the way we studythe human mind/brain (see http://www.neuropsa.com).It also is not shy about fully consideringevidence collected from our fellow species.Gerhard Meisenberg:Modularity is Alive and Well… SupposedlyMEISENBERG seems to agree with our overall thesisand takes the opportunity to provides a wide-ranginganalysis of instances of cortical functions thatneed more attention than we provided, as well asquestioning some of our specific conclusions. Thisrepresents the type of discussion that needs to beexpanded in evolutionary psychology and the keyissue from our perspective, once again, is the degreeto which higher neocortical functions are modular(genetically-dictated functions) as opposed to experientiallymodularized (epigenetically derived specializations).We have no doubt that a great deal of the neocortexbecomes highly specialized (‘modularized’?) as afunction of experience, explaining the power of theKennard Principle in neurology (functional recoveryis more complete following brain damage in youththan in old age), but aside from a variety of discretemotor and sensory/perceptual processes (many ofwhich MEISENBERG elaborates upon, with which wehave no disagreement), we still want to suggest thatno sociobiology-type, cognitive ‘modules’ are yetclose to being demonstrated (as our original intentwas to distinguish examples such as facial/emotionalrecognition and appreciation capacities fromthe more cognitive types, which may not have beensufficiently clear in our target article). Certainly, verybasic sensory and motor processes became specializedin the neocortex of ancestral mammals a verylong time ago, but our main point was that there isessentially no evidence yet that modularized, sociobiologicalcognitive adaptations have emerged inthe neocortex since our neocortical divergence from‘chimp-brained’ anthropoids some two millionyears ago. It presently does not appear that most ofwhat evolved in the neocortex since then was intrinsicallymodular, but rather, general purpose tissuethat could become modularized under the influenceof environmental factors interacting with functionallydedicated limbic and more primitive subcorticalnetwork functions as organisms mature. To ourknowledge, there is no credible evidence against thistype of epigenetic–developmentalist view. It is ofcourse important from a neuropsychological perspectiveto analyze these developmentally emergentspecializations since they are incredibly importantin neurological practice. However, in our estimation,such evidence does not really relate directly to theissue of evolutionary modularity.We have no conceptual disagreement withMEISENBERG when he discusses the emergence ofsmiling (which can actually be exhibited by newborninfants), but we disagree with the suggestionthat the relevant neurology is merely concentratedin the cingulate cortex. There is as much evidencethat the insula and several other cortices participateEvolution and Cognition ❘ 62 ❘ 2001, Vol. 7, No. 1

A Continuing Critique of Evolutionary Psychologyin emotional facial expressions (BOROD 2000; LANE/NADEL 2000) and we are tempted to look for the executiveinfluences over facial emotional displayseven in lower areas of the brain, as seems to havealready been demonstrated for taste induced expressionin a variety of animals (BERRIDGE 2000; STEINERet al. 2001).We agree that a strict dichotomy between corticaland subcortical processes is ‘too simple’ and wasused as a convenient didactic device for efficientcommunication. However, we believe the overalldistinction, with some obvious caveats, remainsboth conceptually and experimentally ‘useful’.Again, the Kennard Principle tends to be less manifestfollowing damage to many subcortical areas—where early damage is generally more devastatingthan damage occurring later. This is probably becausethe essential subcortical functions organismsare born with have already been partially remappedto higher areas as a function of hierarchical neuraldevelopment. Clearly, there is a continuity of functionsalong the whole hierarchical gradient of theneuroaxis, going from specific hard-wired functionsto ever increasingly flexible soft-wired (developmentallyprogrammed) ones, but the simplifying dichotomyforces us to face up to the important fact thatthere are an enormous number of genetically preordainedfunctions in subcortical regions and a comparativepaucity (at least in terms of discrete functionsper unit of brain matter) in neocortical ones.Our main complaint is that the semantics of modularityare very misleading, especially when such entitiesare seen as ‘encapsulated’ special purpose, cerebralfunctions interacting in linear cascades. Thereare no such modules, only highly interactive circuitsand widely distributed networks with many overlappingfunctions (i.e., the semantics of modularityadds nothing to our capacity to discuss brain functionsthat is not already available in more classicneural systems terminology). These networks are obviouslymassively interactive in divergent, convergentand reentrant ways, with feed-forward andfeed-back components that make any linear processingimage essentially wrong and not even heuristic(FREEMAN 1999). At the same time, in the midst ofsuch complexity, we strongly feel that productiveinvestigators will have to continue to invest in simplification,generally recognizing the strategic powerof Descartes’ 3rd rule of science—to think in an orderlyfashion when concerned with the search for truth, beginningwith the things which were simplest and easiest tounderstand, and gradually and by degrees reaching towardmore complex knowledge, even treating, as thoughordered, materials which were not necessarily so. Wethink most investigators realize that the scientificenterprise would be immobilized if there were noattempt to simplify the true complexity of nature.Perhaps our strongest disagreement with MEISEN-BERG is his contention that the evidence at the subcorticallevel of analysis is not as robust as we makeit out to be. In our estimation, the evidence for subcorticalsystems (not ‘modules’!) is enormous (PANK-SEPP 1998). Of course, there is room for controversyabout the interpretation of specific findings, so let ussimply respond to a few raised by MEISENBERG: (1)Although there are abundant debates about details,our current understanding of the mesolimbic appetitivemotivational systems (not ‘pleasure’ systems) istruly remarkable, and the role of negative affectiveprocesses in engaging such systems have been dealtwith in theoretically coherent ways (for overview,see IKEMOTO/PANKSEPP 1999). (2) In mammals, serotoninis a general state control system that modifiesevery behavior ever studied and it is possible to envisionthat this system primarily helps keep informationchannels of the brain reasonably well segregatedfrom each other and helps tone down theimpact of all emotionally challenging stimuli (PANK-SEPP 1982, 1986). (3) In addition to the fine work onfear that MEISENBERG cites, there is abundant evidencefor subcortical anger–rage processes, sexual systems,nurturance systems, as well as those for separation–distress and play (albeit all these urges are effectedthrough the emergence of cortico–cognitive strategies).(4) The intensity and time-courses of socialbonding does vary as highlighted by MEISENBERG’smention of the now classic differences between thesolitary montane- and the highly social prairie-voles,but so far these seem to reflect differences in the underlyingintensities of socio–emotional systemsrather than difference in kind. (5) There is also abundantevidence for very specific systems to regulateenergy balance, water balance, ionic and thermalbalances, not to mention various hormones andgases. All of the above issues are extensively discussedin PANKSEPP (1998a).In sum, we think MEISENBERG’s arguments add importantdimensions that need to be considered inthe maturation of evolutionary views. We agreewholeheartedly with his striking image of manyhigher areas of the brain being ‘guns for hire’ whichis a striking metaphor for general-purpose learningtissues. Again, we disagree with his assessment thatthe evidence for subcortical systems remains tenuous,but admittedly, the ongoing research efforts onthose issues remain too modest and an enormousEvolution and Cognition ❘ 63 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. Pankseppnumber of details remain to be empirically resolved.However, the evidence that these systems are decisivefor the sociobiological competence of animals israther overwhelming. May we simply recall a blatantearly example—a description of cats that had receivedcomplete lesions of the periaqueductal graymatter of the midbrain: “If the lesion is extensive,the cats lie inert, silent and flaccid as a wet rag. Theynever again show any interest in food, but must benourished by stomach tube. They have no spontaneousactivity. They will sometimes swallow if milk isallowed to run down in the back of the throat. Irritationwhich would cause immediate outcry from anormal cat provokes only feeble movements of thehead or limbs.” However, after a few days of recovery,an animal “begins to right itself, first the head andthen the entire body. Body reactions are elicited normally.It may even walk slowly about on occasion ifstimulated, but has no spontaneous activity, attendsto nothing in its environment and never feeds itself”(BAILY/DAVIS 1943, p306). Nothing comparable canbe obtained with even extensive damage to the neocortex.The intrinsic ‘vitality’ of the subcortical terrain,including its importance for the direct generation ofa large variety of affective states, should not be underestimatedif we truly want to understand basicpsychological processes. The capacity of the cortexto ‘learn’ new strategies based upon the dictates ofthe primary-process emotional and motivationalmechanisms of subcortial areas is also truly remarkable.However, any claim that an abundance of Swisspocket-knife type, intrinsic, inclusive fitness types ofcognitive strategies have evolved within the circuitryof the neocortex during the last few millionyears of human brain/mind evolution remains geneticallyimprobable and certainly unsubstantiated.We welcome anyone providing evidence for suchpossibilities, but in their absences, we need not beconvinced of an intrinsically modular organizationsimply through the recitation of well-establishedneuropsychological specializations that are thestock in trade of neurologists working on the maturehuman brain. We think all neuroscientists acceptthat the mature human brain has abundant specializations,as does, to a much lesser extent, the newborninfant, but we may ultimately find that manyof these ‘genetically provided tools’ are, in fact, developmentallydependent on the specializationsthat exist subcortically in infants. Of course, all intrinsicfaculties of the mind/brain get massively refinedby the experiences of each and every child. Theplasticity clearly forces us to consider that the natureof our socio–cultural environments that we constructfor our children are more important than speculationsabout that stratetgies that EEAs may haveprogrammed into those neuronal spaces (PANKSEPP2001a). Thus, an evolutionary psychological analysishas yet to add anything to the well-establishedneuropsychological evidence and it now seems, alltoo often, to be diminishing our appreciation of thevast plasticity of human potential.Ian Pitchford:A Challenge From an InsiderAlthough we acknowledged the probable existenceof a variety of specialized functions in the neocortex(even though, once again, the term ‘module’ doesnot help us think about them in any scientificallyuseful way), we did strongly argue for the importanceof several subcortical systems that mediateemotions and motivations, and allow the cortex tofunction properly. To the extent that GARDNER seeshope in the neuroevolutionary psychobiology programof affective neuroscience, PITCHFORD seems todismiss it as a misguided enterprise. However, hisremarks were restricted to a small subset of issuesthat we addressed. Moreover, being the guidinglight of an evolutionary psychology list server(nibbs-newsletter@yahoogroups.com associated withhttp://www.human-nature.com/darwin/index.html),PITCHFORD may have been tempted to give us astronger tongue-flogging than he otherwise mighthave (at least that is suggested by some personalcommunications: vide infra). Accordingly, we see hiscomments partly as a ‘tribally’ motivated attempt tocapitalize on select aspects of the present debate bysetting up unrealistic polarities while ignoringmany of our main arguments. In any event, he discussedonly one of our seven criticisms, namely the“search for adaptive modules where there is onlygeneral-purpose computational space”, using languageprocessing and the role of developmentalprograms as his main exemplars.For didactic purposes we constructed perhaps toostrong a dichotomy between subcortical and corticalfunctions (as already indicated in our response toMEISENBERG), but that is easy to do when one haspersonally studied how behaviorally robust and outwardlynormal certain animals are after being surgicallydeprived of their neocortical tissue soon afterbirth (PANKSEPP et al. 1994). Comparable damage tosubcortical areas yields a terminally debilitated animal(e.g., see BAILY/DAVIS 1943 description above).In no way is our position ‘dualistic’. The existence ofEvolution and Cognition ❘ 64 ❘ 2001, Vol. 7, No. 1

A Continuing Critique of Evolutionary Psychologyadaptive, subcortical systems neither precludes or diminishesthe importance of cognitive/neocorticalprocesses in guiding animal or human behavior. Wepostulate that many of the unique human mind/brain processes arise from dedicated subcortical systemsand flexible neocortical systems interactingwith each other, as well as human culture. Moreover,we stand behind our conclusion that the epigeneticmolding of cortical functions is far greater than theadmittedly substantial experiential influences thatalso affect subcortical functions (for a more developedcoverage of our developmental views, see PANK-SEPP 2001a).PITCHFORD suggests that we have not adequatelyabsorbed the prescriptions of developmental systemstheory (e.g., GRIFFITHS 1997; OYAMA 1985)—that weare talking about ‘genetic blueprints’ rather than acceptingthe full scope of the developmentalist challenge.We only suggest that he re-read what we haveactually written about the matter and we suspect hewould be hard put to find as strong an advocacy ofthe brain developmental systems viewpoint as oursamong any other evolutionary psychologists. Someof our past writing on the topic can be found in PANK-SEPP/KNUTSON/PRUITT (1998), PANKSEPP/MILLER,(1996) and PANKSEPP (2000a, 2001a). One of our primarytheses is that irrespective of the complex interactionsbetween genes and environment, operatingsystems that give rise to affective states such as angerand hunger (as well as several others) will invariablyreside in subcortical areas of human and mammalbrains. How these systems are manifested and resolvedat the behavioral level will undoubtedly dependon an animal’s life history, but they will invariablybe manifest through the emerging plasticity ofhigher cortico–cognitive brain systems. The subcorticalexecutive systems for emotional/affective processeshave persisted because they stochastically supportbehavioral phenotypes that produce ‘constant,stable outcomes’ as adaptive solutions to environmentaland social demands. As PITCHFORD notes, ontogeneticevents in the form of gene/environmentinteractions support a large spectrum of phenotypes.Genes may not be ‘blueprints’ per se, however theycan serve as reliable guidelines that promote ‘constant,stable outcomes’ within the general contextsthat their corresponding phenotypes were selected.The blueprint-perspective of genes that PITCHFORDcriticizes, especially with regard to its use of the word‘innate’, is a terribly important issue to clarify. Wealso do not support the notion that the genetic materialcontains pre-determined outcomes. However,genes do carry a certain likelihood of contributing tocertain phenotypes. For example, genes that encodeserotonin-binding membrane proteins invariablycontribute to the expression of natural behaviorslike feeding, mating and aggression (LUCKI 1998).The contributions of such proteins are determinedby complex gene/environment interactions (LESCH/MERSCHDORF 2000). In this regard, nucleotide polymorphismsare inherited characters that can appreciablyalter a gene’s expression (for instance, see LE-SCH et al. 1996). However, genes are also up- anddown-regulated through environment and experiencedependent modulation, which can yield similarresults (LESCH/MERSCHDORF 2000). The guidelinethat a genome carries for specifying the amount ofserotonin transporter proteins on the membranes ofneurons comprising aggression circuitry, for instance,can and will be altered by environmental demandsand individual experiences.As originally stated in the target article, we continueto reiterate PITCHFORD’s affirmation that evolutionarypsychology is well advised to root itself inthe conceptual framework of developmental systemstheory. However, environmentalism is boundto be rather less influential when it comes to thesubcortical neural terrain—at least with regard to thetypes of neural systems that exist, although their intensitieswill no doubt be influenced by differentialearly experiences (PANKSEPP 2001a).As PITCHFORD may have detected, we do disagreewith certain variants of developmental perspectivesas advocated by some members of the philosophicalcommunity, who seem to relegate DNA to less of an‘informational’ molecule than most biologists areprone to agree. We feel that if one aspires to understandthe nature of biologically-evolved functions/adaptations, then it is wiser to put a majority of theirscientific effort into molecular biological approachesand the true study of ontogeny in neural systemsrather than strictly to the study of external issues,even though we acknowledge how incredibly difficultthey are to extricate.Moreover, we hopefully all will agree wholeheartedlywith regard to the vast plasticity of corticallearning functions. However, as we will note towardthe end of this commentary (in the “Behavior-Genetics—AnEssential Key” section), carefully harvestedheritability data does highlight that the variabilityof our basic internal dispositions arecontrolled more by genes than by external environments,while our overt behavior is guided muchmore by learning. This is an issue that the modularityadvocates of evolutionary psychology have yet toeffectively discuss or study.Evolution and Cognition ❘ 65 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. PankseppIn sum, although we fully subscribe to the importanceof developmental landscapes in moldinghigher mind/brain capacities, we do not agree withthe full revolutionary fervor of the ‘Ontogeny of Information’critique of genetic influences. The genesare more influential in the construction of organismsthan the classic OYAMA (1985/2000) type ofview seems to accept. We would be surprised if PITCH-FORD would disagree. Surely, we all now agree thatgenes can do nothing without supportive environments.However, a remarkable amount of organismiccompetence naturally unfolds from the genomeand the resulting internal milieu as long as a minimallysupportive external environment is present.There is no comparable robustness that has everbeen demonstrated through variations of prenatalenvironmental factors, at least within a range thatmost reasonable investigators would deem to be normative.However, in pointing out this obvious fact,we do not think that it forces us into the position ofperpetuating the ‘nature versus nurture dichotomy’.As we said in our article, developmental systems theoryis an incredibly important concept, but that inno way should be taken to diminish the importanceof genetic factors. If PITCHFORD does, in fact, believethat external environmental issues are as importantfor the functions of subcortical areas as for corticalones, we disagree with him based on our reading ofa massive amount of available neurobehavioral literature.If he believes that environmental factors aremore important than genetic ones in molding cortico–cognitivefunctions, we would agree with him.PITCHFORD shares an abundance of recent new informationabout cortical issues, but those do not relateclearly to our claim that much of the specializationof the cortex is due to self-organization throughdevelopment whereby functionally dedicated subcorticalsystems interact with a remarkably plasticcerebral mantle. Nothing about the interesting anatomicalissues he summarizes relate to that key issue,and in reading his extensive anatomical coverage,we felt he was creating a verbal smoke-screen (or ‘baitand switch’ as Dan DENNET 1995 might say) ratherthan dealing with the issues we had raised. Thoseanatomical issues are most interesting and important,but they do not unambiguously relate to eitherdevelopmental or intrinsic functional issues. Furthermore,we would note that although PITCHFORDcites work that reports humans have a brain size overthree times the size of a non-human primate of similarsize (RILLING/INSEL 1999), there has been no selectiveincrease of relative frontal lobe size within thehuman species (SEMENDEFERI/DAMASIO 2000). He doesnot cite a single functional study that points clearlytoward a reasonably self-sufficient, evolutionarilydictatedcerebral module that underlies a psychologicalprocess that evolutionary psychologists havepredicted.We are also fascinated by the spindle neuronsfound in the anterior cingulate by NIMCHINSKY andcolleagues (1999) and their potential (though yetundemonstrated) role in socio–vocal communicativeintent (an anterior cingulate function we notedin our target article). We suggest that of the manyspecialized cerebral functions that have been discoveredby neurologists and neuropsychologically-orientedinvestigators during the 20th century, nonehave yet been clarified any further by evolutionarythinking (but see BARKLEY 2001 for some changingcontextualizing trends). In any event, it is more importantfor evolutionary views to generate new neuropsychologicalfindings rather than simply providinga new context for discussing findings discoveredby other disciplines. In this realm, the emerging relateddiscipline of neuro-psychoanalysis seems to befarther along in pursuing such issues (SOLMS 1997;SOLMS/NERSESSIAN 1999; KAPLAN-SOLMS/SOLMS 2000)and we believe such contributions highlight the typeof work that evolutionary psychologists should beginto pursue in earnest.As soon as evolutionary psychologists begin togive us new information about the neuroanatomicallocation, as well as the underlying neuro- and psychodynamicsof their evolved ‘modules’, they shallhave provided biological evidence for what biologicalevolution has created in the human mind/brain. Intheir absence, claims of evolved cortical modularitymust be deemed insubstantial, no matter how manyindividuals have imprinted on the idea. However, weheartily encourage such discoveries and the applicationof the armamentarium of new scientific toolsthat are now available for the study of such problems(TOGA/ MAZZIOTTA 2000). But please remember thatcorrelations do not clearly reflect causations.Although certain human proclivities, such as languageand theory of mind abilities, are surely linkedto our higher cerebral abilities (GALLESE/GOLDMAN1998; BARKLEY 2001), it will be equally important todetermine how those mind/brain functions arelinked to the more ancient systems that we sharewith the other animals. Indeed, we believe that humanfacility for spoken language may be deeplylinked to evolutionarily-derived, emotional/communicativepragmatics exhibited by other mammals.In our cultural evolution, it is certainly possible thathighly-prosodic, sing-song languages emerged longEvolution and Cognition ❘ 66 ❘ 2001, Vol. 7, No. 1

A Continuing Critique of Evolutionary Psychologybefore we developed the practice of communicatingin more measured, less dynamic spoken tones (WAL-LIN/MERKER/BROWN 1999). However, since we knowso little about the neurodynamics that create the capacityfor speech, we agree with those modern linguists(e.g., LIEBERMAN 2000) who believe that our facilityfor speech is an outcome of primitivesubcortical systems for communicative intent interactingwith comparatively more plastic, higher corticalsystems through complex developmental trajectories.Yes, human speech functions do tend to getfocussed in specific regions of the brain during development,but the question that we must all now consideris how such specialization actually emerges. Wethink that aside from cortico-limbic participation(e.g., the anterior cingulate systems for communicativeintent), most of language may emerge via epigeneticplasticity in fairly general-purpose, but highlymulti-modal learning circuits of the neocortex. Certainsubcortical systems (e.g., the basal ganglia) maybe decisive in the emergence of such higher abilities(LIEBERMAN 2000; BREITENSTEIN, et al. 2001).We suspect that PITCHFORD would generally agree,even though he is adverse to admitting that subcorticalfunctions are important for evolutionary psychologiststo consider for as uniquely a human activityas propositional language. To what extent“language may have adapted to the emerging complexitiesof the brain rather than the other wayaround” (p111 of target article) remains an open issue,no matter how strongly certain individuals (includingourselves, of course) believe in certain evolutionaryscenarios. And surely PITCHFORD would agreethat our ability to read and write, not to mentionmost of our numerical abilities (beyond primitivecounting), are largely culturally induced skills thatmay have rather little do to with specific evolutionaryselection for such abilities (even though we suspectthat behavior geneticists could certainly select forsuch skills by increasing the speed and quality of certainhigher cortico–cognitive abilities). If so, are thesewidespread skills of the members of human societiesexamples of adaptations, exaptations or spandrels?To some extent, and understandably, PITCHFORD’sflogging of our view may be based on his failure tobe conversant with what the senior author has writtenin the past (e.g., PANKSEPP 1998a, 2001a). Just tohighlight how committed we are to the developmentalsystems view, we refer PITCHFORD to the seniorauthor’s vigorous advocacy of treating ADHDwith environmental approaches, and the ongoingfight against prevailing pharmacological interventionsfor ‘normal’ childhood impulse-control problems(PANKSEPP 1998b,c). It is quite remarkable to usthat in the same sentence, PITCHFORD indicates thatthe accomplishment of Evolutionary Psychology (i)“compares extremely favorably with that based onthe neurochemical individuation of traits and disordersadmired by” us, but which have (ii) “producedso many unsatisfactory theories and treatments(VALENSTEIN 1998)”. Is he suggesting that EvolutionaryPsychology has also produced an abundance ofthe latter?In any event, we believe that VALENSTEIN’s interestinghistorical analysis of the biological psychiatryrevolution was based on an older era of psychopharmacologyrather than the future neurochemical revolutionwe were discussing based on emergingknowledge concerning the highly specific functionsof various neuropeptides (PANKSEPP 1993; 1998a). Wealso suspect that VALENSTEIN’s analysis was also influencedas much by his personal socio–politico–philosophicalleanings as his scientific ones (for more onthis issue see the “Politics and Tribalism” section).For instance, the senior author’s long-term advocacyof the foraging/SEEKING/expectancy view of lateralhypothalamic reward (PANKSEPP 1981, 1982, 1998a)was never supported by VALENSTEIN, whose work onthe plasticity of those systems originally threw thisfield of inquiry into theoretical chaos (VALENSTEIN/COX/KAKOLEWSKI 1970). Only when work by his ownclose colleagues suggested a very similar view (BER-RIDGE/ VALENSTEIN 1991), did this type of conceptemerge into more open consideration in behavioralneuroscience (ROBINSON/BERRIDGE 1993). VALENSTEINhas long championed the massive role of learningand plasticity in the emergence of psychologicalfunctions in the brain (and has strongly arguedagainst subcortical specificity of functions), and webelieve his analysis of the psychopharmacology revolutionwas motivated to some extent by his philosophical/democraticpoint of view.Moreover, we are most certainly not dualists ofany stripe and Steven J. GOULD has not been ‘ourmentor’ (even though we admire his writing skillsand with the help of SEGERSTRÅLE’s (2000) analysis,better appreciate his humanistically motivated willingnessto present the other side of the argument inthe lucid, energetic and perhaps politically motivatedways that he has). Accepting the vast plasticityof the neocortex does not make us advocates of untenablevariants/hangovers of CARTESIAN thought.Our views are thoroughly naturalistic, monistic andmaterialistic (without abandoning our spiritual/humanisticcompass, which we believe is closely linkedto our deep animalian capacity to experience pro-Evolution and Cognition ❘ 67 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. Pankseppsocial emotions). We accept that the multilayeredcomplexity of the human mind cannot be understoodif we do not come to terms with our ancestralpast.We were dismayed that PITCHFORD purports thatwe do not subscribe to a hierarchical view of brainorganization, intimating instead we advocate a perplexing‘horizontal layers’ view of brain organization.Obviously, the brain is layered hierarchically,with higher functions critically dependent on variouslower functions (please refer to Figures 10.3 —10.5. of PANKSEPP 1998a), and the flexibility of thelower functions has been increased enormously bythe regulatory capacities of the higher functions. Inour estimation, it is the modularity view that seeksinformationally ‘encapsulated’ evolved units inbrain areas that do not have an adequate evolutionaryhistory for their construction, nor an anatomicalplace or reasonable neurodynamic way to beplugged into the rest of the brain.A great deal in evolutionary psychology will rideon the exact nature of the massive neocortical mushroomingsthat characterize the anthropoid/humanoidbrain. Whatever specializations exist in the neocortex,there is a good possibility that they are theones we share with other animals, and the rest islargely a quantitative expansion of these samethemes as opposed to major re-engineering of functions.Mere increases in computational space cancreate seemingly qualitative emergent effects. PITCH-FORD also suggests that “our brains may not simplyprovide an excess of plastic neurons capable of subservingany function, but may be a solution to theproblem of retaining adequate functioning over aprolonged period of time”. This is not an explanationof increasing hominid brain complexity at all.A relatively small, loosely-structured, ungyrified‘brain’ suits a forty pound lobster of approximatelyeighty years just fine! ‘Prolonged function over time’is as creative as any unsubstantiated Panglossian explanation,as it would be an advantageous propertyof nearly every adaptation. More significant toPITCHFORD’s argument is that we posit different andseemingly unrelated processes for the evolution ofsubcortical and cortical brain structures. This couldnot be any further from what we actually think. Naturalselection has had free roam in all areas of thebrain. However, it should be obvious to all that naturalselection has been molding ancient subcorticalbrain structures for a much longer span of time thanthose found in neocortical regions of the brain. Themore that something has been ‘selected’, the moreconsistently, efficiently, economically and reliably itwill serve its function. We can be quite certain aboutthe genetically ingrained nature of many subcorticalfunctions. We can have no comparable confidencefor neocortical ones, especially in humans whereearly development and cerebral maturation go temporally,hand in hand (DAWSON/FISHER 1994;THATCHER et al. 1996; PANKSEPP 2001a).Above all, we subscribe to the position that thefertility of ideas is linked inextricably to our abilityto predict and explain new phenomena, and we believethat from this perspective, neuroevolutionarypsychobiology far outshines Evolutionary Psychology.We are confident that PITCHFORD, were he tallyingup the substantive findings and claims, wouldsee the strength of our strategy and also our robustallegiance to a deep evolutionary view of human andanimal nature. In any event, while seeking out theweakest chinks in our evolutionary armor, he assiduouslyavoided the most important aspects of ourarguments concerning the core-nature of our emotionaland motivational processes. We hope that inthe long run, evolutionary psychology will not dothe same, even though in the short-run we do anticipatea ‘tit–for–tat’ strategy.In any event, it should be a ‘no brainer’ for allevolutionists that if we do not understand our ancestrallyderived attentional, emotional, motivationaland memorial capacities, we can never really understandour own nature. The most dangerous place toseek modularity is among the highly-specialized,neocortical functions of the adult human brain. Thismay be a bitter pill for most evolutionary psychologistswho seem to wish to force a unique humannature onto our species and hence, seem to have anaversion to both animal- and organically-focussedresearch. Their most robust general principles andphenomena (as well as potential allies) will be foundamong the subcortical functions of the brain (andpresumably among investigators who have studiedsuch functions). They simply need to drop their ‘socialscience, anti-biological tribalism’ and assimilatethe types of lessons that apparently are second natureto their colleagues in ASCAP and human/animalneuroethology societies.Scott Atran:The Language ‘Module’ Once MoreObviously, all neuoscientifically informed peopleagree that the human cortex permits levels ofabstraction that are not available to any other species.Human language is the most striking exampleof this ability, as it has attracted more experimentalEvolution and Cognition ❘ 68 ❘ 2001, Vol. 7, No. 1

A Continuing Critique of Evolutionary Psychologyand theoretical attention than any other faculty ofthe human mind. Clearly language does not merelyarise through SKINNERIAN associations, and we are asattracted to the view of a deep-structure for languageas much as anyone, but we agree with CHOM-SKY that it is premature to genetically map it ontoneocortical areas of the brain. Of all the non-trivial‘modules’ that have been proposed (i.e., thosebeside the straightforward sensory and motorcapacities of the brain), those devoted to languageare clearly among the most compelling.ATRAN provides a balanced overview and a vigorousdefense of modularity with language as the centerpieceof his argument. We agree with much of hisanalysis, but we still think that it is wiser to hold offon any strong, genetically-guided modularity view,since the arguments against a uniquely human geneticguidelines for language remain quite compelling.We think many other organisms could also exhibitlanguage if they too had a much more complexneural space for general-purpose, perceptual/cognitiveprocessing within their neocortices and more sophisticatedvoiceboxes. Yes, we agree that languagehas become a very adaptive tool for the human species,but so have many of the other emergent propertiesthat culture permits. We think that all may agreethat reading and writing are not derived from ourgenetic heritage, and it is a pity that some of us havecaved in so rapidly to the view that our vocal expressionsof language, at least in their basic outlines, areconstructed in such a manner. We certainly agreethat many spandrels can emerge from a vast, predominantlygeneral-purpose learning organ (i.e., theneocortex) which ultimately (i.e., epigenetically) willcontain functionally distinct neural circuits.The most critical issue in our minds is whether anewborn baby has the intrinsic neural circuitry forlanguage or only the intrinsic proclivity to babblewhile it perceives the world, points to the world,grasps objects in the world, manipulates the worldin a self-referential way (all very primitive mammalianabilities) and finally begins to associate theseongoing vocalizations to certain activities in waysthat gradually grow into the vast complexities of humanlanguage. The only issue that we felt we hadraised with this example of modularity was whetherour genes contain the rudiments of a uniquely humanlanguage, or whether they merely help constructa well-placed voice box and resonance chamberfor making complex sounds that can interactwith the enormous RAM-space in the cortex, withlanguage as its final product. ATRAN provides a vigorousdefense of a sophisticated variant of the acceptedwisdom and we chose, largely for didactic purposes,to take the contrary position that is certainly not falsifiedby evidence thus far. Our main goal was to highlightthe simple fact that in our evolutionary thinking,we should not stray from the availableneurodevelopmental evidence to conjectures aboutthe potential evolutionary sources of the final producttoo quickly. This is a primary reason why we supportthe comparative study of subcortical processesso ardently, for there is much more assurance of havingrelevance to our neuro-genetic understanding ofhuman behavior.We should all be more willing to consider that agreat deal of communicative intent and its associatedpragmatics may arise from ancient processes weshare with other animals. We believe that humanswould have no lingusitic competence without thebrain systems that exist below the neocortex and ifone can appreciate this point, one may also understandwhy the human neocortex may not require agenetically dictated language ‘module’ to have communicativecompetence (e.g., BROCA’S and WER-NICKE’s Areas may emerge developmentally becauseof other compatible multi-modal functions of thesetissues). Moreover, we hazard the prediction thatonce the neocortical circuits for language are completelyelucidated, it will be impossible to simulatelinguistic competence in a program that instantiatesonly those higher cerebral processes. In other words,the likely result of this thought experiment would bethat by itself, the unique human neocortical architecturecould not generate, certainly not motivate, asubstantially meaningful human language. Indeed,we note that at least a third of classic KANNER-typeautistic children are incapable of speech throughouttheir lives, although they do not typically have anyhigher cerebral dysfunctions—it is simply that theirhigher brain areas are not well connected to subcorticalsystems (BAUMAN/KEMPER 1995).Let us be frank about our position: Just as we believethere are many subcortical faculties that are absolutelyessential for the types of psychological processeswhich ATRAN discusses, we also believe thatthere are bound to be some cortical faculties of considerableimportance for the emergence of humanlanguage. In describing the cortex as a general-purposetype of ‘information processor’, we are notcommitted to the view that it is nothing but a generalpurpose associative tool. Cortical processing isguided by many subcortical processes, and indeedwe have previously envisioned that there are cortical‘spheres of influence’ for higher-order processing ofinformation relevant to various emotional and mo-Evolution and Cognition ❘ 69 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. Panksepptivational survival concerns (PANKSEPP 1988, 1989).We anticipate that those ‘spheres of influence’ mayprocess information in unique ways. We also believethat the frontal lobes have some very general abilitiesthat give it such a unique role in elaboratingvarious executive processes (GALLESE/GOLDMAN1998; BARKLEY 2001). Indeed, many such executiveprocesses probably participate in so many differentcognitive functions that the concept of modularitymerely becomes a hinderance in trying to work outhow some of the basic cortical processes that helpconstruct so many sophisticated cognitive abilitiesvia the experiences of individuals.Thus, we are not adverse to studying such higherfaculties, but we suggest that evolutionary viewshave not and are not likely to have any impact onhow we empirically study these functions, althoughthey can help loosen up the types of intellectual constipationthat have characterized past variants ofwhat might be appropriately called ‘dustbowl cognitivism’(see BARKLEY 2001 for an example of this laxativeexperience). In any event, our position was thatultimately, most of the sophistication of the corticalfunctions is brought to life experientially. An understandingof the neuro-maturational processes thatyield well-honed cortical processes will be much,much more, dependent on general brain mechanismsthat govern neuronal plasticity than any typeof resolved information that is contained in thegenes. That was our main point, and we encounterednothing in ATRAN’s well-crafted defense that dissuadesus from this position.We see our disagreement amounting to littlemore than a matter of emphasis. ATRAN wishes toretain the concept of modularity and sees some typeof implicit advantage in that evolutionary view, althoughhe does not suggest how it might be cashedout empirically. We see both of those issues to besuperfluous for the pursuit of his faculty-psychologyprogram. We regret that he fails to see the potentialpower of subcortical, socio–emotional systems ingoverning many of the processes in which he is interested.Thus, to take his first example of a structuralprinciple, ‘the whole–object constraint’, theunderstanding of which modularity has presumablyfacilitated, we point out that the visual systemis certainly designed during early development toperceive the objects of the world (this is also evidentin other animals). In our estimation, young childrenattach labels to such objects using social, ‘point andname’ games with caretakers. To underestimate basicsocial/motivational/emotional processes in suchlinguistic acquisitions is, we think, shortsighted.Considering that ATRAN dealt with only one smallsegment of our critique, we were surprised that heclaimed that “there is no empirical evidence whateverto indicate” that our “central thesis is true, andmuch to suggest that it is false”. If the ‘seven solutions’we shared are of relevance for human behavior(admittedly most remain to be tested in humans),then a cognitive psychology that continues to neglectsuch issues must surely be deemed nothingshort of myopic. We think that the failure of mostcommentators to deal with the animal data we providedsimply highlights that a great deal of the evolutionaryprocesses that control our behavior appearto be of little interest to evolutionary and cognitivepsychologists. We suspect that many of those processesare absolutely essential for the higher processesin which cognitive scientists are so interestedand we will only note the disastrous effects that damageto those subcortical systems can have on humanmental life (DAMASIO 1999; SCHIFF/PLUM 1999).As we stated in the target article, while socializationis certainly not a direct evolutionary explanationof human language (as ATRAN also emphasizes),it probably was a selective force for the emergence ofcommunicative intent. We assume that all will agreethat relatively little human language would emergewithout socialization. We can also agree, that inmost people when they have learned language, agreat deal of the critical circuitry is concentrated inWERNICKE’s and BROCA’s areas of the left hemisphere,as well as the circuits of the anterior cingulate. Moreover,if all those tissues are destroyed, the child willstill learn language by using, presumably, the correspondingsystems of the right hemisphere (BATRO2000 VARGHA-KHADEM et al. 1997). However, the plotnow thickens. Children born deaf also aspire to communicateand they do quite successfully, partly byusing somewhat different parts of the brain wheresomatosensory skills are usually elaborated (NEVILLEet al. 1998; PASQUAL-LEONE/TORRES 1993). And wheredo multiply impaired individuals like Hellen KELLERelaborate their urge to communicate?We, along with others (LIEBERMAN 2000), believethat so much of language is linked to subcortical andcortico–limbic social urges/pragmatics, that it may bethat language emerged from these ingrained systemsinteracting with our massively sophisticated RAMtypespace of the neocortex. The reason the left hemispherehas such primacy in most people is becausethe left side of the brain is more talented at extrovertedsocial activities (for reasons we will not entertainhere), and because WERNICKE’s area is nestled betweenall of the major perceptual surfaces of the brainEvolution and Cognition ❘ 70 ❘ 2001, Vol. 7, No. 1

A Continuing Critique of Evolutionary Psychologythat harvest exteroceptive information (i.e. vision,hearing and touch). The ultimate function of languageis to symbolize the world in abstractions andto allow us to develop better internal thought for thepursuit of our many basic goals and externalizedstrategies for coordinating group activities.Thus, we are able to buffer information about ourrole in the world in much more complex ways thanany other species, but where is the intrinsic, evolutionarily-selected‘module’? How could genetic selectionhave created such dispositions uniquely forthe human mind/brain? Where are the genes for language?Might all the pragmatics have already beenthere before humanoids walked the face of the earth?Might the main thing we needed to cross the syntactic-culturalRubicon have been a massive endowmentof more RAM-type space in the right parts ofthe brain working within the context of the mountingmedium of cultural evolution?In saying all of this, we must again emphasize thatwe certainly agree that there are many specializedfunctions in the adult human cortex. In our originalarticle, we did not aspire to suggest that there werenone. Not for a moment do we believe that higherfaculties are constructed merely from associativenetworks. We think that the cortex has many specialways to construct meaning (BARKLEY 2001). There areintrinsic mirroring processes (RIZZOLATTI/FADIGA/GALLESE/FOGASSI 1996), which may serve as a majorvector in the acquisition of language (GALLESE/GOLD-MAN 1998). However, we suggest, along with someothers (LIEBERMAN 2000), that the cortex could donone of that—simply none of that—without thesupport of subcortical systems (and that seems to bea novel perspective for many). We remain perplexedwhy so many scholars are willing to commit themselvesto neurologically unrealistic images of humannature, such as the neurobiological and corticalmodularities of evolutionary psychology, which arenot yet supported by essentially any sociobiologicaldata. Our main goal was to interject a much greaternote of necessary doubt into the intellectual activitiesof recent variants of Evolutionary Psychology.We disagree deeply with the ‘separatist’ cortico–cognitivist position advanced by ATRAN. The importanceof subcortical and limbic systems in humanaffairs has been massively underestimated. When hesays “nothing from work on subcortical emotions…has provided, or hints at providing, the slightest insightinto the cognitive structures and processes discoveredso far”, we suggest this view simply reflectsa dismissive mode of inquiry preferred by many intellectualsinterested in the higher human proclivitiesand abilities. The subcortical perspective willprovide a great deal once individuals begin to studywhy human minds dwell obsessively on certainthemes when they are hungry, thirsty, cold, sexuallyeager, craving drugs and worrying about their socialattachment concerns. There are bound to be higher‘centers of gravity’ for the cognitive elaboration ofsuch basic functions (PANKSEPP 1988, 1989), perhapsepigenticially magnified or diminished dependingon early experiences (PANKSEPP 2001a). The wholefield of biological psychiatry has been based largelyon manipulating subcortical systems, many ofwhich directly modulate cortical functions. Therewill be a vast new biological–psychiatry revolutionof even greater magnitude when we learn to manipulateother peptide systems in the subcorticalreaches of the brain, especially those that controlspecific emotional and motivational processes(PANKSEPP 1993; 1998a). These molecules will take usto the appetitive core of our existence. We will derivewhole new layers of deep understanding about theemotional imbalances that are called psychopathologies.Animal models will help us work out many ofthe essential neural details of such imbalances, andthereby the sources of our everyday likes and dislikesthat continue to be among the most robust sourcesof personality variability, as well as the basis of somany of our human interactions.There is an ‘affect logic’ to our cognitive activitiesthat has barely been studied empirically (SHAND1920; WIMMER/CIOMPI 1995). We intentionally leftout one phrase (i.e., ‘or neocortical principles of generalassociation’) in the above quote because we arenot as committed to a fully associative mode of corticalprocessing as some of our commentatorswished to impute to us. One can have general-purposelearning functions in the brain which are notsimply based on association-network heuristics(FREEMAN 1999). Aside from a few well-analyzed sensory/perceptualand motor processes, our knowledgeof how the ‘great intermediate net’ of heteromodalcortical tissue operates remains a great mystery. Weare tempted to add that essentially nothing from workof Evolutionary Psychology has provided or hints at providing,lasting insight into the cortico–cognitive structuresand processes that truly exist in the human brain.Still, the evolutionary view can certainly help generatesuch hypotheses (e.g., BARKLEY 2001). If we coulddrop our destructive species-centrism (which nocommentator denied being a problem), we can learnan enormous amount about how the human mind/brain operates (DE WAAL 2001; GALLISTEL/GIBBON2001; PANKSEPP 1998a).Evolution and Cognition ❘ 71 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. PankseppLet’s Admit It: We Don’t Know SquatAbout the Evolution of Language!We know about three definite things about the evolutionof language: (i) The voice box and rest of theupper respiratory apparatus appear to have undergoneevolutionary shaping such that we can make aremarkable variety of resonant sounds. Of course,other creatures, songbirds for instance, haveachieved similar excellence in sound variety withoutadvancing much toward human-type languageor human-type neocortex. (ii) The areas of the lefthemisphere that are especially important for language(e.g., planum temporale) are already enlargedat birth and palentological evidences indicates theseorgans enlarged quite a long time ago—namely,when our brains started to expand beyond the proportionsof other great apes some two million yearsago (for a summary of that work, see LIEBERMAN2000). (iii) We can’t recall this one right now, but itdoes not matter much, for if that is essentially allthere is to our knowledge of the evolution of language,aside from our widespread readiness to beawed by this uniquely human skill, the evolutionaryevidential base must be deemed slim indeed.Of course, we are all very impressed that in theabsence of brain damage or major sensory deficits(e.g., deafness) language is biased to settle in the lefthemisphere in circuits known as BROCA’s and WER-NICKE’S areas. Indeed, people with a college educationtend have more robust neurons (i.e., larger dendriticfields) in some of these regions compared tohigh school drop outs (JACOBS/SCHALL 1993; JACOBS/SCHALL/SCHEIBEL 1993), and there are many otherbrain/language correlates (GRABOWSKI/DAMASIO2000). However, we should hold off on surmisingthat such language functions reflect biologicallyevolvedskills that have any pre-determined and detailedcognitive resolution. After all, these are alsoamong the most impressive pharyngeal-motor andexteroceptive-associative areas, and it seems quitereasonable for language, which allows us to re-symbolizethe experience of the individual senses, to finda natural epigenetic home in such areas. Many maybe tempted to localize some fine evolutionary ‘needlework’for the deep structure of language here, butno one is quite sure what one might look for in thoselocations—anatomically, electrophysiologically orneurochemically—that might yield data consistentwith an evolutionary scenario.Parenthetically, and to reply to ATRAN’s most daringattempt to dispatch out critique (i.e., that thereis no evidence for the role of subcortical functions incognition), let us simply assert that we have no reasonfor a comparable agnosticism when it comes tosubcortical functions. There is an abundance of anatomicaland neurochemical data for a host of functionsthat speak clearly for strong and powerful homologiesacross all mammalian species (PANKSEPP1998a). We can also be certain that these systemshave great influences on our cognitive processes:when we have strong emotions and motivations inour minds, related topics are essentially all that weare prone to speak or think about. It is an intellectualtragedy that the cognitive sciences have marginalizedthese incredibly important aspects of our mentallives. We suspect many loose pieces in cognitivepuzzles would fall together well if we would simplyfocus on the affective sides of human lives a bit more.But speaking is only one form of communicatingand we know that when the most robust neural locationsfor language processing are damaged, manypeople are able to communicate quite well again,using different, albeit nearby, regions of the cortex.Sign language seems to have a natural tendency tonestle near the classic left-hemisphere speech areas,but reading is vastly different in deaf individuals(NEVILLE et al. 1998). When people have strokes, languageand other functions tend to be restored quitewell by unaffected tissues that surround the damagedarea by becoming more active during communicatoryactions and intents (BUCHNER et al. 1996;NUDO et al. 1996), and language acquisition certainlyexercises the brain in most interesting ways(CASTRO-CALDAS et al. 1998). In short, communicativecompetence can emerge from many areas of thehuman cortex and all of it requires a variety of subcorticalfunctions about which we know very little(GRABOWSKI/DAMASIO 2000; LIEBERMAN 2000). On thebasis of abundant neurological data, there appears tobe a major role for subcortical structures in such processes(BASSO/SALA/FARABOLA 1987; CROSSON et al.1997; BECHTEREVA/ABDULLAEVA 2000). Perhaps this isthe third finding we were floundering for earlier, ormaybe it was the very important role of prosody(e.g., motherese) in promoting language developmentin children (FIFER/MOON 1994). But it does notmatter, for the evidence supporting evolved cognitiveprocesses of language is tenuous and in the absenceof specific genetic details, it is unlikely to get muchstronger.We could carry on further with these lines ofthought, but the key point has now been made a fewtoo many times. So let us, ever so briefly, indulge ourown evolutionary fancies. The multi-modal associationtissue of the cortex that is essential for linguisticEvolution and Cognition ❘ 72 ❘ 2001, Vol. 7, No. 1

A Continuing Critique of Evolutionary Psychologysymbolization will work more effectively if there area greater number of neurons that can ‘crystallize’their tasks efficiently. Thus, we suspect that the evolutionaryadvantage may have been for individualsthat had more tissue in those brain regions, fastercommunicating neurons and perhaps the many factorsthat contribute to an exquisite plasticity thatcan become readily ‘gelled’. And perhaps little more,except for the fact that when these systems becomedisconnected from the deeper feelings of the righthemisphere, they tend to confabulate like crazy(PANKSEPP 2000c). It is most remarkable that psychologicalprocedures, such as psychoanalysis, can getbehind the confabulatory facade to deeper principlesof reality (KAPLAN-SOLMS/SOLMS 2000). If oneputs our whole vision of human linguistic naturetogether, it may help explain why most of us tend tobe better speakers than listeners, and why so manyof us are eager to speak faster and louder than thenext fellow, until there is little more than a blisteringcacophony of sounds.… “Till human voices wake usand we drown.” (T. S. ELIOT, The Love Song of J. AlfredPfrufrock).Parismony and ModularityThe term modularity, at least as defined in the dictionary,provides no realistic perspective on the realcomplexities of the brain. It is a term that emergedfrom the field of engineering, where it was convenientto design complex circuits so they could berepaired by exchanging large components, asopposed to tracking down problems in the detailedconnections of spaghetti-like electronic circuits.Contrary to what PITCHFORD suggests, the term wasnever common in neurology, even though recentlywe do find even some of our respected neurosciencecolleagues falling into that sloppy usage ever morefrequently, now that it has become a meme of massiveproportions in so many academic circles. Noone wants to be left out of such discussions and alltoo many have come to casually use the term, eventhough neuroscience disciplines have had an abundanceof apt brain terms (e.g., systems, networks orcircuits), long before the modularity idea was introducedto us by individuals who had little directexperience with brain research. This ‘digibit-module’metaphor (we assume some programmers stillfondly remember those useful electronic units prevalentin the 1970s) offers no promising empiricalpayoffs. Modularity simply adds nothing useful toexisting neuroscience strategies or agendas, exceptperhaps a false sense of security that we have identifieda natural kind in the mind/brain without havingdone any neuroscience. Is it too much to ask all evolutionarypsychologists to simply drop the modularityconcept? Nothing would be lost and a lotcould be gained, as was evident in one recent contribution(DUCHAINE/COSMIDES/TOOBY 2001). If weneed such a concept, why not simply use a classicdesignator such as faculties, which simply helpsunderscore key problems that needed scientificattention.We submit that if we do not understand the ancestralminds that are still represented in the manyspecies with which we share the earth, we will notreally understand our own. Now that we understandthe massive genetic relatedness among all mammals,we must start to fathom how the basic brain processesthat all mammals share guide our humanmental activities. We suspect that when we begin toaccept the existence of certain primitive attentional,emotional and motivational faculties in humanmind/brains, we will find some parsimonious waysto solve many of the fascinating problems that evolutionistshave brought to the forefront of the psychologicalagenda. Let us simply share a few examples:As highlighted at the end of our target article, webelieve that human rape emerges substantially frompoorly socialized or sociopathic mind/brains thatare inclined to harm others for their own primitivesexual gratification. It will be an important neuroscienceproject to determine to what degree basicaggression systems in the human brain overlap withthose that mediate sexual urges. We already knowthat in certain species the overlap can be substantial(KOLLACK-WALKER/NEWMAN 1995).As evolutionary psychologists have compellinglyshown, males seek beauty and youth in their socio–sexual partners, while females seek signs of devotionand material investment. This may largely be due tothe fact that the primitive systems for male eroticurges are driven more by simple exteroceptive cuesof nubility, not much different than those of otheranimals. Is there really much reason to propose anyintrinsic cognitive, decision-making circuitry? Mightfemales simply have the socio–neurological skills torespond to primitive social rewards more readilythan males—the pleasures of being together, eatingtogether, sleeping together and being tolerant ofeach other? These are all primitive affective featuresof the mammalian condition that do not require sophisticatedspecial-purpose, cognitive ‘modules’.This is not to suggest that the origin of such choicesare not based on reproductive issues, but simply toEvolution and Cognition ❘ 73 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. Pankseppassert that simple emotional systems with a modicumof some general-purpose cognitive skill, mayeasily yield some of the most striking folk-psychologicaldiscoveries of evolutionary psychology.Yes, childless, rouge males are more likely to harmthe young whereas fathers and especially mothersare much less likely to do so, especially to their own(DALY/WILSON 1988). Could this simply reflect theprimitive social-bonding mechanisms of the mammalianbrain that promote anti-aggressive attitudesby arousing ‘peace-promoting’ neurochemical activitiesin ancient neural systems? One of the mostspectacular findings in this area is the discovery thata single sexual episode can gradually transform infanticidalmales into tolerant ones, so that they areleast likely to kill young at the time their own offspringenter the world (MENNELLA/MOLTZ 1988). Weanticipate that when investigators probe such matters,they will find that satisfying sexual activitiesarouse the chemistries for nurturant feelings in thehuman brain, just as they do in other animals(CARTER/LEDERHENDLER/KIRKPATRICK 1997).We have no problems with the idea that evolutionarily-derived,socio–sexual adaptations existwithin our mind/brain dynamics, we simply questionthe wisdom of conceptualizing these processesas evolved, cortico–cognitive ‘modules’. At the sametime, we do agree with carefully analyzing how humancognitive activities change as a function of suchaffective changes (which will eventually be capableof being precisely manipulated in humans viaknowledge derived from the study of the corresponding,emotion-modulating chemistries in animalmodels). There are other interesting emotionalexamples to consider from the vantage of the principleof a cross-species parsimony (PANKSEPP 1998a).We certainly believe that such ancient emotional dynamicsare bound to have remarkably strong effectson the cognitive dynamics of human minds. However,the study of such processes will require developmentof a more vigorous, empirically deep psychology(PANKSEPP 1999; SOLMS/NERSESSIAN 1999).Group SelectionIn the initial paper, we chose to leave a variety of ourarguments open-ended, so we could attract pointedcriticism about issues about which we all truly knowvery little. The most poignant example was our willingnessto consider a viewpoint that is viewed as ananathema by most evolutionists. Namely, the conceptof group selection. In truth, we are quite agnosticon this issue and simply urge our colleagues to bemuch more open-minded on the topic than manyhave typically been. First, let us be unambiguous oncertain issues. We certainly agree that evolutionprogresses as a function of the survival of gene patterns.The functional unit of selection must be theindividual that carries genes and at least in ‘higher’animals, that negotiates (typically emotionally)with other members of a species to pass on its genes.In such interactions, affect may be the main currencyof decision making and behavioral choice.Thus, it would seem essential to consider the role ofemotions, especially those emerging from dyadicinteractions, as a fundamental source of evolutionaryprogression in our species and many others. Inshort, our introduction of a variant of the groupselection concept is only tangentially related to theclassic concept.In its historical meaning, group selection wasposed and modeled as a possible explanation of whyindividuals in social groupings may behave in a waythat is overtly suggestive of altruism. WYNN-ED-WARDS (1962) postulated that individuals may concedetheir own opportunity to pass on their owngenetic guideline, by opting to contribute time andresources to other members of the group to which itbelonged—a contribution that would facilitatepropagation of the species. Soon after, HAMILTON(1964) argued that seemingly altruistic acts could beexplained by the concept of inclusive fitness. In thisway, individuals would only help out a group memberto the degree that the cost of a specific ‘generous’act was less than the genetic passage to be gained(i.e., the degree of genetic-relatedness should be apredictor of the extent to which an individual willbe altruistic). From this vantage, altruism could beexplained in terms of the genetic success of individuals,and the potential benefits of reciprocity.What remains a moot and often an acrimoniouspoint is whether gregarious animals act altruisticallywhen there is negligible genetic gain at stake. In ourprevious paper, we opined that certain affective/behavioralfunctions could only propagate within aweb of group dynamics. Indeed, if these processesonly occur when individuals are genetically related,they certainly can be reduced to selection at the levelof the individual. However, if some can also operatestrongly between genetically-unrelated individuals(e.g., arising perhaps from pair bonding and othersocial coalitions) then one can imagine, for instance,dyadic selection (a minimalist form of group selection)to be an influential force in evolution. Eitherway, we feel that this process is substantially differentand thus deserving of its own distinction whenEvolution and Cognition ❘ 74 ❘ 2001, Vol. 7, No. 1

A Continuing Critique of Evolutionary Psychologythinking about mechanisms of selection. Henceforth,we will refer to selective processes that operateon neuro/behavioral systems that only function successfullyin such groups as social selection (not to beconfused with group selection although there is thepossibility of linkages).It is clear that the study of intrinsic social processeswas massively marginalized during the previouscentury. Anyone that has spent time studyingsuch matters should recognize that many socio-affectiveprocesses can only operate within groups.Normally, one cannot tickle themselves or sociallyplay alone, and one can only deeply grieve for theloss of another, but not as much for material losses.A preliminary characterization of the underlyingbrain systems has been achieved for some of theseneural processes (PANKSEPP 1998a), but there hasbeen massive resistance in psychological and neurosciencecommunities for pursuing such topics of ultimateconcern. Many still assert that specific socialaffects do not exist—that they are all learned or sociallyconstructed. This is most certainly wrong,even though all the details of our social existence aresurely learned. And when we have the conceptualliberties that the human-type cortex allows, we canobviously choose to go against the dictates of thesesystems. We are perhaps the only species that has theoption of being libertarians.Our advocacy of the social selection hypothesiswas advanced largely to help open scientific inquiryinto the many unstudied mysteries that exist in allmammalian brains. We should not close our mindsto reasonable possibilities because of the appealingrigor of certain mathematical equations. Whether insome instances social selection and group selectionexert a synergistic pressure on certain brain functionsremains an intriguing possibility. While it continuesto be only a possibility for other species, wefeel that it is in fact very plausible that group selectionhas already occurred in our own species. However,the important distinction is that such selectiveprocesses may have been driven primarily by culturalevolution and not by biological evolution (justconsider the caste systems in some cultures, not tomention the holocausts that have been inflicted onothers). It seems intuitive that gene frequencies andthus collective genomic phenotypes can be alteredby seemingly capricious cultural choices. The potentialgrouping of genes in human beings will be asunpredictable as their new selective force—culture.There should be no reason why this cannot be accepted,without moral exacerbation, as an emergentproperty of the interactions between dedicated brainoperating systems, flexibly non-dedicated systems(where emergent properties can also be exquisitelyhoused), environment and culture. It is within suchrealms where the concept of developmental programswill become most robustly manifest. Cortico–cultural evolution has obviously provided a new andall too often terrible order to the evolved dynamicsof the mind/brain (PETERSON 1999), and of course,the genes have also played their part. Fortunately, westill have the hope of better educating our highercerebral systems.Behavior Genetics—An Essential KeyIt is understandable that evolutionary psychologistsdid not want to get entangled in the genetic determinismthat has historically colored sociobiologicalthought. However, there is no intellectually-soundreason to be coy about this topic. Genes are obviouslyoverwhelmingly important in the way themind/brain is constructed as is epigenetic development.To think otherwise is to indulge in politicizednonsense.The robust heritability of a majority of humanpersonality traits is stunning, while there is very littleheritability for the specific things we do in ourenvironments. For years, BOUCHARD (1998) has beenfinding clear evidence for ‘nature’ in the constructionof human personality, while the role for ‘nurture’has gotten slimmer and slimmer (almost nonexistentby some of his most recent ‘adoptees raisedin the same environment’ analyses). Namely, identicaltwins reared apart show the same reasonablyhigh degree of heritability on practically all traits asdo twins reared together (r’s typically between .4 and.6), while two unrelated adoptees raised in the samefamily exhibit practically zero correlation on suchtraits (BOUCHARD personal communication withJ.P.). Also, it is becoming clear that our genetic dispositionshelp us actively create social and materialenvironments that are satisfying or less distressing.Yet, no one knows what has really been selected forat the genetic level, but we suggest that at a systemslevel, it is the vigor of certain basic motivational,emotional and attentional functions. We will notdwell on these important issues since they have beencovered so well elsewhere (BOUCHARD 1998), but willshare one striking recent example:BOUCHARD’s most recent work on religiosity wasinitially a bit of a mess until he broke it down intointrinsic and extrinsic religiosity. The intrinsic variety(e.g., spirituality?) had a high genetic loading,while the extrinsic variety (e.g., church attendanceEvolution and Cognition ❘ 75 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. Pankseppand other external practices) exhibited a high environmentalloading. The two forms of religiousnesswere inversely correlated—probably not surprisingfor many evolutionists who understand the utility ofdeception and the probable importance of corticalprocesses in generating such tendencies. Also, intrinsicreligiosity was positively related to authoritarianism(BOUCHARD et al. 1999). A liberal collegeeducation had a small effect on dampening that oldtime prophet instinct (i.e., a temporary nurture effect),but at eight years past those liberating collegedays, the old ways were—hang on Evolutionary Psychologyfans—coming back with a vengeance, especiallyif you had lots of kids (personal communicationwith J.P.). Once again kids force us back to ourgenetic roots. Clearly, practically any human or animaltrait can be amplified or diminished genetically.However, in the past we rarely knew what was reallyselected, but the new molecular biology will soonchange that.Politics and TribalismAn enormous amount of ink and mental energy waswasted during the sociobiology wars because of thepolitical inclinations of certain participants (SEGER-STRÅLE 2000). We cannot easily escape our politicalbiases, but intellectual and social disasters can easilybe made if we do not own up to our ‘mere beliefs’.For this reason, we especially appreciate SEGER-STRÅLE’s participation in these discussions. Forinstance, the harsh treatment of the role of genes inour destinies at the hands of politically motivatedMARXISTS is a story that presently has only been halftold (hopefully, some forthcoming memoirs will setthe records straighter). We are the kind of speciesthat can perform intellectual, emotional and bodilymonstrosities in behalf of what we deeply believe(PETERSON 1999). All too easily, we develop tribalaffiliations which color all our subsequent beliefsand arguments. None of us are immune to suchissues, but they are often very apparent when wecome to issues as contentious as sociobiology andEvolutionary Psychology (LEWONTIN 1993).Probably most who read journals such as this arewell sensitized (or habituated) to those aspects ofthe human spirit, but the present debate has certainlynot been absent of those features. Let us be sobold as to highlight a striking instance that we couldnot avoid noticing. Our harshest critic in these discussions,PITCHFORD, asserts that “Neuroevolutionarypsychobiology’s reassignment of the tabula rasato the neocortex revivifies CARTESIAN dualism andrepresents a desperate strategy likely to be extremelydeleterious to scientific investigation. Evolutionarypsychology should reject this approach…”To some extent, this assertion is notsimply a cold-headed response to the strengths andweaknesses of existing data and arguments, but alsoan indication of tribal affiliation, since he is a ‘listmaster’ for the extended evolutionary psychologycommunity. In personal communications, PITCH-FORD was kind enough to say, upon first reading ourarticle, that “although I disagree with a few of thedetails I do agree with broad thrust of the piece. Inmy view EP has made a number of fundamentalerrors: (1) by adopting the method of reverse engineeringin place of a consideration of phylogeny (2)by adopting functionalism, or the study of substrateneutral processes, in preference to the study of neurobehaviouraland neuropsychological homologies,(3) by adopting a commitment to the notionof the monomorphic mind, thereby excluding aconsideration of both polyphenism and adaptivegenetic variation as studied by game theorists, and(4) by adopting genic selectionism in preference todevelopmental systems theory. I think the issue ofdomain-specific versus domain-general mechanismsis of less importance than these four mainissues, as the brain surely contains both, and theirrelative contributions to our cognitive–emotionalcapacities is an empirical matter” (personal communicationwith J.P., 1/3/01).Then, four months later, after one of us (J.P.) hadshared some surprise about the throttling he hadprovided, PITCHFORD wrote back that “In my viewyour perspective embraces almost every issue of importance.The commentary does concentrate on areasof disagreement, but had the word limit allowedI would have had much more to say about affectiveneuroscience as the essential foundation for evolutionarypsychology. The emphasis on functionalismthat evolutionary psychology has adopted from cognitivescience can result in a neglect of cross-speciescomparisons, and in a reluctance to embrace importantbranches of evolutionary biology such as ecologyand game theory. This can make it seem abiologicalat times, but I see this as an unwelcome tendencyrather than a defining trait. However, most of thefaults you identify in ‘seven sins’ can be found in thework of evolutionary psychologists” (personal communicationwith J.P., 4/23/01). We do not share suchpersonal communications with any desire to embarrass(indeed we agree completely with the gist ofthose arguments), but they do highlight that in ourhuman (male?) eagerness to put forward strongEvolution and Cognition ❘ 76 ❘ 2001, Vol. 7, No. 1

A Continuing Critique of Evolutionary Psychologyviews, we often veer far from the middle road. We alltend to do that. We are all prisoners of our times andtypically of intellectual tribes that have strong expectationsconcerning proper behaviors of theirmembers. And all too often we are also captives ofour political biases and our remarkable left-hemispherelanguage skills, which all too often comesclose to mere confabulation rather than reasonableargumentation (for perhaps too creative a critique ofthis human dilemma, see PANKSEPP 2000).And so what are our own politics? (1) we are committedto accepting our animal nature and at timeswe may veer rhetorically a bit too far in the zoomorphicdirection; (2) we are committed to accepting theaffective complexity (and cognitive also) of other animalsand we may often sound a bit too anthropomorphicbecause we do believe that it is fundamentallycorrect that other mammals have many basicemotional feelings not all that different than ourown; (3) we have a commitment to a deep sense ofhuman freedom and plasticity, and we may oftenclaim that humans have more cortico–cognitivechoices than they, in fact, do (but we should all hopethat they might); and (4) we have a commitment tohuman justice and paradoxically, that may lead ourprose to be a bit more heated than it should be. Wetruly regret if we were unfair to any of our commentatorsand we are sorry if some of their importantissues were not addressed as fully as they could havebeen.The Sin of AnthropocentrismIn closing, let us simply reiterate our thanks to allthe commentators for participating in this discussionand refresh our call to the many prominent figuresin the field who, so far, have chosen to remainsilent on these important issues. 1 We are personallycommitted to the importance of evolutionaryviews for understanding our deep nature. We donot believe that it can be done without a full confrontationwith the emotional tendencies of ourfellow animals. We thank evolutionary psychologistsfor opening up the debateabout human nature once more,and we encourage them to helpcorrect the deep flaw in our educationalsystems and societalinstitutions that continue to marginalizethe essential understandingof the basic underlyingissues—concepts that must bederived from our human understandingbut detailed knowledge of which can typicallyonly be derived from well-conducted researchon and with our fellow animals. We were quitepleased that GARDNER has found our ‘rat laughter’research to be intriguing, and that story line continuesto yield interesting findings (BURGDORF/PANKSEPP 2001; BURGDORF et al. 2001).Our recognition of human potentials will bemuch more profound when we realize how our general-purposeneocortical expansions have enabledour own capacity to elaborate on the geneticallyingrainedsolutions that evolution so ‘kindly’ provided.Clearly, our expansive neocortex has permittedthe emergence of an unprecedented culturalevolution. This neuro-evolutionary passageemerged too rapidly, or so it seems to us, for anyhighly-resolved, sociobiologically-engraved strategiesto have emerged within the expanding neocorticaltissues. Evolutionary psychologists should beginto entertain the full complexity of our animaliannature. With the recent emergence of more balancedviews (e.g., BADCOCK 2000), the future of thefield seems bright, especially if it can rectify and freeitself from the many fledgling mistakes it has alreadymade.In closing, we also note that many of the issuesthat we raised in our target article were concurrentlybeing addressed by one of the fathers of modularityand cognitive science. We are delighted thatJerry FODOR recently chose to provide a frank analysisof what has or has not been achieved by thecognitive and evolutionary psychology perspectives:“In fact, what our cognitive science has doneso far is mostly to some throw [sic?] light on howmuch dark there is. So far, what our cognitive sciencehas found out about the mind is mostly thatwe don’t know how it works” (FODOR 2000, p100).We suggest that this is largely because evolutionarypsychology has devoted so little thought to understandingthe basic neuro-emotional and motivationalprocesses upon which both human and animalminds are built. This neglect has, we suspect,been largely due to the dualistic, anti-organic biasin psychology and philosophy,as well as themore recent siren-songAuthors’ addressJaak Panksepp and Jules B. Panksepp,J.P. Scott Center for Neuroscience, Mind &Behavior, Departments of Psychology andBiological Sciences, Bowling Green StateUniversity. Emails: jpankse@bgnet.bgsu.eduor julesp@caspar.bgsu.eduattractions of computersand computationalism.To continue FODOR’s lament(p39): “you mightthink that cognitive scientistswould be worrieda lot about the limitationsEvolution and Cognition ❘ 77 ❘ 2001, Vol. 7, No. 1

Jaak Panksepp and Jules B. Pankseppof the Classical computational theory of mind.Speaking for myself, I’m worried half to death. Infact, it seems to me, much of the field is in deepdenial; a condition to the prevalence of the pervasivegood cheer of books like PINKER’s and PLOTKIN’soffers striking testimony. As usual, the characteristicmechanism of denial is suppression.” We expectto receive no less for the critique we have provided,but do, of course, desire more. In any event, we wishthe field well.Notes1 We thank all commentators for sharing their perspectivesand trust that the thinking of some of the remaining seveninvestigators who were invited to comment on our ideashave been influenced as well (i.e., David BUSS, LedaCOSMIDES, Bobbie LOWE, Randy NESSE, Steven PINKER, JohnTOOBY and E. O. WILSON). The editors are willing toconsider sharing their comments in future issues of thisjournal.2 We typically use Evolutionary Psychology in capitals whenwe refer to the revolutionary, social-psychological versionof the discipline, and lower case when we refer to the moregeneric variety.ReferencesBadcock, C. (2000) Evolutionary Psychology: A Critical Introduction.Polity Press: Cambridge, UK.Baily, P./Davis, E. W. (1943) Effects of lesions of the periaqueductalgray matter in the cat. Proceedings of the Society forExperimental Biology and Medicine 51: 305–306.Barkley, R. A. (2001) The excecutive functions and self-regulation:An evolutionary neuropsychological perspective.Neuropsychology Review 11: 1–29.Basso, A./Sala, S. D./Farabola, M. (1987) Aphasia arisingfrom purely deep lesions. Cortex 23: 29–44.Batro, A. M. (2000) Half a Brain Is Enough: The Story of Nico.Cambridge Univ. Press: Cambridge UK.Bauman, M. L./Kemper, T. L. (1995) Neuroanatomical observationsof the brain in autism. In: Panksepp, J. (ed) Advancesin Biological Psychiatry, Vol. 1, Greenwich CT: JAIPress, pp. 1–26.Bechtereva, N. P./Abdullaeva, Y. G. (2000) Depth electrodesin clinical neuropsychology: neuronal activity and humancognition function. International Journal of Psychophysiology37: 11–29.Berridge, K. C. (2000) Measuring hedonic impact in animalsand infants: microstructure of affective tast reactivity patterns.Neuroscience and Biobehavioral Reviews 24: 173–198.Berridge, K. C./Valenstein, E. S. (1991) What psychologicalprocess mediates feeding evoked by electrical stimulationof the alteral hypothalamus? Behavioral Neuroscience105: 3–14Bouchard, T. J. (1998) Genetic and environmental influenceson adult intelligence and special mental abilities. HumanBiology 70: 257–279.Bouchard, T. J./McGue, M./Lykken, D./ Tellegen, A. (1999)Intrinsic and extrinsic religiousness: Genetic and environmentalinfluences and personality correlates. Twin Research2: 88–98.Buchner, R. L./Corbetta, M./Schatz, J./Raichle, M. S./Petersen,S. E. (1996) Preserved speech abilities and compensationfollowing prefrontal damage. Proceedings of the nationalAcademy of Sciences 93: 1249–1253.Borod, J. C. (2000) The Neuropsychology of Emotion. OxfordUniversity Press: New York.Breitenstein, C./Van Lancker, D./Daum, I./Waters, C. H.(2001) Impaired perception of vocal emotions in Parkinson’sdisease: Influence of speech time processing and executivefunctioning. Brain and Cognition 45: 277–314.Burgdorf, J./Knutson, B./Panksepp, J./Ikemoto, S. (2001)Nucleus accumbens amphetamine microinjections unconditionallyelicit 50 kHz ultrasonic vocalizations in rats. BehavioralNeuroscience. In Press.Burgdorf, J./Knutson, B./Panksepp, J./Shippenberg, T.(2001) Ultrasonic vocalizations index pharmacologicalaversion in adult rats. Psychopharmacology 155: 35–42.Burgdorf, J./Panksepp, J. (2001) Tickling induces reward inadolescent rats. Physiology & Behavior 72: 167–173.Carter, S./Kirkpatrick, B./Lederhendler, I. I. (eds) (1997)Neurobiology of affiliation. Annals of the New York Academyof Sciences 807. New York Academy of Sciences: NewYork.Castro-Caldas, A./Petersson, K. M./Reis, A./Stone-Lander, S./Ingvar, M. (1998) Learning to read and write during childhoodinfluences the functional organization of the adultbrain. Brain 121: 1053–1063.Charney, D. S./Nestler, E. J./Bunney, B. S. (eds) (1999) Neurobiologyof Mental Illness. Oxford University Press: NewYork.Cosmides, L./Tooby, J. (2000) Evolutionary psychology andthe Emotions. In: Lewis, M./Haviland, J. (eds) The Handbookof Emotions, 2nd edition. Guilford: New York, pp.91–116.Crosson, B./Zawacki, T./Brinson, G./Lu, L./Sadek, J. R. (1997)Models of subcortical functions in language: current status.Journal of Neurolinguistics 10(4): 277–300.Daly, M./Wilson, M. (1988) Homicide. Aldine: New York.Damasio, A. R. (1999) The Feeling of What Happens, Bodyand Emotion in the Making of Consciousness. HarcourtBrace: New York.Darwin, C. (1998) The Expression of the Emotions in Manand Animals, 3rd edition. Oxford University Press: NewYork. Originally published 1872.Dawson, G./Fisher, F. W. (eds) (1984) Human behavior andEvolution and Cognition ❘ 78 ❘ 2001, Vol. 7, No. 1

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Shulamith KreitlerAn Evolutionary Perspectiveon Cognitive OrientationIntroductionThe purpose of this paperis twofold. The first goal isto describe how the cognitiveguidance of behaviorhas developed in an evolutionarysense. The conceptionthat cognitionplays a role in regard tosurvival and behavior liesat the core of the evolutionaryepistemology theoriesthat consider knowledgeas an evolutionaryprocess subserving survival(CAMPBELL 1974;LORENZ 1977; POPPER1984; RIEDL 1980). Thesetheories provide the bioethologicalbasis for thescientific endeavor to describehow cognitionguides behavior. Startingwith a cognitive theory of motivation on the humanlevel, we will examine its evolutionary developmentand implications. In view of the novelty of the goal,that which we can expect to do in a first exploratorypaper of this kind is to confront the issue of the evolutionaryaspects of the cognitive guidance of behaviorand trace some of the major tracks along whichthe cognitive guidance of behavior seems to have developedin the framework of evolution. In order toclarify our question, it is advisable to emphasize thatwhile it is well-known that in the course of evolutionboth cognition and behavior have developed each inits own right and in terms of its own characteristics,it is neither well-known nor obvious that the cognitiveguidance of behavior did. This is precisely ourthesis. Starting from the assumption that in livingorganisms cognition guides behavior (KREITLER/KRE-AbstractThe paper deals with evolutionary aspects of the cognitiveguidance of behavior. It is assumed that cognitionguides behavior because of empirical evidenceand because the impressive development of cognitionin the course of evolution suggests that it plays a rolein regard to survival. The theory of cognitive orientation—atheory of cognitive motivation on the humanlevel—is presented as a basic framework for the evolutionaryanalysis. The major advantages of this theoryare that it provides a comprehensive description of theprocesses intervening between input and output andhas ample empirical support. The major evolutionaryaspects discussed concern the differentiation betweenmotivation and cognition, the development of meaningand the development of action, emphasizing therole of cognition in regard to each.Key wordsCognition, development, meaning, belief, behavioralprogram.Evolution and Cognition ❘ 81 ❘ 2001, Vol. 7, No. 1ITLER 1976), the questionof the evolutionary developmentof the cognitiveguidance of behavior appearsto be both legitimateand relevant. It is legitimatebecause any phenomenon,structure orprocess that has been observedon the humanlevel has analogues or primaryrudimentary manifestationson earlier evolutionarylevels (MITHEN1996; THORPE 1974). It isrelevant because consideringthe evolutionaryunderstructure and aspectsof a basic humanprocess provides increasedinsights and opportunitiesfor deeper understandingof its structuraland functional properties.Thus, our exploration will be guided by questions,such as “What happened to the cognitive guidanceof behavior in the course of evolution?”, “Whathas evolution done to it?”, “How was it affected byevolution?”, “Which of its components were shapedand transformed by evolution?”Our second goal has to do with SKINNER. Years agoduring a Sabbatical stay at Harvard University I hadthe chance to discuss psychological issues with SKIN-NER, who at the time was already retired from HarvardUniversity (1975–1976). As it happened I waslucky enough to have stayed in the same neighborhoodin Cambridge, MA where he lived and thus hadample opportunity of discussing science with him.After he became familiar with the theory of the cognitiveguidance of behavior that Hans KREITLER andI had developed—the theory of cognitive orienta-

Shulamith Kreitlertion—he said one day (I bring his views in a freeparaphrase of my notes from the time): “The cognitiveorientation theory is the extreme opposite of thebrand of behaviorism that I have developed andwhich came to be called SKINNERIANISM. It is oppositebecause it starts on the top, uses an apparently unparsimoniousset of theoretical concepts, assumesprimacy of the cognitive system, and attributes acentral role to cognition in the functioning of organisms.In contrast, SKINNERIANISM starts at the verybottom, uses an apparently monastically frugal setof a-theoretical concepts, does not accord any statusto the cognitive system, whose existence it barelyacknowledges and considers as a theoretical outrage,and attributes no role to it in the functioning of organisms.However, an integral part of the developmentof an approach in science (viz. theory) is to testits extensibility. Therefore, I was intent to test howfar up the evolutionary ladder I can get, in order tofind out how much I can explain with my theoreticallymeager tools on the higher levels of psycho–biological functioning. That’s why I tried to explaineven politics in behaviorist terms and why I wroteWalden Two, a book about a utopian society basedonly on operational conditioning. You should do thesame with the cognitive orientation theory, thoughin an inverse sense. Your task is to test how far downthe evolutionary ladder you can get with the cognitiveorientation theory, in order to find out howmuch you can explain with your theoretically richtools on the lower levels of psycho–biological functioningpreceding the human. Since you start with abig load of concepts, stepping down the evolutionaryladder should give you a chance to see whichcomponents developed and why. Frankly, I envy youthis task”. So, this is the task with which this paperdeals. Notably, the procedure suggested by SKINNERcorresponds to the use of evolutionary thinkingwithin the top-down system advocated by DEKAYand SHACKELFORD (2000). Handling this task was delayedso much probably because we had first to learnmuch more about the cognitive orientation theorybefore we could put it to the test of ‘extensibility’.Cognition: What It Is not and What It IsThe basic thesis of the cognitive orientation (CO)theory is that cognitive contents and processes guidebehavior. Cognitive contents refer to information,beliefs, attitudes, opinions, values, and meanings inthe broad sense of the term, whereas cognitive processesare operations and transformations of differentkinds that deal with the production, testing,elaboration, transformation, analysis, storing,retrieval and uses of cognitive contents. Cognition,as defined above, is not restricted to any particularspecies and occurs on different evolutionary levels.In principle, we should expect its barest primarymanifestations in the simplest organisms (RIEDL1980, 1981, 1987).In order to understand better the role of cognitionin the motivational context, it is necessary to emphasizethat cognition does not entail rationality and noteven reasonableness. The fact that humans maythink rationally does not mean that they always ormostly do or that this kind of thinking prevails in themotivational context. Rationality constitutes one setof rules which may guide cognition, but there areother sets, such as Aristotelian logic, Boolean logic,mythological thinking, symbolic thinking, magicalthinking etc. (CASSIRER 1953; KREITLER 1965; LÉVI-STRAUSS 1979).Likewise, the involvement of cognition in motivationdoes not mean that motivation depends on orinvolves a decisional process which considers, for example,benefits and costs, advantages and disadvantages,pros and cons, weighing each carefully and calculatingthe end-result according to normativedecision rules (as some models of cognitive motivationassume, e.g., BECKER 1974; ROGERS 1983; RONIS1992).Further, cognition does not entail consciousness,nor is it a product or function of consciousness. Consciousnessis itself a function of cognition, and mayaffect cognitive functioning but is not a necessarycondition for cognition (KREITLER 1999). Cognitiveprocesses run their course without consciousness asmost other organismic processes. They may becomeconscious to a certain extent, under specific conditions,when this subserves the needs of the organism.Similarly, cognition does not entail voluntary control.Though the exercise of so-called voluntary controlinvolves cognition, the involvement of cognitiondoes not mean that one has voluntary control overany aspect of the motivational dynamics of behavior.Finally, cognition is not restricted to the verbal medium.Indeed, cognitive processes may occur withoutrelying on any formal medium of expression, andwhen some of their aspects or end-results are expressedor communicated this may occur by meansof any form of expression, verbal or nonverbal.So, given that cognition is not necessarily logical,reasonable, conscious, voluntarily controlled andverbal, which positive property can be attributed toit? We venture to suggest that it is meaning-dependent.Evolution and Cognition ❘ 82 ❘ 2001, Vol. 7, No. 1

An Evolutionary Perspective on Cognitive OrientationMeaning is a concept that plays a major role incognition. It is defined as a pattern of cognitive contents(called ‘meaning value’, e.g., a color, an emotion)focused on some input (called referent, e.g., astimulus, an object) that is expressed verbally ornonverbally, and forms together with the input orsubject a meaning unit. Examples of meaning unitsare The sea—blue; Table—made of wood; Car—withfour doors; Virus—scary. In a given context meaningconsists of a sequence of meaning units, each ofwhich may be characterized in terms of the contentsassigned to the subject (e.g., causes, sensory qualities)and the relations between it and the subject(e.g., its directness, generality). The characterizationsconstitute a set of variables (called ‘meaningvariables’) which enable assessing the meaning ofany input or the meaning assignment tendencies ofan individual. Meaning includes the more interpersonalpart as well as the more personal–subjectivepart, and may vary in contents, structure, varietyand complexity (KREITLER/KREITLER 1985, 1986,1987a, 1989, 1990a, 1990b; KREITLER 1999).Meaning provides the raw materials for the contentsof cognition and the functional frames for theprocesses of cognition. Accordingly, cognition maybe defined as a system which produces, elaborates,transforms, stores and uses meanings. Hence, theproperties, contents, processes, functioning andproducts of cognition are functions of meaning.When we conceive of a cognitive model of motivation,we talk in fact of a model of motivation inwhich meaning fulfills a major role. Basically, the COmodel is a model based largely—though not exclusively—onthe construction, transformation and applicationof meanings. It describes the processes interveningbetween input and output not as a seriesof rational decisions or problem solving, but as outputsof meanings and clustered orientative beliefs,which give rise to motivational dispositions manifestedin behavior.The Cognitive Orientation Theory:General CharacteristicsIn its original presentations CO was a theory of cognitivemotivation of behavior elaborated in regardto overt behaviors (KREITLER/KREITLER 1965, 1972,1976, 1982). Later developments led to the elaborationof models applicable to further specificdomains, such as emotions, physiological processesof disease and health, psychopathology and cognitivebehaviors (KREITLER 1999; KREITLER/KREITLER1986, 1989, 1991a, b, 1997). Accordingly, at presentit would be justified to distinguish between the differentmore specific CO models and the comprehensiveCO theory, which refers to the generaltheoretical concepts and procedures characteristicfor all these models. In the present context we willfocus on the basic CO theory and CO model ofbehavior.CO is a cognitive model of behavior designed toprovide an account of the major processes interveningbetween input and output and to enable understanding,predicting and changing human behavior.This introductory statement emphasizes several importantcharacteristics of the CO model. First, asnoted, the original application domain of CO hasbeen human behavior. Secondly, it is a model focusedon actual observable overt behavior, as distinguishedfrom self-reported behavior or decisions toact, commitments to action and other replacementsof behavior. Thirdly, the purpose of the model is toprovide concepts and methods that would be directlyapplicable for understanding retrospectively,predicting prospectively, and changing at presenthuman behavior. While different psychologicalmethods allow in principle indirectly the attainmentof the specified goals, the special characteristicof the CO model is that it enables attaining the mentionedgoals by the same methods (rather than onegoal by one method, and another goal by anothermethod) and that it functions by direct applicationof the methods so that interpretational pitfalls areavoided on the way. Fourthly, the CO model is acomprehensive model that addresses all the majorlinks and stages intervening between the input andthe output, in contrast to other motivational modelsthat focus on one or another stage or aspect of theprocess (e.g., planned behavior, AJZEN 1985; healthbelief model, BECKER 1974).The basic thesis of the CO theory is that cognitivecontents and processes guide behavior. The rationalefor this thesis is twofold. The first reason is rooted inthe empirical–scientific approach of psychology. Onthe human level there is a large body of researchdemonstrating the contributions of cognition to behaviorand the utility of this assumption (for a reviewsee KREITLER 2002).The second reason is rooted in the more generalevolutionary approach. It assumes that since cognitionhas undergone such a spectacular developmentin the course of evolution (WILSON 1978, p87), it ishardly possible that it does not contribute to behaviorwhich is the prime tool of survival. In this sense,CO is embedded in evolutionary epistemology,which conceives of a continuum from biological toEvolution and Cognition ❘ 83 ❘ 2001, Vol. 7, No. 1

Shulamith Kreitlercognitive processes and attributes to cognition a rolein survival (KOVÁ Č 2000; LORENZ 1977; PLOTKIN1997; RIEDL 1980, 1987).Accordingly, CO is a behavior-oriented theory,which means that whatever the role of cognition,CO is not a theory of cognition but of motivationand the focus is always on behavior. The purpose ofapplying cognition is to understand, describe, predictor change behavior.In addition to the basic thesis about the involvementof cognition in motivation, the CO theory assumesthat there is a basic distinction between themotivational disposition for behavior and the performanceof behavior. Both are indispensable for anyact, at least on the human level. The disposition providesthe directionality and answers, as it were, thequestions “Why did this behavior occur?” or “Whydid the behavior occur specifically in this direction?”The performance of behavior is the operational manifestationof the directionality and answers, as itwere, questions, such as “Why did behavior take theform that it did?”, “Why was the behavior executedspecifically in this manner or with these means?”The distinction between the directionality of behaviorand the manner of its enactment is evident alsoon lower evolutionary levels, even in regard to reflexes,defensive responses and tropisms. Referring,for example, to the flight of bees, GALLISTEL (1990,pp527–528) writes: “The behavioral purpose is establishedby the bee’s motivational state, which is conceivedof as a set of signals that selectively potentiatesa functionally cohesive set of acts” (see ibid alsopp87–88 concerning the migration of birds). The distinctionitself, its sharpness and manifestations arean evolutionary product (see “Evolutionary aspectsof the CO theory”).The motivational disposition and the performanceof behavior differ greatly in their components,structure, and dynamics. Also, the involvementof cognition in motivation differs in regard tothe disposition and the performance, as will becomeevident in the next section.The last general characteristic of the CO theory tobe mentioned in this context concerns the multilevelednessof its structure and functioning. Ratherthan assuming a dominant single course of actionon a relatively high level, according to the CO modelthe input is first addressed in terms of the lower andsimpler levels of functioning, that are replaced bymore complex ones only if they fail to deal with thesituation, according to concurrent or previous experience.The lower and simpler levels of functioningcorrespond to those that may be identified in lowerevolutionary levels. They persist in more advancedspecies, such as the human, possibly in order to handlesimpler situations or more basic survival tasks. Beit as it may, the persistence of the lower levels sideby side with the higher levels as functional optionswithin the same organism are in accord with thebasic evolutionary principle whereby higher developmentalproducts do not necessarily displace or replacelower ones, but may rather complement, transformand limit their scope. An example is providedby the different stages of evolution contained withinthe brain of homo sapiens, sometimes called the reptilian,mammalian and human brain (MCLEAN1973). This phenomenon may be called “preservationof the fittest”. Moreover, the CO theory assumesthat issues are handled first on the lower levels, oralternately, that whatever can be handled on a lowerlevel is not handled on a higher level. This set ofprinciples built into the CO model demonstrates theextent to which the evolutionary perspective hasbeen built into the theory down to its fundamentals.The CO Theory: How Does It Function?Major Theoretical StagesIn the present context only a brief presentation ofthe CO model of behavior is provided. It will bedescribed in its original form, as it applies on thehuman level. Later sections will focus on evolutionaryaspects.The CO theory provides detailed descriptions ofthe processes intervening between input and output.These can be grouped into four stages, eachcharacterized by metaphorical questions and answers.The first stage is initiated by an external orinternal input and is focused on the question “Whatis it?” It consists in identifying the input in terms ofa limited and primary ‘initial meaning’ as one of thefollowing: (a) a signal for a defensive, adaptive orconditioned response, or (b) a signal for molar action,or (c) as irrelevant in the present situation, or(d) as new or especially significant, and hence as asignal for an orienting response.The first alternative consists in the evocation of adefensive or adaptive response, unconditioned orconditioned. If feedback indicates that the input hasbeen properly handled, the behavioral act is broughtto an end. The second alternative consists in theemergence of a meaning that identifies the input outfrom the start as one that cannot be handled bymeans of a conditioned or unconditioned responseand calls for a response on a higher level (viz. molar),that requires more preparatory elaboration. TheEvolution and Cognition ❘ 84 ❘ 2001, Vol. 7, No. 1

An Evolutionary Perspective on Cognitive Orientationthird alternative consists in the emergence of ameaning that includes no components enabling, indicatingor compelling for action. Hence, the inputis dismissed as irrelevant. The fourth alternativearises when the input cannot be identified, for example,because too little information is available, or noresemblance could be established between it andpatterns of stimuli stored in memory. Hence, it istransferred to a higher level of elaboration.The second stage is initiated by feedback indicatingfailure of the conditioned or unconditioned responsesto cope with the situation, or by a meaningsignalling the need for molar action, or by an inputthat has failed to be sufficiently identified despite theorienting response. It is focused on the question“What does it mean in general and what does itmean to or for me?” An enriched process of meaninggeneration sets in, based on extended elaboration ofboth the interpersonally-shared and the personalkinds of meaning, in terms of meaning units in theform of beliefs. Beliefs are cognitive units (namely,they include a referent and a meaning value), expressedverbally or not, which express some messagepresented as true (acceptable, adequate, etc.) or not.The beliefs that emerge at this stage may refer to theindividual’s goals, norms, oneself, or others and reality.By examining the extent to which the person’sgoals, norms, beliefs about oneself and reality areinvolved, meaning generation leads eventually to aspecification whether action is required or not. Insofaras meaning elaboration indicates that the situationis relevant in regard to the person’s goals,norms, beliefs about self or general beliefs, the indicationis pro-action or at least, going on with theprocess of elaborating the input, even though thenature of the action or its direction remain unspecified.In this context, relevance of beliefs means thata goal arises which may seem to be attainable in thesituation, a norm belief is evoked which implies thenecessity of undertaking action, or a belief aboutoneself or a general belief arises which indicates possibledangers in the situation if no action is undertakenor if action is undertaken. Thus, the output ofthe second stage is indication for action, blocking ofaction, or no indication for action.If there is indication for action or at least no blockingof action, the third stage sets in. It is focused onthe question “What will I do?” The answer is soughtby means of beliefs characterized in terms of relevantcontents and a specific form. Relevance in contentsdenotes deep underlying meanings of the involvedinputs rather than the superficial obvious explicitmeanings. The deep meanings are characterizedBelief Subject RelationSelf I FactualNorms Non-I DesirableGoals I DesiredGeneral Non-I FactualFigure 1: Structure of the Four Types of Beliefs.first, by being of the personal–subjective type ratherthan the interpersonally-shared type and second, bybeing relatively remote though not dissociated fromthe surface meanings. For example, when the inputis ‘dieting’, the surface meaning is likely to be “eatingless”, and the deeper personal meanings, which donot necessarily refer specifically to eating, could be“renouncing pleasures”, or “accepting restrictionson one’s actions”. Meaning elaboration of the inputin successive steps leads to the emergence of differentdeeper meanings.As noted, the beliefs are characterized also interms of form. The beliefs that play a role in the thirdstage are formally of four types: (a) Beliefs about goals,which express actions or states desired or undesiredby the individual (e.g., “I want to be respected byothers”); (b) Beliefs about rules and norms, which expressethical, esthetic, social and other rules andstandards (e.g., “One should be assertive”); (c) Beliefsabout self, which express information about oneself,such as one’s habits, actions, feelings or abilities(e.g., “I rarely become excited”, “I was born in Israel”);and (d) General beliefs, which express informationconcerning others and the environment (e.g.,“The world is a highly dangerous place”). The fourtypes of beliefs may be characterized formally interms of the kind of subject and the kind of relationbetween the subject and predicate. Figure 1 presentsthe formal structure of the four types of beliefs.Accordingly, the cognitive elaborations in thethird stage refer to beliefs of four types that representdeep underlying meanings of the involved inputs.The meaning elaborations consist in matchings andinteractions between beliefs (‘belief clustering’),based on clarifying the orientativeness of the beliefs(i.e., the extent to which they support the indicatedcourse of action). If the majority of beliefs of a certaintype support the action, that belief type is consideredas positively oriented in regard to that action. Alternately,it may be negatively oriented or lack any orientativeness.If all four belief types support a certainaction, or at least three support it whereby the fourthEvolution and Cognition ❘ 85 ❘ 2001, Vol. 7, No. 1

Shulamith KreitlerBeliefs about SelfBeliefs about NormsBeliefs about GoalsGeneral BeliefsFigure 2: The Behavioral Intent.BehavioralIntentis neutral, a cluster of beliefs is formed (‘CO cluster’)orienting toward a particular act. It generates a unifiedtendency orienting toward the performance ofan action, which is called behavioral intent, and canbe considered as a vector representing the motivationaldisposition towards a given behavior (see Figure2).In many cases the process of forming a CO clusteris greatly reduced because there exist in memory almostcomplete CO clusters, formed on the basis ofpast recurrences of behaviors in similar situations.The meanings of the input allow for retrieving suchalmost-complete CO clusters and adapting them tothe present situation by the addition or transformationof a few beliefs. In other cases, when there arenot enough beliefs orienting toward the course ofaction in at least two belief types, no CO cluster willbe formed. The process will discontinue, especiallyif there are no personally relevant beliefs in the cluster.Similarly, when there is a paucity of beliefs of acertain type, an incomplete CO cluster will beformed which cannot be acted upon but may be discardedor stored for possible use in another situation.Another resulting possibility is the occurrence ofconflict (‘intent conflict’) when two CO clusters andconsequently two behavioral intents are formed.Another intriguing possibility consists in the formationof an inoperable cluster due to the inclusion of‘as if’ beliefs in one or more belief types. If, for example,the ‘as if’ beliefs come to replace other beliefs inthe beliefs about self and general beliefs, we may geta CO cluster orienting toward virtual action such asdaydreaming.The fourth stage is initiated by the formation of thebehavioral intent, and is focused on the question“how will I do it?” The answer is in the form of abehavioral program, namely, a hierarchically structuredsequence of instructions governing the performanceof the act, including both the more generalstrategy (‘program scheme’) as well as the more specifictactics (‘operational program’). Previouslyformed habits are major components embedded inthe programs. An important role is attributed to thebehavioral intent at this phase. It guides the selection,retrieval, modification and sometimes constructionof behavioral programs, and serves as aframe of reference for supervising program executionand evaluating outcomes.Different programs are involved in executing anovert molar act, a cognitive act, an emotional response,a daydreamed act, conflict resolution, etc.There appear to be four basic kinds of programs: (a)Innately determined programs, which are involvedprimarily in the output on the submolar level, e.g.,those controlling reflexes or classically conditionedresponses; (b) Programs that are partly innate andpartly acquired by learning, e.g., those controllinginstincts (as described by LORENZ 1965, or TINBERGEN1951), behaviors corresponding to defense mechanisms,and language behavior; (c) Programs acquiredthrough learning, e.g., those controlling culturallyshaped behaviors (running elections, etc.),or personal habits (e.g., forms of relaxing, makingfriends, arranging one’s cupboard); and (d) Programsconstructed by the individual ad hoc, in linewith situational requirements, often using existingprogram schemes, including plans for plan construction.Implementing a behavioral intent by a programrequires selecting and retrieving a program, and oftenadapting it to prevailing circumstances. A ‘programconflict’ may occur between two equally adequateprograms or between one that is about to beenacted while another is still in operation.The retrieval, adaptation and execution of theprogram, as well as determination of the end-pointin view of the behavioral intent also require cognitivecontrol.Major Advantages of the CO TheoryThe major cited advantages will be considered fromthe perspective of the human level, the level forwhich the theory was originally developed and inwhich it was primarily applied. The advantages arepartly applied and partly theoretical.The major advantage from the applied viewpointis that the CO theory enables predictions of behaviorsand changes in behavior by using systematicprocedures.Evolution and Cognition ❘ 86 ❘ 2001, Vol. 7, No. 1

An Evolutionary Perspective on Cognitive OrientationInput notidentifiedActionrequiredInputWhat isthe Input?MeaningactionWhatdoes it meanfor me?Whataction?4 types of beliefs:Cog. orientationclusterBehaviorialIntentHowto performaction?BehavioralprogramInputInputidentified Reflex, dealt withConditioned ExitResponse, etc.Input identifiedfor molaractionInput notdealt withInput not requiredExitBehaviorFigure 3: A Schematic Flow-Chart of the CO ModelA large body of research demonstrates the predictivepower of the CO theory in a variety of domains,e.g., coming on time, reactions to success and failure,curiosity, achievement, studying mathematics,planning, assertiveness, conformity, cheating, overeating,breast-feeding, cessation of smoking, self-disclosure,rigidity, defensive responses, undergoingtests for the early detection of breast cancer, sexualresponses, compliance in diabetes patients etc. Allstudies refer to actual observed behaviors. The participantswere adults, adolescents, children, retardedindividuals, schizophrenics, individuals with differentphysical disorders, etc. (KREITLER/CHAITCHIK/KRE-ITLER/WEISSLER 1994; KREITLER/CHEMERINSKI 1988;KREITLER/KREITLER 1976, 1982, 1988, 1991a, 1993,1994a, 1994b; KREITLER/NUSSBAUM 1998; KREITLER/SCHWARTZ/KREITLER 1987; KREITLER/SHAHAR 1976; LO-BEL 1982, NURYMBERG/KREITLER/WEISSLER 1996; TIP-TON/RIEBSAME 1987; WESTHOFF/HALBACH-SUAREZ1989).All studies confirmed the hypothesis that behaviorwould occur if it was supported by at least threeThemes1)2)3):n)BeliefsaboutSelfBeliefsaboutNormsBeliefsaboutGoalsFigure 4: The Predictive Matrix of Beliefs.GeneralBeliefsbelief types and a behavioral program was available.The success of the predictions is based on applyingthe special standardized procedure developed in theframework of the CO theory (KREITLER/KREITLER1982; in press). The procedure consists in assessingthe motivational disposition for the behavior (viz.behavioral intent) by means of a CO questionnaireand examining the availability of a behavioral programfor implementing the intent. A CO questionnaireassesses the degree to which the participantagrees to relevant beliefs orienting toward the behaviorin question or rejects those that do not orienttoward it. The beliefs differ in contents and form. Incontents they refer to themes which represent meaningsunderlying the behavior in question (called‘themes’). In form they refer to the four types of beliefs,namely, beliefs about goals, beliefs about rulesand standards (or norms), beliefs about the self, andbeliefs about others and reality (called general beliefs).Thus, a CO questionnaire mirrors the predictionmatrix (see Figure 4). It usually consists of fourparts presented together in random order, each representingone of the four types of beliefs and containingin random order beliefs referring to the differentthemes. The participant is requested to checkon a 4-point scale the degree to which each beliefseems to him/her true (or correct).The themes of the CO questionnaire are identifiedby means of a standard procedure applied to pretestparticipants who are known to manifest the behaviorin question or not. The procedure consists in interviewingthe participants about the meanings ofthe key terms and then in turn sequentially abouttheir responses concerning the meanings (see Figure5 for a schematic representation of the procedure).Repeating the questions about meanings leads todeeper-layer meanings, out of which those that recurin at least 50% of the interviewees are selected for theEvolution and Cognition ❘ 87 ❘ 2001, Vol. 7, No. 1

Shulamith KreitlerInterpersonally-shared meaning ofbehavior = key wordsLevel 1:Personal meanings of key-words_Level 2: Personal meanings of responses on level 1_Level 3: Personal meanings of responses on level 2Figure 5: The Process of Identifying Themesby means of Meaning Generation.final questionnaire. Examples of themes for successin dieting are acceptance of limitations, and viewingthe world as a safe place. It is important to emphasizethat the beliefs in the questionnaire do not refer inany way to the behavior in question but only to thethemes which represent the underlying meanings.Identifying the themes is followed by constructing aCO questionnaire, which has to be examined for itspsychometric properties, including reliability andvalidity before it is ready for application. Althoughfor every kind of behavior it is necessary to constructa particular CO questionnaire, a single CO questionnairemay predict a broad range of relevant behaviors,e.g., the CO questionnaire of curiosity predicted14 different curiosity behaviors (KREITLER/KREITLER1994b).The availability of the behavioral program is establishedby means of questionnaires, observation,information from others or role-playing.The advantages of the prediction procedure generatedby the CO theory are the following: (a) it hasprovided a great number of significant predictions ofactual behaviors; (b) the predicted behaviors were ofmany different kinds and from different domains;(c) the participants whose behaviors were predictedwere of different ages and intelligence levels, evencognitively challenged individuals; (d) applying theprocedure does not require any special ‘mind-set’,preparation, intention, or even average intelligenceon the part of the participants; (e) the procedure doesnot enable the participants to tailor their responsesso that it would or would not correspond to theirbehaviors because it is impossible to unravel fromthe statements in the CO questionnaire the kind ofinvolved behavior; (f) the procedure may be appliedin regard to any behavior whatsoever provided thatit can be assessed; (g) applying the prediction procedureis straightforward and does not require the involvementof further criteria or constructs or assumptionsor the creation of particular conditions inregard to any behavior.The CO theory has also enabled successful modificationsof behavior, such as rigidity, impulsivity,curiosity and eating disorders (BACHAR/LATZER/KREITLER/BERRY 1999; KREITLER/KREITLER 1988, 1990C,1994B; ZAKAY/BAR-EL/KREITLER 1984). The essentialcomponents of the procedure consist in mobilizingsufficient support for a specific course of action byevoking in the participant beliefs orienting towardthis course of action. The major conditions on whichsuccess of the intervention depend are that theevoked beliefs (a) refer to the themes (viz. underlyingmeanings of the action), (b) refer to all four types ofbeliefs (viz., goals, norms, self and general), and (c)that they have originated in the participant by a processof meaning generation. The procedure for interventionor change of behavior has been standardizedand can be applied in individual or group sessions(KREITLER/KREITLER 1990c). Again, there are no limitationson the types of behaviors or of participantsin regard to which it can be applied.The major theoretical advantages of the CO theoryare that it shows precisely what is the role ofcognition in regard to motivation and behavior. Theassumption that attitudes, values, informations andopinions are important in regard to behavior isdeeply rooted in Western culture and underlies education,persuasion and dissemination of informationthrough the media and otherwise. However, alarge body of studies in the last century showed thatattitudes were not related to behavior (e.g., COREY1937; LAPIERE 1934). This evidence posed a seriousproblem for behavioral scientists (KREITLER 2002).The CO theory provided a plausible and operationalaccount of the role of cognition in regard to motivationand behavior. Thereby, the CO theory also clarifiedthe role of several additional basic concepts inregard to motivation and behavior, mainly the self,values and norms, and attributions, causal and others.Evolutionary Aspects of the CO TheoryThe evolutionary aspects of the CO theory will bediscussed in two complementary forms. The firstwill be in terms of general developmental trends,the second—in terms of the stages interveningbetween input and output in line with the CO theory(see section “The CO theory: How does it function?Major theoretical stages”). In both cases thediscussion will be based partly on extrapolationsfrom observations on ontogenetic development,and partly on information about animal behavior.However it may be, the purpose is merely to traceEvolution and Cognition ❘ 88 ❘ 2001, Vol. 7, No. 1

An Evolutionary Perspective on Cognitive Orientationthe blueprint of evolutionary aspects involved inthe cognitive guidance of behavior.General developmental trends. The first and possiblythe most important trend is the differentiationof action and motivation. This differentiation doesnot seem to exist at the lowest evolutionary levels.On these levels we find actions, which may beprompted by inner or outer states that are not differentiatedfrom the action and cannot be identified assuch. As stated most precisely by WIMMER (1995,p43), “information about [homeostatic] disturbanceand motoric pattern are combined in a unit whichbecomes segregated at higher levels of organization,e.g., taxis behavior”. The differentiation occurswhen we first obtain some kind of representation ofthe inner or outer stimulus. This constitutes the basisfor the development of motivation in later phases,as distinct from action. The two arms—of motivationand of action—undergo different transformationsand development in the course of evolution.The development of motivation depends largely onmeaning, whereas the development of action dependslargely on structures.Accordingly, the second basic trend is the evolutionof meaning. This evolution takes differentforms, some of which overlap, whereas others occurin a sequence. The development is partly structuraland partly dynamic. The structural developmentconsists basically in shaping the major outlines ofthe meaning system. This entails differentiating thecontents of the meaning values into subsets whichwe call meaning dimensions, differentiating the relationsof the contents to the referents in terms ofdirectness, generality, positivity and the nature ofthe relation (propositional, normative, desired), andmoving from the nonverbal actional means of expressionto the use of diversified nonverbal and verbalmeans of expressing meaning values and meaningunits. On higher levels, the whole system splitsinto two major branches of the interpersonallysharedmeaning and the subjective–personal meaning,which may intertwine and interact to differentdegrees. The more dynamic development consists inmoving from meaning values to meaning units(each of which includes a meaning value centeredaround a referent), from meaning action to meaninggeneration, and from surface or label meaning toelaboration of the deeper underlying meanings. Thelatter development brings aboutan increasing involvementof personal–subjective meaning in additionto the interpersonally-shared one which dominatesthe surface-label meaning.A special line of development is taken by meaningunits. It will be mentioned because it is of particularimportance for the cognitive guidance of motivation.This line leads from meaning units to beliefs.Both meaning units and beliefs share the basic structureof a referent related to a meaning value. However,they differ in function. Meaning unit is a conceptfrom the domain of meaning and its functionis the expression or communication of meaning perse. In contrast, belief is a concept from the domainof action or behavior, and its function is to expressor communicate some cognitive contents—e.g., astatement, a view, an attitude, an opinion, an information,a value, an expectation etc.—with behavioralimplications.The primary meaning unit is a belief which statesa fact, namely, the relation of the meaning value tothe referent is factual, existential. Hence, it could beeither a belief about self or a general belief. It is difficultto decide which one was the first. But in viewof the primary focusing on oneself it seems likelythat the belief about self preceded the general belief.Be it as it may, each of these primary units split toform by differentiation two distinct varieties. Thus,the belief about self differentiated into a belief aboutgoals and a belief about self, both of which share the‘I’ as a referent and differ only in the kind of relationbetween the meaning value and the referent (viz.desired vs. factual, respectively). Similarly, the generalbelief differentiated into a belief about normsand a general belief, both of which share the non–Ias a referent and differ only in the kind of relationbetween the meaning value and the referent (viz.normative/desirable vs. factual, respectively). As aresult, four types of beliefs emerged.Complementing the evolution of meaning is thethird trend, which refers to the evolution of action.The major characteristics of evolution in this spherehave been extensively studied. One characteristicconsists in increased variety of action units. On thelower evolutionary levels we find genetically and innatelydetermined units of action that are rigidly enactedin a specific manner. New units are added tothe repertory which modify the innately given actionalunits or combine with them in different waysand thus form, for example, conditioned responsesand habits. Further, the units of action which initiallymay be bound to specific inputs or situationsbecome gradually free of such constraints, and maybe used in a variety of contexts, or for the attainmentof purposes completely divorced from those withwhich they may have been related when acquired.Another characteristic development consists in in-Evolution and Cognition ❘ 89 ❘ 2001, Vol. 7, No. 1

Shulamith Kreitlercreased complexity of organization of the actions.For example, several action units may combine intolarger units by forming a sequence or chain of unitsor a hierarchically structured ‘family of habits’,wherein one or more action units may be replacedby other units when the former do not attain thegoal.The increased organizational complexity enablesalso increased flexibility in form, structure and enactment,which is manifested, for example, by selectingaction units in line with situational characteristicsand adapting the enactment of the actionunits to particular contextual conditions in whichthe action takes place. Increased flexible adaptabilitymay reach a level where its implementation cannotdepend any more on tuning-in with the situationalconstraints as one goes along, following the trial–and–error method. The next major evolutionarystep in this respect consists in the emergence of thebehavioral program which completely changed thescene of action by introducing from above an organizationalprinciple that takes control over actionfrom “the top, organizing the enactment in terms ofan overall scheme”. This change may have occurredat the level of worms. The program starts to play arole in the selection of the appropriate action unitsfor the behavior and in the way habits are embeddedwithin it (GALLISTEL 1980; MILLER/GALANTER/PRIBRAM1960).Within the sphere of behavioral programs a furtherevolution takes place whereby learning gets anincreased role and contribution. The primary programsare innate ones, such as reflexes. These programsbecome slowly more flexible and adaptable.For example, conditioning brings about the bindingof these primary programs to new signals so thatthey come to be evoked by learned stimuli in thecontext of new situations. These and other processesof learning invade, as it were, the innate schemes,enlarging and transforming them in different ways.Moreover, the innate programs are gradually amplifiedby newly evolving programs that are increasinglybased on learning. Thus, the next stage consistsof programs that are partly innate and partlylearned, such as instincts, behaviors correspondingto defense mechanisms, language and social behaviors.The components due to learning become graduallymore salient and push the innately-given componentsinto the background so that they becomebarely perceptible. For example, in an instinct thelearning-based ‘appetitive behavior’ is distinctly differentiatedfrom the end-stage behavior which representsthe more innate components. However, in adefense mechanism behavior such as displacement,the innate element is represented not in the actualmanifest behavior but in its underlying cause orgoal. In different social behaviors, such as courting,threatening or impressing others the innate elementis even more covert (HINDE 1970, pp370–382; MORRIS1969).The next evolutionary stage is characterized bythe emergence of behavioral programs that arepurely learned, such as, driving a car or using theinternet. It is likely that even here there are somebarely perceptible but identifiable innate elementsthat have stealthily invaded the scheme (e.g., pushingbuttons, turning wheels). The next evolutionarystage consists in giving up ready-made programs andrelying increasingly on ad-hoc programs newly createdin line with the present circumstances. Thus,program retrieval is gradually replaced by planning.A series of studies on the development of planningshowed that the major developments occur in regardto information gathering (focusing on items of informationnecessary for proceeding smoothly withthe action), information organization (for example,in terms of if–then considerations, or defining domainsof information), and plan structuring (chronologicalordering, number of steps, major and alternateplans, etc.). The development of planningand acquisition of planning skills are promoted bymotivation and increased awareness of needs (KRE-ITLER/KREITLER 1987a–c).The advantage of the ad-hoc programs is their perfectfit to the situation, their disadvantage—thatthey have to be created under the pressure of ongoingbehavior. The task is however made easier in viewof the more advanced development of programs forthe formation of programs. These programs makeuse of elements accumulated in the course of previousdevelopment, from the bottom, as it were, but assoon as they emerge they start controlling the processesfrom the top, as it were. However, in this fieldtoo, the more developed manifestations do not completelyreplace the lower-level products, which maystill be adequate for handling specific situationsmore adequately than higher-level behaviors (seethe principle of “preservation of the fittest” in “TheCO Theory: General Characteristics”).The described development of programs emphasizedtheir increased shaping by learning. The developmentof programs may also be examined from thecomplementary viewpoint of the changes they undergo.First, it is evident that behavioral programsincrease in number, so that the organism has programsadequate for an increasing number of differ-Evolution and Cognition ❘ 90 ❘ 2001, Vol. 7, No. 1

An Evolutionary Perspective on Cognitive Orientationent situations, and also more programs for the sameor similar situations from which one can select theadequate one. Secondly, behavioral programs growin complexity in the sense that they get more hierarchicallyorganized with an increasing number oflevels. This involves not only an increasing numberof units encompassed within the program but alsoconsideration of special conditions, alternatives and‘if then’ eventualities. With time, following differentiationprocesses in complex systems (RIEDL 1997,p152), a differentiation occurs between the levelsthat deal with the overall strategy of action (viz. ‘operationalprogram’) and the levels that deal mainlywith the operational details involved in the specificationsof the enactment, mostly in terms of physiologicalprocesses (viz. ‘program scheme’). The increasedcomplexity of organization enablesactivating not only the program as a whole but partsof programs, or shaping new programs by integratingparts from other programs. At a higher level ofdevelopment, behavioral programs themselves undergofurther evolution in that they are organized,for example in terms of similarity or relevance forspecific situations. Eventually this leads to the thirdimportant development which consists in formingprinciples for the construction of programs. Thisfrees performance to a certain extent from relianceon ready-made programs and introduces the optionof planning, with all the freedom and control it involves.Specific developmental trends. The present sectionis devoted to highlighting diverse evolutionarytrends affecting the phases defined by the CO modelas intervening between input and output. Since theCO model is a cognitive model of motivation, tracingthe evolutionary trends involved in the phasesof the model would highlight more specifically thanbefore the evolutionary development of the cognitiveguidance of motivation.In the first phase, the primary actions would beactivities dependent completely on internal stimuli,if indeed one may consider the internal perturbationsof disturbed homeostasis as stimuli. In the bestcase, they are equivalents or precursors of stimuli.The differentiation between the motivation and actionstarts at this phase. WIMMER (1995) provides amost original and detailed reconstruction of the differentiationin evolutionary perspective. At the lowestlevel we start with the primary action unit, whoseactivity is regulated by internal stimuli. These internalstimuli (the earliest precursor of emotion), reflectboth the inner state of the organism (homeostasis or‘disturbance’) and the outer state (they are the ‘perception’of the environment mediated by the organism’shomeostasis, p43). Moreover, they contain theroots of motivation in the form of ‘expectation’ ofinformation and of response, in the form of rigidmodes of reaction (RIEDL 1980). The evolutionary developmentshows that originally the locomotory activityof kinesis, for example, in taxis behaviors, isrigid, automatically regulated and appears to be undirected.At higher levels of taxis behaviors it is replacedby a type of action which shows the earmarksof being directly determined by the impinging stimulus(LORENZ 1981), reflecting the impact of the organism’sinternal state. This is probably the earliestprecursor of goal-directedness. Notably, it correspondsto ‘intentionality’, that has been identifiedas a basic characteristic of behavior, at least from thelevel of organisms evidencing directed movementsor taxes (DIAMOND 1991; FREEMAN 1999, p41). Themanifestation of intentionality marks the emergenceof motivation as distinct from the action perse. WIMMER (1995, p43) cites in this context the exampleof the behavior of the turbellarian wormswhose movement toward a feeding site depends ontheir internal state: the hungry worm will move towardthe site, whereas a satiated worm will not.Hence, WIMMER concludes that it is the differentiationof the internal state (e.g., hungry, satiated)which leads to the development of more differenttypes of perceptions (sensitivity to different constellationsof stimuli) and of motoric reactions (approachor avoidance reactions). With the developmentof perception, it becomes possible for externalstimuli to initiate action. In terms of the CO theory,it is the motivational disposition which drives developmentof the grasp of reality and of the performancecomponent.Further evolutionary trends that may be relevantin regard to the first phase of the CO theory dependon the development of meaning elaboration andfeedback evaluation. At the lowest level, the inputwhich initiates action has meaning which is minimal(it consists perhaps of only one meaning value)and is couched in actional terms. Hence, we call it‘meaning action’. Its manifestation is precisely inevoking action, say, a reflex. It consists in identifyingthe input as a signal for a specific action, namely, inevoking action and it has no independent existenceoutside action. This applies to stimuli that evoke reflexesor primitive defensive reactions. ‘Releasers’ (orsign stimuli) are at a slightly higher evolutionarylevel insofar as they act as triggers for instincts,which are chains of actions manifesting some degreeEvolution and Cognition ❘ 91 ❘ 2001, Vol. 7, No. 1

Shulamith Kreitlerof plasticity. At higher evolutionary levels we mayfind meanings that are not identical with a specificaction, for example, a meaning that is evaluated asirrelevant and hence is dismissed without evokingany response, or a meaning that is evaluated as requiringa more complex type of behavior and henceis transferred to the next phase.Similarly, also feedback evaluation is subject todevelopment. Basically, feedback evaluation dependson applying cognitive processes and meaning.Thus, evaluating the outcome of action requiresmatching the outcome with the intent, adjustingboth so that they can be compared, judging the degreeof similarity, and evaluating the result also interms of the current situation and its potentialities.This description fits higher evolutionary levels. Onlower levels the precursor of feedback would consistsimply in recurrent enactment of the action if thedisturbance exists, regardless of whether the actionhad been enacted before and what the goal may havebeen. The action is simply repeated until the goal isattained or a ‘stop circumstance’ sets in (e.g., the organismis distracted, the organism stops because offatigue, the action is inhibited by an internal inhibitingmechanism). In this context it is relevant to citethe observations about infants 5–7 months old tryingagain and again without success to retrieve a toythey had and which was taken away from them (DI-AMOND 1991). Hence, at low evolutionary levels, itmay happen that an action does not attain the goalin any sense, yet it is not followed by another action.At a later evolutionary level, feedback already in thecase of instincts is based on a rigid type of matchingthe actual activity or outcome with the original ‘expectation’or ‘optimal’ pattern. This geneticallybasedyardstick (WIMMER 1995, p45) or ‘inbornteacher’ (LORENZ 1981, p299) is amplified by moreflexible and extended forms of feedback, based onlearned programs stored in memory. One type consistsin evaluating the correspondence between actualaction and stored pattern (KLIX 1980, p102), anotherin estimating the accuracy of reaching the goal(ibid, p104). WIMMER (1995) emphasizes that the latteris anchored in the brain centers of reward andpunishment, so that it uses elementary emotionalreactions as criteria for success and failure of actions.The development of these feedback mechanisms hashelped to dissociate behaviors from specific stimuliand has promoted the occurrence of behaviors incontexts other than those in which they have beenacquired (e.g., conditioned actions) (ibid, pp45–46).With further developments the feedback mechanismsbecome more flexible and expanded, consistingin examining the outcome situation more extensivelyand comparing it in several senses with theoriginal goal(s). At this level, the feedback operationalready includes the vestiges of preparing the nextaction designed to handle the original goal in viewof the experienced failure of the previous action.In the first phase of the CO theory there are at leasttwo occasions for feedback: after the execution of thenon-conditioned or conditioned response, and afterevocation of the orienting reflex. The previous observationsabout the evolution of feedback indicatethat it requires higher evolutionary levels to diagnosefailure in these actions, and in case of failure, itrequires even higher levels to transfer the handlingof the situation to more advanced phases (viz. tomolar action).In line with the CO theory, the second phase isfocused on a more extended elaboration of themeaning of the input and the situation in which itis embedded. This phase is designed to examine, onthe bases of the meanings which emerge, whetheraction is at all indicated. This phase bears evidencefor an advanced evolutionary level. In addition to anextended meaning system, it presupposes the abilitynot to act. Studies with infants suggest that this abilityis based on a more basic ability, which is not toreact in line with innate and conditioned prepotentresponse tendencies. The latter gives one the possibilityof reacting in line with internal tendencies.Extending the potential of inhibiting actions toplanned, considered and possible actions, largelywidens the scope of intentionality and control. Bothabilities of inhibiting action are probably not possiblewithout the frontal cortex (DIAMOND 1991).There seem to be two major evolutionary trendscharacteristic of the second phase. The first and mostobvious one concerns meaning elaboration. Thebroader and more comprehensive the meaning systembecomes, the more extensive gets the meaningelaboration in this context, involving more aspectsof meaning and both types of meaning—the interpersonally-sharedand the personal, in the form ofmeanings as well as beliefs. Correspondingly, the examinationof whether the individual is involved inthe situation is based on an ever-broader basis. Thesecond aspect that undergoes evolution is the criteriafor acting or not acting. If on more primary levelsthe decision follows criteria, such as direct contributionto survival, evolution brings into play furthercriteria, such as potential for promoting the attainmentof some personal goal, the possibility to defendone’s values or to enhance one’s sense of self-identity,etc.Evolution and Cognition ❘ 92 ❘ 2001, Vol. 7, No. 1

An Evolutionary Perspective on Cognitive OrientationThe third phase is focused on determining the directionalityof behavior. One important evolutionarytrend concerns the emergence of underlyingmeanings. Their range and depth increase so thatthey come to represent the matrix of meanings outof which the behavioral intent can be shaped. Thishowever raises a dilemma. On the one hand, it isdesirable for shaping the behavioral directionality toevolve a broad and deep set of underlying meanings.Yet, on the other hand, awareness of this completeset would render action difficult insofar as it constitutesa heavy memory load and uses large mentalresources of consciousness. The solution to the dilemmacould be to develop the whole matrix but tokeep it unconscious. Possibly, the demands ofsmooth action and preservation of mental resourcesmay have promoted development of the unconsciousness.Other interesting lines of development concernthe formation of the CO cluster. Most important ofall is the issue of the number of belief types that arenecessary for creating a CO cluster. At the lowestlevel, as a rule only one belief type dominates thescene, most probably goals (= representing approachor avoidance, “I want” or “I don’t want”). The nextbelief type to make its contribution to behavioraldirectionality is rules and norms (= representingconstraints of right and wrong, allowed or forbiddenand later also of utility, desirability etc.). Each coulddominate the scene in itself. An important developmentoccurs however when beliefs about goals andbeliefs about rules and norms do not point in thesame direction, or when they clash. A study withchildren showed that it is only under these conditionsthat the other belief types general beliefs andself beliefs enter the scene and begin to make a differencein predicting the child’s behavior (MAROM1978).There are other pairs of belief types that couldform inconsistencies of consequence for the developmentof the CO cluster. One is the inconsistencybetween beliefs about goals and general beliefs,which occurs when prevailing circumstances do notsupport the attainment of a goal. This may havebeen the case for the apes studied by DELGADO (1974)whose goal to express aggression was not matchedby the information that the target or would-be victimis a strong ape who would not easily succumb.Another example is the inconsistency between beliefsabout norms and beliefs about self, which occursfor example when one becomes aware of notliving up to the requirements. Again, there is the gapbetween beliefs about norms and general beliefs thatmay occur when reality does not live up to normativerequirements. A last example concerns the gapbetween beliefs about self and general beliefs, whichoccurs when informations about oneself and othersare inconsistent. In each of these cases the drive forsolution is based on bringing into consideration additionalinformation, in the form of further types ofbeliefs. Hence, the drive behind the formation of aCO cluster representing the overall directionalityimplied by four types of beliefs is the inconsistenciesin directionality between any two pairs of belieftypes. Studies showed that when individuals functionunder difficult conditions, such as the impactof drugs or medication, drunkenness, fatigue, or intenseemotions, they tend to form CO clusters basedon fewer than all four types of beliefs. The inconsistenciesbetween belief types serve to mobilize for theCO cluster more beliefs. This enables forming a COcluster if there are enough beliefs supporting thecourse of action rather than if all beliefs or all belieftypes support it.The increase in the number of belief types onwhich the CO cluster is based is probably attendedalso by other developments in the formation of theCO clusters. We will mention in particular four thatare of special importance. The first concerns the anchorof the CO cluster. Originally, the anchor musthave been a goal belief. In the course of development,other belief types could assume this role, inline with the characteristics of the behavior consideredfor enactment. For example, general beliefswhen the behavior is conformity, norm beliefs whenthe behavior is helping others, beliefs about selfwhen the behavior is assertiveness. Secondly, alsothe salience of the belief type in the formation of theCO cluster developed from a fixed order to an orderthat varies in line with the behavior and circumstances,and later possibly also personality. Thirdly,the range of implications of the beliefs that exerts animpact on the processes of belief interactions andclustering undergoes change. The primary and mostimportant kind of implication is orientativeness,namely, the degree to which the belief supports thebehavior. Development sets in operation furthertypes of implications (e.g., morality, utility, emotionalimpact), which though initially they play asecondary role, they increasingly contribute to theformation of the CO cluster and to shaping the directionalityof behavior. Finally, development occursalso in regard to the amount of consistencyamong the orientativeness of the beliefs required forthe formation of a CO cluster and the consequentemergence of the behavioral intent. On the lowestEvolution and Cognition ❘ 93 ❘ 2001, Vol. 7, No. 1

Shulamith Kreitlerdevelopmental level of the CO cluster the requirementis for perfect or almost perfect consistency inthe orientativeness of the beliefs. All relevant beliefsmust support the behavior; otherwise no CO clusteris formed. With evolution the standards of requirementsare loosened, until the stage is reached whenconsistency is required only within the CO clusterbut not among all beliefs relevant for the behavior.Thus, the primary question “Do all beliefs supportthe behavior?” is replaced by the question “Is theresufficient support for the behavior?” This developmentenables the accumulation or generation of beliefsof all kinds, without prior consideration of thebehaviors which they could support or not. Subsequently,further beliefs are generated or acquiredwithout regard for their actual or potential contradictionwith the already existing beliefs. Removal ofthe constraints on belief accumulation enables agreat enrichment of cognitive contents. The impacton cognitive development is remarkable.Furthermore, it can be expected that also the criterionfor ‘sufficient support’ becomes more flexiblewith time so that assessing sufficiency is based notonly on the number of involved beliefs but also onadditional aspects, such as the situational context,kind of behavior, and meaningfulness of the beliefsThe fourth phase is focused on behavior performance.The major evolutionary developments referto the activation, adaptation and enactment of thebehavioral programs.Activation involves identifying and evoking theadequate program. On the lower evolutionary levels,each behavioral program is related to a specific stimulusor range of stimuli. Under these conditions,identifying the adequate program consists in retrievingit as a response when the proper stimulus appears.Identifying and retrieving are not yet differentiated.When behavioral programs undergodissociation from the releasing stimuli, retrieval becomesa matter of search in memory for the adequateprogram. The selection process involves focusing ona pattern of behavior that is marked as ‘adequate’and searching in memory for a pattern that matchesthe constructed concept to a satisfying or ‘satisficing’degree. The ‘adequate’ program is initially searchedfrom among those stored in memory, and in laterevolutionary levels may be shaped or constructed adhoc.Adaptation of programs involves an interactionbetween the program and the available circumstancesthat results in transforming the program sothat it increasingly matches the circumstances. Developmentbrings about an ever-increasing adequacyin the fit between the behavioral program andthe context in which the behavior takes place. Accordingly,adaptation consists in a gradual departurefrom rigid action units or sequences in favor of flexible,changing, and variable behaviors. The differencebecomes most apparent when one comparesthe relative flexibility in the operation of instincts,especially in the appetitive stage, with the rigidity inthe operation of reflexes. It is as if the stored behavioralprogram becomes a paradigm for a set of variationsthat assume unique shapes when set in operationin reality.Enactment of programs involves an interactionbetween the program and the emerging reality contingentupon setting it in operation. Initially, enactmentis examined only in terms of the end-point(i.e., has the action been terminated or not). Laterdevelopments introduce control over intermediateand primary stages of enactment. Control consistsfirst in determining whether the action takes placeor not, but it evolves into a more complex operationwhich includes also evaluation of the operation itselfand its outcomes and the possibility of modifyingit, if necessary. Thus, control assumes graduallyan autonomous status from the performance itself.Thereby it becomes increasingly more cognitive andenables the introduction of additional criteria forthe implementation of control. On the lower evolutionarylevels the main criteria for evaluating ongoingperformance is the occurrence of the activity itself(namely, whether the action occurs and runs toits end), implemented by utility (the pragmatic evaluationin terms of whether the straightforward goalwas attained). Evolution enables the introduction offurther criteria, mainly whether the action and/orthe goal attainment are emotionally satisfying,whether they conform to social desirability, whetherthey are useful also in promoting the attainment ofother goals, whether they conform to standards ofreasonableness or rationality, whether they were sufficientlyeconomical in the use of resources (such as,time, money, energy), whether they satisfy aestheticrequirements, etc. Notably, the application of considerationsof this kind for evaluating the performanceand its outcomes serve to enhance the integrationof behavior into the system of personality.ConclusionsOne major conclusion of this paper is that the cognitiveguidance of behavior is an evolutionary product,reflecting multiple developmental trends.Major among these are the differentiation betweenEvolution and Cognition ❘ 94 ❘ 2001, Vol. 7, No. 1

An Evolutionary Perspective on Cognitive Orientationmotivation and action, the development of meaning,and the development of action. In the course ofevolution, motivation unfolds in a manner thatenables its increasing suffusion and consequentshaping by cognitive contents and processesanchored in the components and dynamics ofmeaning. Meaning assignment processes are, however,subjected to the overall motivational frameworkthat determines the purpose, manner andproducts of the meaning elaboration. This meansthat cognitive elaboration in the context of motivationdiffers in contents and processes from cognitiveelaboration in other contexts. Thus, in contrastto ‘pure’ problem solving and other contexts inwhich cognition is operative, meaning elaborationin the motivational framework capitalizes on thefollowing: a specific balancing of interpersonallysharedand personal–subjectivemeaning; the use ofmeaning action; treatingmeaning units as beliefs; activatingfour types of beliefs;unraveling underlying meaningsof the inputs to a specificlevel of depth; focusing onAuthor’s addressShulamith Kreitler, Department of Psychology,Tel-Aviv University, Tel-Aviv 69978,Israel, Tel: +972-3-5227185, Fax: +972-3-5225371. E-mail: krit@netvision.net.ilorientativeness of beliefs as a particular aspect of themeaning of beliefs; and clustering of beliefs into aCO cluster expressible in terms of a goal-directedvectorial disposition.In the course of evolution, action develops in thefollowing ways: organizing action units; freeingthem from original inputs or inputs related to theiracquisition; increasing complexity of organization;development of behavioral programs, amplifyingthe innate programs by partially-learned and fullylearnedprograms; amplifying and replacing storedprograms by planning, which produces ad-hocplans, some of which may be stored for future use;cognitive control of the enactment of programs, includingthe evaluation of action outcomes.In view of the above, it is justified to conclude thatthe cognitive guidance of behavior underscores andrenders explicit an importantmanifestation of the contributionof cognition to behaviorand to survival. Thus, it highlightscognition’s unique rolein terms of evolution, as it isshaped by evolution, for evolution.ReferencesAjzen, I. (1985) From intentions to actions: A theory ofplanned behavior. In: Kuhl, I./Beckman, J. (eds) ActionControl: From Cognition to Behavior. Springer: Berlin, pp.11–40.Bachar, E./Latzer, Y./Kreitler, S./Berry, E. M. (1999) Empiricalcomparison of two psychological therapies: Self psychologyand cognitive orientation in treatment of anorexia and bulimia.Journal of Psychotherapy Practice and Research 8:115–128.Becker, M. H. (1974) The health belief model and personalbehavior. Health Education Monographs 2: 324–508.Campbell, D. T. (1974) Evolutionary epistemology. In:Schlipp, P. A. (ed) The Philosophy of Karl Popper. OpenCourt: LaSalle IL, pp. 413–463.Corey, S. M. (1937) Professed attitudes and actual behavior.Journal of Educational Psychology 28: 271–280.Cassirer, E. (1953) An Essay on Man: An Introduction to aPhilosophy of Human Culture. Doubleday, Anchor Books:Garden City NY.DeKay, W. T./Shackelford, T. K. (2000) Toward an evolutionaryapproach to social cognition. Evolution and Cognition6: 185–195.Delgado, J. M. R. (1971) Physical Control of the Mind: Towarda Psycho-Civilized Society. Harper: New York.Diamond, A. (1991) Neuropsychological insights into themeaning of object concept development. In : Carey, S./Gelman, R. (eds) The Epigenesis of Mind: Essays on Biologyand Cognition. Erlbaum: Hillsdale NJ, pp. 67–110.Freeman, W. J. (1999) How Brains Make up their Minds.Phoenix, Orion Books: London.Gallistel, C. R. (1980) The Organization of Action: A New Synthesis.Erlbaum: Hillsdale NJ.Gallistel, C. R. (1990) The Organization of Learning. A BradfordBook, MIT Press: Cambridge MA.Hinde, R. A. (1970) Animal Behavior: A Synthesis of Ethologyand Comparative Psychology. 2nd edition. McGraw-Hill:New York.Klix, F. (1980) Erwachendes Denken. Eine Entwicklungsgeschichteder menschlichen Intelligenz. VEB DeutscherVerlag der Wissenschaften: Berlin.Ková č, L. (2000) Fundamental principles of cognitive biology.Evolution and Cognition 6: 51–69.Kreitler, S. (1965) Symbolschöpfung und Symbolerfassung:Eine experimentalpsychologische Studie [The Creationand Perception of Symbols: An Experimental PsychologicalStudy]. Reinhardt: Basel, München.Kreitler, S. (1999) Consciousness and meaning. In: Singer, J.A./Salovey, P. (eds) At Play in the Fields of Consciousness.Erlbaum: Mahwah NJ, pp. 175–206.Kreitler, S. (2002) The cognitive guidance of behavior. In:Jost, E. et al. (eds), The Yin and Yang of Social Cognition:Perspectives on the Social Psychology of Thought Systems.Erlbaum: Mahwah NJ.Kreitler, S./Chaitchik, S./Kreitler, H./Weissler, K. (1994)Who will attend tests for the early detection of breast cancer?Psychology and Health 9: 463–483.Evolution and Cognition ❘ 95 ❘ 2001, Vol. 7, No. 1

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Chris MacDonaldEvolutionary EthicsValue, Psychology, Strategy and Conventions1. IntroductionThe purpose of this paper 1is to present an exampleof what moral theorywould look like if it tookevolutionary theory seriously.In it, I strive toavoid the errors to whichso many previous attemptshave succumbed.If I succeed in doing so, itis because of my intentmerely to show how DAR-WINIAN thinking improvesthe empirical basisupon which we constructmoral theories. At nopoint do I try to derivefoundational moral valueseither from nature inthe broadest sense orfrom the process of evolutionitself. My project isbased on the widely-held assumption that moral decisionmaking requires both values and facts. Knowinga bit about evolution can help make it clear whatthe relevant facts are. Here, I do not intend to discussbiological evolution in particular, but evolution inthe broadest sense. Evolution takes place whereverthere is variation, replication, mutation, and differentialsurvival according to inherited traits. Thusevolutionary theory applies to (at least) biologicalorganisms, certain computer programs, cultures,and ideas.At the heart of my project lies the claim that moralstandards are the products of biological and culturalevolution. Biological evolution has given us brainscapable of certain cognitive tasks, and capable ofmanifesting certain emotions. On top of this biopsychologicalsubstrate, cultural evolution has builtAbstractThe purpose of this paper is to present an example ofwhat moral theory would look like if it took evolutionarytheory seriously. First, I examine briefly the implicationsthat accepting evolutionary theory has forvalue theory. Secondly, I suggest that an evolutionaryperspective can shed new light on moral psychology.Finally, I suggest that an evolutionary perspectiveclears the way for a moral perspective that focuses onthe function and development of HUMEAN social conventions.I strive to avoid the errors to which so manyprevious attempts at grounding ethics in evolutionhave succumbed. If I succeed in doing so, it is becausemy intent is merely to show how DARWINIAN thinkingimproves the empirical basis upon which we constructmoral theories. At no point do I try to derive foundationalmoral values either from nature in the broadestsense or from the process of evolution itself.Key wordsEthics, convention, psychology, function, game theory.Evolution and Cognition ❘ 98 ❘ 2001, Vol. 7, No. 1a wide range of particularnorms and conventions.The field of ethics can beinterpreted as being devotedto influencing thecourse that the evolutionof these norms and conventionstakes.The basic claim of thispaper is that attention toevolutionary theoryshould have two mainconsequences for moraltheory. First, attention toevolutionary theory (as itapplies to ethics) shoulddraw attention to the fieldof evolutionary psychology,as a way of enrichingour understanding ofmoral psychology. Secondly,it should draw ourattention to the study ofsocial conventions, understoodas regularities of behaviour, sustained by ashared interest in coordination and an expectationthat others will cooperate.Why social conventions? To begin with, asGAUTHIER (1991, p18) argues, we no longer acceptthe world-view upon which absolutist conceptionsof morality depend. 2 Thus we no longer have accessto plausible notions of inherent and intrinsic valueupon which to base ideal moral codes. 3 An evolutionaryview can only be founded on a subjectiveand relative theory of value. 4 Such a value-theoryleads us to ask what moral rules—what social structures—wouldaid in the pursuit of whatever it is thatwe value. 5 What moral rules have we agreed upon,and what moral rules could we agree upon, in pursuitof what we want? That is, the evolutionary perspectivemust lead us to focus on the extent to which

Evolutionary Ethicsmorality is a useful human construct, a necessaryartifice.Our second reason for paying to attention to conventionsis that an evolutionary perspective forcesus to take an explicitly historical view of our moralstandards. Much of moral theory is weakened by theextent to which it fails to appreciate the importanceof considering moral agents as historically and sociallysituated. Moral theory mistakenly assumes, forthe most part, that agents (or societies) are in a positionof radical choice vis-à-vis their moral principles,or more generally, their social arrangements. Attentionto social conventions—actual, existing, workingbits of cooperative behaviour—is a way of takingseriously the moral work already done by culturalevolution.2. ValueWe begin by looking at the implications of evolutionarytheory for value theory. The most fundamentalshift that an evolutionary perspectivebrings about—or should bring about—in ourunderstanding of value lies in the undermining ofwhat RACHELS (1990) calls the ‘Human Dignity Thesis’.According to the Human Dignity Thesis,humans have special value not shared by nonhumannature; also according to the Thesis,humans have this special value just by beinghuman, which implies that all human lives havethis value to some extent or another. Both of thesevalue claims have wide-reaching implications forethics. But, according to RACHELS, an evolutionaryperspective casts doubt upon the two claims that hecalls the ‘twin pillars’ holding up the Human DignityThesis. First, the acceptance of evolutionarytheory renders unnecessary (as an explanatoryhypothesis) the Western religious claim thathumans were made in the image of God. Secondly,in situating humans on a developmental spectrumof cognitive capacities, as creatures with capacitiesdifferent in degree but not in kind from other animals,evolutionary thinking undermines the secularclaim that humans are of special value onaccount of their unique rationality.Although the theory to which, I argue, the evolutionaryperspective drives us is the theory of valueassumed by our most well-developed social science—economics—itis also a theory of value that isat odds with most modern moral theories. A full theoryof value is beyond the scope of this paper, butthe basics—borrowed from David GAUTHIER’s accountof value—can be stated briefly. The value theoryto which an evolutionary perspective drives us isone that holds that all value is dependent upon theaffective relationships of the individual, and is thereforeboth subjective (as opposed to objective) andrelative (rather than absolute).First, that value is subjective means that the ascriptionof value is taken to be attitudinal, not observational(GAUTHIER 1986, p25). That is, a thing’sbeing valuable depends upon some sentient beinghaving a positive attitude toward it. GAUTHIER(1986, p46) puts the point quite simply: “Value… isa measure of preference”. Thomas HOBBES is perhapsthe classic exponent of the subjective view of value.HOBBES (1985, p24) writes, “For these words ofGood, Evill… are ever used with relation to the personthat useth them: There being nothing simplyand absolutely so; nor any common Rule of Goodand Evill, to be taken from the nature of the objectsthemselves; but from the Person of the man”. Forthe value subjectivist then, nothing is ever valuablesimpliciter; it is always valuable to someone. Accordingto GAUTHIER (1986, p56), we should decide“whether any adequate account of our experienceor environment must refer to objective value” byconsidering “whether any reference to objectivevalue occurs in the best explanation we can providefor our actions and choices” (emphasis original).The value objectivist, according to (GAUTHIER 1986,p55) “…holds that any adequate account of our experienceor our environment must refer to value orto the valuable as being independent of sentient beingsand their affections”. GAUTHIER suggests insteadthat “…the only serious candidate for an explanatoryschema for human action is… [oneaccording to which] choice maximizes preferencefulfilment given belief. Objective value plays no rolein this account” (p56). GAUTHIER continues: “Objectivevalue, like phlogiston, is an unnecessary part ofour explanatory apparatus, and as such is to beshaved from the face of the universe by OCKHAM’srazor”.I would argue that objective value died (or at anyrate, should have died) in 1859, with the publicationof DARWIN’s The Origin of Species. DARWINIAN theorypulls down the two supporting pillars of the HumanDignity Thesis. Under the scientific world-view withwhich we are left, objective value is simply an unnecessaryexplanatory hypothesis.On the theory of value to which evolutionarythinking drives us, not only is value to be taken tobe subjective, but it is also relative. On the relativeconception of value, according to GAUTHIER (1986,p50), “…each person has his own good (and bad),Evolution and Cognition ❘ 99 ❘ 2001, Vol. 7, No. 1

Chris MacDonaldand… the goods of different persons are not parts ofa single, overall good”. Note that to hold that valueis individualistic in this sense does not imply eitherethical or psychological egoism; that is, it does notimply that individuals care only about their ownwell-being. As GAUTHIER puts it, “A state of affairsthat is good for a single individual is then good fromthe standpoint of each person who prefers or wouldprefer the enhancement of that individual’s well-being”(p51). (Nor does understanding value as relativenecessarily lead to ethical relativism. Moral rulesmay be shared, even if we differ in our attributionsof value.) In opposition to the relative conception ofvalue is the absolutist conception, which holds, accordingto GAUTHIER that “…values are the same forall persons, or for all sentient beings” (p50). The conceptionof value as relative fits nicely with the competitiveframework of evolutionary theory, accordingto which individuals may value the welfare ofothers, but only contingently.As noted above, most modern moral theories areat odds with this subjective and relative theory ofvalue. Utilitarianism, for example, makes the claimthat the happiness of any given individual is a goodthing simpliciter; that is, it is to be regarded as a goodby every rational being. Thus for Utilitarians, valueis subjective but also absolute. KANTIANS, for theirpart, hold that the respect due to all rational agentsfinds its source in their inherent value. The KANTIANtheory of value must then be both objective (since itmakes no mention of anyone’s preferences or affectivestates) and absolute (since it holds rationalagents to have value for everyone). For a fuller defenceof the individualist theory that takes value insteadto be both subjective and relative, I refer thereader to GAUTHIER (1986)’s Chapter II section 4.3. Evolutionary PsychologyHow can we tell whether a given rule or standard isconsistent with our moral psychology? Empiricalinvestigation—trial and error—is one way. 6 But evolutionaryconsiderations can be useful in guidingour empirical investigations. Thus one set of methodsthat can be used to study the psychologicalmechanisms behind our moral behaviour is that ofevolutionary psychology. 7Evolutionary psychology is predicated on the ideathat, contrary to the beliefs of behaviourists such asPAVLOV and SKINNER, the human mind is anythingbut a blank slate. Rather than seeing it as a generalpurposecomputational device, evolutionary psychologistssee the mind as a collection of specialpurposealgorithms, each designed (in the metaphoricalsense 8 ) to solve a particular problem facedduring our evolutionary history.According to evolutionary psychologists, evolutionhas given us certain cognitive capacities (suchas the ability to detect aggression in others, or theability to remember wrongs) and emotional dispositions(such as the capacity for sympathy and spite)that factor significantly in our moral lives. Investigationof these capacities and dispositions can be informedby evolutionary theory. Knowledge of thesorts of problems that the human brain evolved tosolve can lead to hypotheses concerning what sortsof particular cognitive mechanisms and predispositionswe can expect to be included in our psychologicalmakeup. Such hypotheses can be tested againstanthropological data and against psychological evaluationof test subjects. We can then evaluate particularmoral standards in terms of the degree to whichthey are compatible with known capacities and dispositions.In applying this test, I am proposing theuse of an evolutionary perspective to investigate humannature. There is nothing novel about wantingto look at human nature in designing moral systems.The idea here is that our shared nature places limitson what we can learn to do—on how we can learn tobehave. The novelty, of course (at least for philosophers)lies in looking to evolutionary theory for abetter understanding of that nature.3.1 Cognitive capacitiesWhat sorts of limits on moral behaviour might beseen in our shared, evolved nature? As one example,COSMIDES and TOOBY (1992) find compelling evidencefor a special human aptitude for detectingcheating in situations of social exchange. Similarstudies might just as easily find that humans are particularlyinept at a particular kind of reasoning.Given such knowledge, it would seem foolish (froman instrumentalist point of view) to propose a moralstandard that ignored our aptitudes while relyingheavily on forms of reasoning at which we generallydo badly.For example, it might one day be discovered thathumans lack the aptitude for categorical reasoningof the sort demanded of us by KANT. That is, it mightturn out, upon empirical investigation, that humansare relatively inept at figuring out whether, ina particular case, the maxim by which one acts isone “which [one] can at the same time will that itshould become a universal law” (KANT 1959, p39). Ifsuch a hypothesis about our cognitive capacitiesEvolution and Cognition ❘ 100 ❘ 2001, Vol. 7, No. 1

Evolutionary Ethicswere validated, I take it that this would be a harsh—though perhaps not fatal—criticism of that aspect ofKANTIAN ethics (at least from the point of view ofanyone who wants to propose useful moral rules).3.2 Emotional predispositionsNext, let us look at the emotional dispositions withwhich our evolutionary past has left us. Since‘ought’ implies ‘can’, it would seem foolish to havea moral rule which demanded behaviours of whichwe were emotionally (or otherwise) incapable, ornearly so. A case in point: all available evidencepoints to a general human predisposition to givepreferential treatment to friends and family. 9 Notonly is this tendency apparently universal amonghuman cultures, it also shows up (under the title‘kin altruism’) in a large number of animal species.Flying in the face of this apparently universalhuman tendency is the Benthamite claim that“each is to count for one and none for more thanone”. This Utilitarian standard requires preciselythat my brother not count for more than a strangerin my moral deliberations. I submit that if we wantour moral rules to be respected and adhered to, theyat least ought not to demand that we forgo stronglyingrained behaviours unnecessarily.It may be objected that humans display all sortsof nasty predispositions that ought not to be excusedsimply because they are ‘natural’. I agree, and notsimply because I want to avoid committing the naturalisticfallacy. For example, regardless of how ingrainedour tendencies toward aggression are, weneed to place moral restrictions on aggressive behaviour.Yet in the case both of aggression and of favouritism,the crucial point is to limit socially destructivebehaviour. Thus, while our sense of justicedemands that we limit favouritism in cases such asthe awarding of government contracts, there will bemany situations in which preferential treatment offamily and friends seems not just forgivable, but appropriate.Likewise, while it is socially imperative toprohibit the killing of innocents, it seems unnecessaryto forbid certain other displays of aggression, forexample organized sports.Further, consider the fact that there may be anumber of moral rules that could mediate any givensocial situation. For example, we might determinesuitable patterns of economic distribution in one ofthree ways:1) by assigning to each individual the right tosome level of quality of life (i.e., a form of Deontology);2) by claiming that a certain pattern of distributionmaximized the net benefits to society as a whole(i.e., Utilitarianism); or3) by claiming that a certain pattern of distributionwould be agreed to in a (hypothetical) fair bargainingsituation (i.e., Contractarianism).Each of these rules is in some sense plausible, andeach might even point to a similar outcome. So instrumentally,it might not much matter which onewe choose. I suggest that if one of them turns out tobe psychologically implausible, this can help uschoose among these competing moral standards.4. Game Theory and Social ConventionsWhat can we say, from an evolutionary perspective,about the particular rules that will work well asmoral rules? The evolutionary perspective leads usto be interested in the function of moral rules; ourfocus should be on determining how different ruleswill function within a social milieu. Game theorycomprises a set of tools that can be used to study,from a theoretical point of view, the social mechanismsof morality. We can usefully think of a moralrule as a strategy for solving a particular social-coordinationproblem. Game theory, as the study of strategicdecision–making, 10 can illuminate moralsituations by reducing them to their basic strategicform. From this perspective, game theory can actuallyserve two important functions with regard tomoral theory.4.1 The function of moralityFirst, we can use game theory to illuminate thefunction(s) of morality. For example, GAUTHIER(1986) uses game theory to argue that morality is asolution to problems of strategic interaction such asthe Prisoner’s Dilemma. Economist Ken BINMORE(1994) argues that morality is a device for choosingamong various stable equilibria in multi-persongames. (A game is said to be in equilibrium wheneach player’s choice of strategies is the best responseto her opponent’s choice of strategies.)An example may help to illustrate this point.Imagine a game in which our goal is to agree upon asocial contract (in the non-technical sense), whichinvolves negotiating a set of rules for us to live by.We can reduce this problem to its strategic essenceby representing it in simplified form as a game. Letus look at the following 2–person game, where x, y,and z are possible actions that each of us could take.If we think of this game as representing negotiationEvolution and Cognition ❘ 101 ❘ 2001, Vol. 7, No. 1

Chris MacDonaldMexyzYoux y z0 1 30 1 31 2 11 3 12 1 05 1 0changed your behaviour unilaterally. The only wayeither of us can improve our own situation is for bothof us to agree on a new equilibrium. BINMORE’s claimis that moral norms are coordination mechanismsthat allow us to deal with quandaries such as this.According to BINMORE, moral norms are coordinatingconventions for the selection of equilibria. Thisis the functionalist description of moral standards towhich an evolutionary perspective leads us. But howcan we endorse a perspective that ‘reduces’ moralityto convention? We turn to this question next.4.2 Social conventionsFigure 1: Choosing a Social Contract. The payoff pairs are{payoff to me, payoff to you}, where payoffs are assumed tocorrespond to preferences. 5 is best, 0 worst.toward a social contract, then x, y, and z representdifferent sets of rules that we could each choose tolive by.In this case, the outcomes {z,x} (at bottom left),{y,y} (centre), and {x,z} (top right) are all ‘equilibria’.That is, for each of those pairs, my choice is my bestpossible reply to your choice, and your choice is yourbest possible reply to my choice. For example, if Ichoose z, you are best off if you choose x, and if youchoose x, I am best off if I choose z. Equivalently, wecan say that if you and I are already in a situationwhere I am playing x and you are playing z, neitherof us can benefit by unilaterally changing his behaviour.The same is true for the outcome pairs {y,y} and{x,z}.(This game would be a simple matter of coordination,were it not for the fact that the payoffs in twoof the three equilibria are asymmetrical. If the payoffsin all three equilibria were symmetrical, then wewould find it easy to agree which equilibrium wasbest: it would just be a matter of coordinating ourefforts. 11 As it stands, the example above poses a hardmoral problem.)In the example at hand, we know that we’re betteroff cooperating than not cooperating, but we disagreeover which project to cooperate on (i.e., whichsocial contract to choose). If we are currently bothplaying y, (i.e., in the centre of the matrix) then Iwould prefer that we move to the lower–left of thepayoff matrix, but I can’t move us there by myself.Changing my behaviour unilaterally would makeme worse off. Similarly, you would prefer that wemove to the upper–right of the payoff matrix, butyou would be worse-off than you are now if youThe term ‘convention’ has been much abused incommon parlance and in various literatures. It isoften used as a synonym for ‘custom’ or ‘fashion’ or‘habit.’ As used by moral philosophers, it often has aderogatory flavour; the principles justified withinthe framework of one or another moral theory, forexample, are often contrasted with ‘mere social convention’.Both the sloppy and pejorative usages areunfortunate. We should seek to reclaim this undervaluedconcept, for use in both moral theory andapplied ethics. For our purposes here, we will givethe concept more content, differentiating conventionsfrom traditions, customs, norms, prejudices,and other such regularities of behaviour and belief.Conventions, here, will be defined in the HUMEANsense as regularities of behaviour, sustained by ashared interest in coordination and an expectationthat others will cooperate. Conventions, then, canbe defined in terms of these three key concepts: aregularity of behaviour; an interest in coordination;and an expectation of cooperation. The degree towhich various conventions will embody these threecharacteristics will vary, but all three characteristicswill always be present to some degree. Examples includethe everyday conventions of lining up atbanks, keeping promises, and buying alternatingrounds of beer at the pub, as well as such seriousconventions as avoiding killing non-combatants inwar. 12 Each of these ways of acting is useful for allconcerned; none of them is strictly arbitrary, nor isany of them necessarily optimal. Like these examples,many of our accepted moral obligations cannotbe understood except as the product of a convention.They are best seen as adequate responses to socialexigencies. The unifying characteristic of each ofthese situations is, roughly, a surprising degree ofsocial cohesion or order. In each situation, the interestsof the individuals involved would, on some descriptionof the situation, lead us to expect a veryEvolution and Cognition ❘ 102 ❘ 2001, Vol. 7, No. 1

Evolutionary Ethicsdifferent—much more turbulent—outcome. 13Where conventions exist to regulate behaviour thataffects others in significant ways, those conventionsmay be thought of as ‘moral’ conventions, and willbe seen as having significant moral force.Note also that implicit in the definition of a conventionis the fact that any given convention will bebut one of several possible ones for that situation. 14That is, a convention is not a regularity of behaviourstrongly determined either by human nature or theenvironment. But neither (contrary to one commonusage of the term) are conventions entirely arbitrary.Some norms may be arbitrary, but conventionsnever are. As defined here, a convention is sustainedin part by an interest in coordination: thus conventionsalways serve some useful purpose. GAUTHIER,for one, agrees with this strengthening of the term.GAUTHIER (1977, pp333–334) writes that “…to supposethat what is conventional must therefore bearbitrary is entirely contrary to the spirit of contractarianism,which finds only in convention a sufficientrationale for society”. Further, “[t]o considersociety arbitrary is to suppose that it affords no sufficientfulfilment or meets no fundamental need ofmost or all of its members”. A working conventionis always one among several solutions to a problem—butit is a solution none the less.5. ConclusionI have argued that an evolutionary perspective hasthree main implications for moral theory. An evolutionaryperspective has implications for value theory(suggesting a subjective, relative theory ofvalue), can guide us in interpreting the extent towhich various candidate moral rules are consistentwith human moral psychology, and can help usunderstand the function of different moral rules instrategic interaction. And each of these implicationspushes us toward a conventionalist understandingof ethics.A full examination of the significance of socialconventions for moral theory is beyond the scope ofthis paper. I will limit myself to making a few broad,programmatic remarks.First, we should pay attentionto the extent to which existing socialconventions structure socialinteraction and moral choice. Oftentimes, conventions will be sowell entrenched (and so psychologicallyingrained) that we willnot see ourselves as facing decisionsat all. Further, existing conventions structureindividual moral choices in that it is by reference toexisting standards that we justify our actions to others.Justification is not an abstract quality: it is alwaysa matter of justification to someone. 15 To the extentto which we are concerned with justifying our actionsto members of our own communities, we willneed to pay attention to the actual standards—existing,acknowledged social conventions—that applythere, rather than to hypothetical or ideal ones. 16 Weshould never be satisfied to rely upon existing standards;existing social conventions can often be improvedupon. But where conventions exist, they willneed to be heeded; their contribution must be analysed,and they should be ignored only when a plausiblealternative is in sight.Secondly, we should examine the extent to whichconventions should be seen as having normativeforce. Conventions represent—by definition—workingsolutions to social problems. Conventions aremutually beneficial, and thus have some motivationalforce. But conventions will not always beequally beneficial. Some conventions, for example,will be asymmetrical in that they reflect, rather thanmitigate, power imbalances between the parties involved.The possibility of asymmetrical conventionsis both a challenge to the notion of conventions asnormative, and a challenge to the common presumptionthat the notion of equity is essential tomorality.Finally, we need to investigate the possibility ofseeing conventions as a moral technology, and askwhether we can build new conventions to remedyexisting social problems. If existing conventionsserve a useful function, then perhaps we can takewhat we have learned about conventions and applythat knowledge in a forward–looking manner. Oneof the implications of this line of thought is thatgood conventions make good moral choices easier.It might even be said that the ‘real action’ of moralitygoes on long before the individual is ever faced witha decision. This points to a need for attention to thebuilding, rather than the following, of moral rules.Is it possible to engineer new conventions? Are theresome kinds of conventions which are impossible tobuild and maintain, andAuthor’s addressChris MacDonald, Department of Bioethics,Dalhousie University, 5849 University Ave.,Halifax, Nova Scotia, Canada, B3H 4H7.Email: chris.macdonald@dal.cawhich therefore oughtnot to be sought after? Ofcourse, this possibility dependsupon further investigationof the processesby which conventions (asa sub-set of social normsEvolution and Cognition ❘ 103 ❘ 2001, Vol. 7, No. 1

Chris MacDonaldmore generally) come into being, stabilize, andchange. This is fertile territory for evolutionarythought. For although I have given no account hereof the origins of conventions, it seems likely thatmost of them have their origins in some blind(though by no means random) social–evolutionaryprocess. We need to investigate further the social–evolutionary processes by which conventionsevolve. Finally, it seems likely that social thinkinginformed by an evolutionary perspective will best beable to incorporate a suitable focus on the functionof moral standards. And only through such a functionalistperspective can we reclaim moral theory asa practical discipline.Notes1 Parts of this paper were published, in Dutch in 1999, as“Evolutionaire Ethiek: Psychologie en Conventies” (EvolutionaryEthics: Psychology and Conventions), in Ethiek enMaatschappij (Ethics and Society), no. 2, vol. 4, pp. 53–61.The author would like to thank Prof. Peter DANIELSON forthe generous intellectual support that led to the writing ofthis paper. Thanks also to Bryn Williams-Jones, Jason ScottRobert, and Michael Stingl for their helpful comments onvarious versions of this paper, as well as to the anonymousreviewers for this journal.2 GAUTHIER himself refers to such absolutist conceptions as‘KANTIAN/Christian’ conceptions. Clearly, this is a shorthandway of speaking; Christian views vary enormously,and many modern KANTIANS are not Christians.3 Of course, evolutionary success itself does provide an objectivenotion of value, but such success is only necessarilyvaluable from the perspective of genes, or possibly species.Nothing forces us to value it.4 On this view, value is strictly speaking subjective. In anyparticular case, values may be shared—that is, they may beintersubjective—but this will be a contingent matter. Formore on this view of value, see GAUTHIER (1986)’s ChapterII section 4.5 Clearly, such a theory of value does not depend upon evolutionarythinking. HOBBES and HUME—writing before DAR-WIN—held such views. My argument is simply that theevolutionary perspective provides further reason for holdingto such a theory of value.6 For a number of empirical investigations based on computermodelling, see DANIELSON (1999).7 Evolutionary psychology can be seen as a more respectableheir to the role previously played by sociobiology. For moreon this, see MACDONALD (1998).8 It is commonplace in evolutionary theory to talk of evolutionor selective forces ‘designing’ various features. Thissort of talk is merely a metaphor, however—a useful shorthandway of talking about nature’s blind generate–and–testproblem solving.9 It may be that this is a behavioural tendency, without beingwhat I have called an emotional predisposition. That is,emotional commitment to kin may not be the proximalmechanism responsible for kin altruism; the relevantmechanism may be one that involves non-emotional effecton behaviour. If this is the case, it affects only the wordingof the point I make here.10 My choice situation is ‘strategic’ when the success of mystrategy depends on which strategy you choose. GAUTHIERdefines a strategic choice as one “in which the actor takeshis behaviour to be but one variable among others, so thathis choice must be responsive to his expectations of others’choices [and vice versa]”. (GAUTHIER 1986, p21) In parametricchoice, on the other hand, the actor takes his behaviorto be the sole variable in a fixed environment.11 This would then be an instance of what game theorists calla game of pure coordination. A real-life example of a gameof pure coordination is the problem of choosing on whichside of the road to drive.12 I leave it to the reader to think through the sense in whicheach of these cases exemplifies the characteristics of regularity,interest, and expectation.13 Indeed, since the bank queue approximates an n–personiterated Prisoner’s Dilemma, traditional game theory tellsus that it is flatly irrational for bank customers to line uppatiently. In my town, at least, I suspect that someone whojumped the queue would be unlikely to face serious opposition,and unilateral deviation is very unlikely to destabilizethe entire convention.14 Aside from queuing, other possible conventions herewould include drawing straws, the game of Rock–Paper–Scissors, taking numbers, sorting by age, and so on.15 As RAWLS (1971, p580) writes, “justification is argumentaddressed to those who disagree with us”. For a discussionof the pervasiveness of this approach to justification as amethodology in contemporary political thought, see NOR-MAN (1998).16 For a concrete example of the ways in which existing conventionsshape choices, see MACDONALD (2000).References:Binmore, K. (1994) Game Theory and the Social Contract,Vol. 1: Playing Fair. Cambridge MA: MIT Press.Cosmides, L./Tooby, J. (1992) Cognitive Adaptations for SocialExchange. In: Barkow, J. H./Cosmides, L./Tooby, J.(eds) The Adapted Mind: Evolutionary Psychology and theGeneration of Culture. Oxford: Oxford University Press,pp. 163–228.Danielson, P. (ed) (1999) Modelling Rationality, Morality,and Evolution. Vancouver Studies in Cognitive Science,Vol 7. Oxford: Oxford University Press.Gauthier, D. (1977) The Social Contract as Ideology, Philosophyand Public Affairs 6: 130–164.Gauthier, D. (1986) Morals by Agreement. Oxford: OxfordUniversity Press.Gauthier, D. (1991) Why Contractarianism? In: Vallentyne,P. (ed) Contractarianism and Rational Choice: Essays onDavid Gauthier’s Morals by Agreement. Cambridge: CambridgeUniversity Press, pp. 15–30.Evolution and Cognition ❘ 104 ❘ 2001, Vol. 7, No. 1

Evolutionary EthicsHobbes, T. (1985) Leviathan. London: Penguin Books. Firstpublished 1651.Kant, I. (1959) Foundations of the Metaphysics of Morals.Translated by Lewis White Beck. New York: Macmillan/Libraryof the Liberal Arts Press.MacDonald, C. (1998) Evolutionary Perspectives on Ethics.Encylcopedia of Applied Ethics, Vol. 2. San Diego: AcademicPress, pp. 189–196.MacDonald, C. (2000) Clinical Standards and the Structure ofProfessional Obligation. Professional Ethics 8(1): 7–17.Norman, W. (1998) Inevitable and Unacceptable?: MethodologicalRawlsianism in contemporary Anglo–Americanpolitical philosophy. Political Studies 46(2): 276–294.Rachels, J. (1990) Created From Animals: The Moral Implicationsof Darwinism. Oxford: Oxford University Press.Rawls, J. (1971) A Theory of Justice. Cambridge MA: HarvardUniversity Press.Evolution and Cognition ❘ 105 ❘ 2001, Vol. 7, No. 1

Zusammenfassungen der Artikelin deutscher SpracheJaak Panksepp/Jules B. PankseppDie sieben Sündender ‚Evolutionären Psychologie‘In diesem Artikel wird eine fundamentale Kritik aneinigen der basalen Voraussetzungen der ‚EvolutionärenPsychologie‘ (EP) geübt. Die Kritik gründetauf Resultaten ethologischer, ontogenetischer undneurobiologischer Untersuchungen. Der EP wirddabei vorgeworfen, Erkenntnisse dieser Disziplinennicht zu berücksichtigen und eine ‚Überinterpretation‘der menschlichen Natur vorzunehmen, dievöllig von der Idee der ‚inclusive fitness‘ beherrschtist. Vor allem die neurologischen Hintergründe vonaffektiven und motivationalen Prozessen von Menschenund Tieren stehen dabei im Hintergrund undführen zu einer ungerechtfertigten Ausdehnung desModulkonzeptes und der damit verbundenen erkenntnistheoretischenImplikationen. Kritisch betrachtetwird auch das Konzept von genetisch determiniertenModulen im Bereich des Neocortex, welcheaus der Perspektive der EP ganz bestimmteFunktionen (special purpose functions) erfüllen, derenevolutionäre Hintergründe aus den Überlebensbedingungendes Pleistozäns ersichtlich gemachtwerden können.Dagegen legen die Ergebnisse neurobiologischerForschungen am menschlichen Gehirn bzw. amSäugergehirn ganz allgemein nur die Existenz vonsubkortikalen Modulen nahe. Im Bereich ‚höherer‘Hirnregionen erscheint aus neurobiologischer Perspektivenur die Annahme der Existenz einesSprachmodules als angebracht.Der Neocortex wir im Gegensatz zur EP eher als‚Generalist‘ interpretiert, d.h. ein auf keine spezifischenFunktionen hin evoluiertes Organ, welchesontogenetischen Einflüssen und Anforderungen gegenüberin hohem Ausmaß offen ist.Die sieben Sünden der EP werden in folgendenBereichen geortet:1. Überbetonung der pleistozänen Umgebungsbedingungen,welche sich formend auf menschlichesSozialverhalten ausgewirkt haben sollen2. Extreme Zentrierung der EP auf die menschlicheSpezies3. Adaptationismus4. Massive Modularität5. Vermischung von Emotion und Kognition6. Das Fehlen glaubwürdiger neuronaler Perspektiven7. Anti-organische Tendenzen bzw. der Mythos vomcomputationalen RepräsentationalismusUllica SegerstråleFortgesetzte Konflikteum Modelle und die RealitätDie Schlacht um die ‚gute Wissenschaft‘ die im Rahmender Kontroverse um die Soziobiologie geschlagenwird setzt sich nun im Bereich der Neurowissenschaftenfort. Wissenschaftliche ‚Realisten‘ prallendabei mit eher theoriefreudigen ‚Modellierern‘ zusammen.Aus der Sicht von PANKSEPP/PANKSEPP solltendie Vertreter der EP – um entsprechend wissenschaftlichund empirisch fundiert zu sein – die Anatomieund die evolutionären Hintergründe desmenschlichen Geistes berücksichtigen.Außerdem ähnelt die PANKSEPP’sche Kritik an denkognitiven Neurowissenschaften den ethologischenEinwänden an der Soziobiologie, die nicht nur dieultimaten Funktionen von Verhalten in Betracht ziehen,sondern ebenso die Entwicklung, die Evolutionsgeschichteund proximaten Ursachen des Verhaltens.Dies mag auf einen potentiellen Konfliktbereichzwischen Ethologie und EP hinweisen.Dabei sind PANKSEPP/PANKSEPP in guter Gesellschaftanderer kritischer Positionen, wie den an Entwicklungsprozessenorientierten Wissenschaftlern,die davon ausgehen, daß die Modularität im erwachsenenGehirn nicht auf eine vorgegebene Bereichsspezifitäthinweist, sondern aus epigenetischenProzessen resultiert.In der Tat könnte gegenwärtig eine neue interdisziplinäreAllianz entstehen, indem die Kritiker vonEP und Soziobiologie auf ihrer Suche nach einemeher ganzheitlichen und pluralistischen Paradigmadie Ethologie entdeckt haben.Evolution and Cognition ❘ 106 ❘ 2001, Vol. 7, No. 1

Zusammenfassungen der Artikel in deutscher SpracheSchließlich sollte man nicht vergessen, daß dieEP ein sich entwickelndes Forschungsprogrammdarstellt und eventuell eine Annäherung von kognitiverund affektiver Neurowissenschaft zustandekommt. Ein möglicher Vermittlungsbereich könntedas Feld der nonverbalen Kommunikation sein.Linda Mealey‚La plus ça change...‘:Antwort auf eine Kritik der EPPANKSEPP/PANKSEPP stehen nicht alleine in ihrer Kritikan der ‚Evolutionären Psychologie‘. Die meistenKritiker, wie auch PANKSEPP/PANKSEPP sind dabeinicht gegen Evolution oder Psychologie bzw. gegendie Verwendung evolutionärer Konzepte in der Psychologieeingestellt. Die Kritik richtet sich dabei vorallem gegen eine bestimmte, bedeutsame Gruppevon der EP nahestehenden Wissenschaftlern, dieeine gemeinsame Haltung zu zentralen Kernproblemeneinnehmen.Ich teile die Bedenken von PANKSEPP/PANKSEPPhinsichtlich der Rigidität und doktrinären Einschränkungendie in manchen Versionen der EP gegebensind. Jedoch ist diese Debatte nicht neu undich glaube, daß sie, wie auch andere wissenschaftlicheKontroversen durchaus förderliche Eigenschaftenhaben.Carlos StegmannKommentar zu Panksepp/PankseppPANKSEPP/PANKSEPP behaupten, daß die EP – um innerhalbdes Rahmens wissenschaftlicher Forschungzu verbleiben – mit den Erkenntnissen der Neurowissenschaftenund der Hirnforschung übereinstimmenmuß. Dieses Argument wird im Kommentarerweitert. Die menschliche Spezies stellt eine integraleEinheit dar, die aus unterschiedlichenSchichten besteht. Dazu gehören sowohl dieSchichte des soziokulturellen Verhaltens von Gruppen,wie die Schichte des individuellen Verhaltensund die Schichte der neuronalen Mechanismen. JederVersuch einer Erklärung der menschlichen Natur,inklusive der Psychologie muß daher interdisziplinärausgerichtet sein.Psychische Phänomene müssen aus der soziokulturellenSchichte heraus erklärbar sein, wie beispielsweisediejenigen die Friedrich HAYEK in Bezugauf die ‚extended order‘ der menschlichen Gesellschaftentdeckt hat. Ebenso müssen diese Gesetzeauf die neurophysiologischen Gesetze rückführbarsein, die auf dieser Schichte anzutreffen sind.Nur derartige interdisziplinäre Ansätze stelleneine Kompaß bereit anhand dessen man durch dasdunkle Meer des menschlichen Gehirns und der zugehörigenGeistphänomenen navigieren kann.Russell Gardner, Jr.Affektive Neurowissenschaft,Psychiatrie und SoziophysiologiePANKSEPP/PANKSEPP betonen in anerkennenswerterWeise daß die EP nicht die stammesgeschichtlich altenBereiche des menschlichen Gehirns in Betrachtzieht. Die EP vermischt subkortikale emotionale Faktorenmit Kognitionen, die im Verlauf der Evolutiondes Menschen verstärkt wurden. Weiters beachtetdie EP die kommunikativen und sozialen Bereicheder menschlichen Entwicklung zu wenig, obwohlneue Arbeiten deutlich machen, daß ein signifikanterAnteil der Varianz der Hirngröße von Primatenmit der jeweiligen Gruppengröße zusammen hängt.Jaak PANKSEPPs frühere Arbeiten über emotionaleKommunikation könnten dabei zur Entwicklng einergrundlegenden psychiatrischen Wissenschaftbeitragen, die derzeit leider kaum existiert.In Übereinstimmung mit PANKSEPP steht die Forderungnach einem basalen, wissenschaftlichenRahmen für die Psychiatrie mit dem Titel Soziophysiologie,welche sowohl die Ähnlichkeiten als auchdie Gemeinsamkeiten des Menschen mit anderenTieren untersucht.Gerhard MeisenbergGrade der ModularitätDie Unterscheidung zwischen kognitiven und modularenFunktionen, wie sie ursprünglich von FO-DOR (1983) vorgeschlagen wurde ist gültig. Jedochwerden zwei Formen modularer Funktionen unterschieden.:(1) ‚Primäre‘, genetisch determinierte Module,die überwiegend subkortikal vorliegen und hauptsächlichMotivation (z.B. Konditionierung vonAngst) bedingen.(2) ‚Modularisierte‘ Funktionen, die vor allemkortikal vorliegen und durch wiederholten sensorischenund /oder kognitiven Input geformt werden.Diese Funktionen sind vor allem für sensorischeAnalyse und motorische Kontrolle verantwortlich(z.B. Sprache).Evolution and Cognition ❘ 107 ❘ 2001, Vol. 7, No. 1

Zusammenfassungen der Artikel in deutscher SpracheDie strikte Unterscheidung zwischen genetischdeterminierten subkortikalen Schaltkreisen undnicht modular organisierten kognitiven Funktionen,wie sie von PANKSEPP/PANKSEPP vorgeschlagenwird, wird zurückgewiesen. Ebenso wird das von derEP vertreten Modell des ‚Schweizer Armeemessers‘als zu vereinfachend abgelehnt.Im Gegensatz dazu wird behauptet, daß vieleHirnfunktionen (inklusive der kognitiven Funktionen)‚Mehr-Zweck‘–Mechanismen darstellen. Diese‚guns for hire‘ stehen in Form von spezialisiertenModulen zur Verfügung um adaptives Verhalten zuerzeugen.Es wird behauptet, daß das nützlichste Konzeptein ausgedehntes ethologisches Modell darstellt, indem spezialisierte Gehirnsysteme entweder sensorischenoder kognitiven Input verwenden und damitentweder Verhalten oder kognitiven Output erzeugen.Ian PitchfordKeine Evolution. Keine KognitionObwohl sowohl Natur- wie auch Geisteswissenschaftlerjeglichen Glauben an unterschiedlichematerielle und immaterielle Substanzen leugnenwürden, werden gegenwärtige Debatten größtenteilsmit einer Terminologie geführt, die antikengriechische Philosophen wohl sehr vertrau gewesenwäre. Dabei werden menschliche Merkmale ingöttlich – transzendente und tierisch – körperlicheunterteilt. Die modernen Fassungen dieser Unterteilungenlauten dann rational, kognitiv, diskursiv, autonomund kreativ – für den göttlichen Anteil, sowieemotional, instinktiv, determiniert etc. für dentierisch – körperlichen Teil.Dieser grundlegende Dualismus erhält die, in fastallen Diskussionen über den Menschen anzutreffendendrei zentralen Dichotomien Körper/Geist,Kognition/Emotion und Natur/Kultur aufrecht.Das Bemühen der neuroevolutionären Psychobiologiedas Gehirn in einen determinierten affektivenTeil und einen plastischen, nicht direkt funktionsgebundenenNeocortex zu unterteilen, indem Konzeptewie ‚exaptation‘, Emergenz und dgl. mehrverwendet werden steht genau in dieser quasitheologischenwestlichen philosophischen Tradition.Dieser dualistische Ansatz stellt keine schlüssige Basisdar, von der ausgehend die EP kritisiert werdenkann.Scott AtranDer ‚Fall‘ Modularität:Sünde oder Erlösung?Die These von evoluierten kognitiven Modulen beruhtauf unterschiedlichen, konvergierenden Beweisen:functional design (Analogie), Ethologie(Homologie), Universalität, Unabhängigkeit vonPerzeptionen, spezifische Pathologie (zerebrale Störungen),Hemmungswiderstand (Hyperaktivität),Leichtigkeit der kulturellen Übertragung.Zwar mag keiner der Einzelfaktoren notwendigsein, jedoch ist die Datenlage zwingend und eindeutigfür Bereiche wie beispielsweise ‚folk biology‘ und‚folk psychology‘.Im Gegensatz dazu wird die zentrale These vonPANKSEPP/PANKSEPP durch keinerlei empirische Hinweisegestützt. Diese These geht davon aus, daßmodulare Strukturen – wie sie von der EP (nebenzahlreichen kognitiven Psychologen und Entwicklungspsychologen)interpretiert werden – epigenetischeProdukte subkortikaler Emotionen (diebei allen Säugetieren vorkommen) und neokorticalerunspezifischer ‚Mehr-Zweck‘–Operationensind.Keine einzige bedeutsame empirische Entdekkungim Bereich Sprache oder anderer höherer kognitiverFunktionen ist bei den Untersuchungen zuden evolutionären Ursprüngen und Funktionen offengeblieben.Jaak Panksepp/Jules B. Panksepp:Weiter Kritik an der EP:7 Sünden für 7 Sünder – plus/minus 2Die vielfältigen, von den Kommentatoren behandeltenProblembereiche ließen zwei zentrale Fragenoffen: eine Debatte über Modularität sowie die Evolutionvon Sprache. Dahingehend bietet eineneokortikale – entwicklungsbezogene – systemischePerspektive, zusammen mit einem Verständnisder basalen emotional – motivationalen Systeme(die wir mit anderen Säugetieren teilen) mehr Einsichtenin die Formen intrinsischen Funktionierensmenschlicher Gehirn/Geist Prozesse als jegliche Variantekortikaler Modularität.Es erscheint sehr unwahrscheinlich, daß die Entwicklunghöherer kortikaler Funktionsbereiche,welche Sprache bedingen, ohne zugrundeliegende,subkortikale Systeme abgelaufen ist. Obwohl Menschen‚kommunikative Instinkte‘ bzw. über ‚Anla-Evolution and Cognition ❘ 108 ❘ 2001, Vol. 7, No. 1

Zusammenfassungen der Artikel in deutscher Sprachegen zum Spracherwerb‘ verfügen können ist die Existenzvon sog. ‚Sprachmodulen‘ nicht belegt. Dahingehenderscheint es angebracht, daß dieVertreter der EP neurowissenschaftliche Erkenntnissein ihre Überlegungen einbeziehen.SEGERSTRÅLE – eine historische Perspektive: SEGERSTRÅLEstellt diese Kontroverse in den aktuellen historischenKontext. Wir stimmen dabei mit ihr überein,daß die Ethologie in die Fundamente psychologischenDenkens einbezogen werden soll. Wir stimmennicht mit ihrer Behauptung überein, daß ‚wirklicheWissenschaft‘ nur auf einer ‚anatomisch–physischenEbene‘ gemacht wird, sondern räumentheoretischen Perspektiven ebenso einen zentralenStellenwert ein.SEGERSTRÅLE bemerkt zutreffend, daß unsere Bemühungengegen einen wissenschaftlichen ‚Tribalismus‘gerichtet sind, der eine Fülle von Datenmaterialüber ‚nonverbale Kommunikation‘ sowie diezugrundeliegende neurophysiologische Basis ausdem tierischen Verhaltensbereich völlig mißachtet.Wir wenden uns gegen die anthropozentrische Perspektiveder EP und gehend dabei davon aus, daßzahlreiche, von der EP postulierte, in höheren kortikalenBereichen angeordnete Module bereits in subkortikalenArealen (die sich ebenso in anderen Tiergruppenfinden) vorliegen. Dabei ziehen wir nichtin Zweifel, daß es innerhalb des Kortex beträchtlichesensorische und motorische Spezialisierungengibt (inklusive diverser Gesichts-, Geruchs-, Berührungs-und Stimmlagen–Erkennungsdispositionen).Jedoch sind diese Dispositionen weder soziobiologisch(im eigentlichen Wortsinn) noch nurbeim Menschen anzutreffen.Wir stimmen SEGERSTRÅLE dahingehend zu, daßes an der Zeit ist „to relax the modularity idea“.MEALEY – Was ist in einem Wort? MEALEY gibt eineausgewogene Übersicht über die grundlegendenund einige spezielle Argumente. Dabei stimmen wirnicht mit ihrer Verteidigung der modularen Sichtweiseüberein.Neben allen anderen ‚Sünden‘ ist dabei die ‚SpezieszentriertePerspektive‘ zentral. Wir vermissendabei die Integration neurowissenschaftlichen Datenmaterialsin die akademische Psychologie –ohne diese Integration kann die Psychologie niemalseine ‚grundlegende Wissenschaft‘ werden.Gegen den Modularitätsbegriff wie ihn MEALEYverwendet wenden wir ein: er vermittelt ein unzureichendesund vereinfachendes Bild der Hirnfunktionen,die als zusammengeschlossene (encapsulated)Informationsverarbeitungssysteme interpretiertwerden. Dabei werden die zahlreichenWechselwirkungen innerhalb des Gehirns völligmißachtet. Einen ähnlichen Fehler machte auchdie frühe Neurowissenschaft mit dem Postulat vonsog. ‚Zentren‘ – welches bald nicht mehr verwendetwurde.Der Modularitätsbegriff stellt im Bereich der Psychologiekein neues und sinnvolles Konzept dar.Dagegen scheint es sinnvoll etwa den Begriff des‚Triebes‘ wieder einzubringen, da dieser eher den Erkenntnissenneurowissenschaftlicher Untersuchungenentspricht.STEGMANN – eine philosophische Perspektive: Aus einerphilosophischen Perspektive heraus stimmenwir mit STEGMANN überein, jedoch aus der wissenschaftlichenPerspektive heraus üben wir Kritikdaran. Derartige holistische Perspektiven sind ofteiner systematisch – experimentellen Analyse entgegengesetzt.Ebenso wenden wir uns gegen die Annahme, daßkomplexere Schichten auf darunterliegende Systemebenereduzierbar sein sollen. Dieser Form desReduktionismus stellen wir den modernen philosophischenBegriff der ‚supervenience‘ entgegen. Wirwären sehr daran interessiert zu erfahren, inwieweitSTEGMANNs Ideen empirischer Überprüfung zugeführtwerden könnten.GARDNER – eine neuropsychiatrische Perspektive: GARD-NER geht von der – im Bereich der Psychiatrie besondersdringlichen Notwendigkeit der Integrationunterschiedlicher Perspektiven hinsichtlich Gehirn/GeistPhänomenen aus. Nach wie vor werdendiese Phänomene getrennt behandelt und die Notwendigkeitdiverser Brückenschläge ist eine der großengegenwärtigen Herausforderungen.Es ist eines der wesentlichen Anliegen des Seniorautorsdie tiefen biologischen Muster psychiatrisch/emotionaler Störungen aufzuzeigen. Überaus hilfreichist dabei die Einbeziehung von Datenmaterialienaus tierischen Verhaltensanalysen. Die dabeivorgefundenen basalen emotionalen Systeme, diewir mit allen Säugetieren gemeinsam haben erweisensich dabei besonders für die Psychiatrie vongroßem Wert. Die Entstehung der neuro-psychoanalytischenBewegung erscheint als ein hoffnungsvollerSchritt in diese Richtung.MEISENBERG – Modularität ist am Leben und guterDinge – angeblich. Meisenberg scheint mit den größtenTeil unserer Überlegungen übereinzustimmenEvolution and Cognition ❘ 109 ❘ 2001, Vol. 7, No. 1

Zusammenfassungen der Artikel in deutscher Spracheund unternimmt eine weiterreichende Analyse kortikalerFunktionen.Wir selbst zweifeln nicht daran, daß große Teiledes Neokortex – als Folge von Erfahrungen spezialisiertsind. Jedoch stellen wir die Existenz soziobiologischbegründeter, kognitiver Module in Zweifel. Esgibt keinerlei Hinweise darauf, daß sich modularisierte,kognitive Adaptationen im Neokortex – imVerlauf der letzten 2 Jahrmillionen entwickelt haben.Wir gehen davon aus, daß dasjenige, was sichin diesem Zeitrahmen entwickelt hat eher unspezifische,d.h. nicht spezialisierte Funktionsbereiche waren,die erst unter Umwelteinflüssen und mit subkortikalerBeteiligung modularisiert wurden.Wir stimmen mit MEISENBERGs Überlegungenhinsichtlich der Entwicklung des Lächelns überein.Jedoch stimmen wir ihm nicht zu, daß die relevantenneurologischen Strukturen vor allem im cingulärenKortex angeordnet sind. Demgegenüber nehmenwir an, daß auch noch tieferliegende Gehirnregionenam Lächeln beteiligt sind.In striktem Widerspruch stehen wir zu MEISEN-BERGs Behauptung daß das Datenmaterial welchessich auf die subkortikale Ebene bezieht nicht so solideund fundiert ist wie wir das behaupten.Atran – noch einmal das ‚Sprachmodul‘. ATRAN lieferteine heftige Verteidigung der Modularitätsidee undbezieht sich dabei vor allem auf die Sprache. DasZentralproblem dabei kann folgendermaßen formuliertwerden: beinhalten die Gene die Rudimenteunserer einzigartigen menschlichen Sprache oderhelfen diese Gene mit komplexe Laute zu erzeugen,die mit dem enormen Speicherraum des Kortex interagierenund als Endprodukt Sprache generieren.Nach unserem Dafürhalten ist es nicht notwendigdie Existenz von genetisch fixierten Sprachmodulenanzunehmen. Sprache entwickelt sich ausdem Zusammenwirken von subkortikalen Hirnstrukturenmit dem Neokortex.Wir stimmen auch nicht mit ATRANs ‚kortikal –kognitivistischer Sichtweise‘ überein, nach der keinerleiBefunde aus dem subkortikalen Bereich Einsichtenin das Funktionieren kognitiver Strukturenvermitteln konnte. Die Bedeutung und der Einflußsubkortikaler Aktivitäten wurde und wird nach wievor weitgehend unterschätzt.PITCHFORD – die Herausforderung eines Insiders. PITCH-FORD bezieht sich vor allem auf eine der von uns postuliertenSünden der EP – die Suche nach adaptivenModulen Bereichen, in denen jedoch nur nichtfunktionsgebundene Operationen vollzogen werden.PITCHFORD thematisiert dabei vor allem dieSprachfunktion und die Rolle von Entwicklungsprozessen.PITCHFORDs Behauptung die Autoren hätten diezentralen Argumente der ‚developmental systemstheory‘ nicht entsprechend berücksichtigt wird zurückgewiesen,da vor allem der Seniorautor dazuzahlreiche Publikationen vorgelegt hat. Dabei stimmenwir größtenteils mit den Argumenten der ‚developmentalsystems theory‘ überein – ein Kritikpunktjedoch richtet sich gegen jene überzeichneteArgumentation in der von ‚Ontogenie von Information‘die Rede ist.Leider geht PITCHFORD nicht auf unsere Kernargumenteein.Noch einmal weisen wir darauf hin, daß der gefährlichsteOrt der Suche nach Modularität derjenigevon hoch spezialisierten neokortikalen Arealenim Gehirn erwachsener Menschen ist. Wir sind derMeinung, daß sich die allgemeinsten Funktionsprinzipieneher auf subkortikalem Niveau finden.Shulamith KreitlerEine evolutionäre Perspektiveauf kognitive OrientierungDer Beitrag beschäftigt sich mit den evolutionärenAspekten der kognitiven Steuerung des Verhaltens.Aufgrund empirischer Befunde und der eindrucksvollenEntwicklung von Kognition im Verlaufder Evolution kann davon ausgegangen werden,daß die kognitive Verhaltenssteuerung inBezug auf Überleben eine Rolle spielt. Eine Theoriekognitiver Orientierung – auf dem Niveau desMenschlichen – wird als grundlegender Rahmenfür eine evolutionäre Analyse präsentiert. Die großenVorteile einer solchen Theorie bestehen in einerumfassenden Beschreibung der zwischen Inputund Output wirkenden Prozesse, für die esvielfache empirische Belege gibt. Die wichtigstenevolutionären Aspekte, die besprochen werden,betreffen die Unterscheidung zwischen Motivationund Kognition, die Entwicklung von Bedeutungsowie die Entwicklung von Handeln, in jedemFall unter besonderer Berücksichtigung derRolle von Kognition.Evolution and Cognition ❘ 110 ❘ 2001, Vol. 7, No. 1

Zusammenfassungen der Artikel in deutscher SpracheMac DonaldEvolutionäre Ethik: Werte, Psychologie,Strategie und KonventionenAbsicht dieses Artikels ist es deutlich zu machen,welche Auswirkungen eine strikte Anwendung derEvolutionstheorie auf eine Theorie der Moral hat. Ineinem ersten Schritt werden die Implikationen derEvolutionstheorie für die Werttheorie und das moralischeVerhalten aufgezeigt. Danach wird der Versuchunternommen eine evolutionäre Perspektivehinsichtlich des Problems der Entwicklung sozialerKonventionen darzustellen. Moralische Standardswerden dabei als Produkte der kulturellen und derbiologischen Evolution begriffen, wobei basale moralischeWerte weder von der Natur noch vom Evolutionsprozeßselbst abgeleitet werden. Die Anwendungdes DARWINISTISCHEN Denkens erfolgt dabeivor allem auf die empirische Basis, auf der aufbauendmoralische Theorien entwickelt werden.Bezug genommen wird dabei auch auf die ‚EvolutionärePsychologie‘ welche viel zum Verständnismoralischen Verhaltens beitragen kann.Weiters erweist sich die Untersuchung sozialerKonventionen als zentrales Element. Konventionenwerden dabei als Regularitäten des Verhaltens interpretiert.Diese werden durch mit anderen Menschengeteilte Interessen aufrechterhalten wobei die Erwartungder Kooperationsbereitschaft zugrundeliegt.Evolution and Cognition ❘ 111 ❘ 2001, Vol. 7, No. 1