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Reduction and Elimination in Philosophy and the Sciences

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Are Lamarckian Explanations Fully Reducible to Darw<strong>in</strong>ian ones? The Case of “Directed Mutation” <strong>in</strong> Bacteria — Davide Vecchi<br />

environmental factors are irrelevant to mutational<br />

responses (it has been established that <strong>the</strong>re exist very<br />

specific processes of mutational response triggered by<br />

environmental stresses like starvation), nor on <strong>the</strong><br />

hypo<strong>the</strong>sis that <strong>the</strong>re is no target<strong>in</strong>g of specific genomic<br />

regions, while grow<strong>in</strong>g evidence is accumulat<strong>in</strong>g <strong>in</strong> favour<br />

of see<strong>in</strong>g mutational responses as evolved adaptive<br />

responses of <strong>the</strong> organism. What can be safely said today<br />

is that r<strong>and</strong>om variation is not <strong>the</strong> norm. Mutational<br />

processes are highly biased <strong>and</strong> targeted affairs. Still, can<br />

a revised <strong>and</strong> softened version of Darw<strong>in</strong>ism suffice to<br />

dispel <strong>the</strong> ghost of Lamarck? As Hall (1998) argues,<br />

adaptive mutation is a different process from growthdependent<br />

or “normal” bacterial mutation, even though it<br />

clearly is a Darw<strong>in</strong>ian process. The important questions on<br />

this score are: how is <strong>the</strong> hypermutation state triggered? Is<br />

it triggered by <strong>the</strong> environmental stress or by <strong>the</strong> selective<br />

pressure? Is it regulated by specific genes, or ra<strong>the</strong>r by<br />

some unspecified <strong>and</strong> unknown, possibly <strong>in</strong>nate, capacity<br />

of <strong>the</strong> organism? We still don’t know. For <strong>the</strong> same<br />

reasons, I contend, we cannot answer <strong>the</strong> question of<br />

whe<strong>the</strong>r we need an explanation of HSH <strong>in</strong> terms couched<br />

via <strong>the</strong> language of <strong>in</strong>tentionality <strong>and</strong> choice, or whe<strong>the</strong>r a<br />

Darw<strong>in</strong>ian explanation couched <strong>in</strong> terms of chance <strong>and</strong><br />

selection is all we need. Or so I will argue.<br />

It could be argued that <strong>the</strong> phenomenon of adaptive<br />

mutagenesis is purely Darw<strong>in</strong>ian. From this perspective it<br />

could be asked: what does <strong>the</strong> Lamarckian language of<br />

<strong>in</strong>tentionality, purpose <strong>and</strong> choice add to explanations<br />

concern<strong>in</strong>g adaptive mutations? Noth<strong>in</strong>g is <strong>the</strong> answer:<br />

Lamarckian concepts are simply redundant. To say that<br />

organisms choose to mutate <strong>in</strong> particular ways, that <strong>the</strong>y<br />

deliberately try to reach a particular adaptive goal, that<br />

<strong>the</strong>y <strong>in</strong>tend to change <strong>and</strong> adapt <strong>in</strong> specific ways, is<br />

equivalent to <strong>in</strong>vite <strong>in</strong>appropriate teleological talk. Even<br />

though emphasis on <strong>the</strong> phenomenon has highlighted <strong>the</strong><br />

need to discrim<strong>in</strong>ate between mutations with different<br />

characteristics concern<strong>in</strong>g <strong>the</strong>ir basic properties<br />

(environmental conditions, time <strong>and</strong> site of occurrence), it<br />

is clear that <strong>the</strong> directional outcome is a product given by<br />

<strong>the</strong> extreme reduction of design search space, by <strong>the</strong><br />

temporal <strong>and</strong> local <strong>in</strong>crease <strong>in</strong> efficiency of <strong>the</strong> search, <strong>and</strong><br />

by a more general strategy of mutation rate <strong>in</strong>crease.<br />

A variant of <strong>the</strong> same dismissive attitude could<br />

<strong>in</strong>stead stress that <strong>in</strong>stead of explanatory redundancy<br />

(based on <strong>the</strong> application of Ockham’s razor), Lamarckian<br />

explanations are reducible to Darw<strong>in</strong>ian ones. This would<br />

v<strong>in</strong>dicate <strong>the</strong> hypo<strong>the</strong>sis that Darw<strong>in</strong>ism is a more<br />

fundamental process than Lamarckism. In fact, it seems<br />

that <strong>the</strong> “<strong>in</strong>tentionality” of Lamarckian systems like those<br />

supposedly govern<strong>in</strong>g adaptive mutational responses must<br />

arise ei<strong>the</strong>r by means of <strong>the</strong> cumulative effects of past<br />

natural selection act<strong>in</strong>g on ultimately r<strong>and</strong>om mutations, or<br />

more simply by a more error-prone pathological response<br />

of cells. Therefore <strong>the</strong> epistemic reducibility of Lamarckian<br />

explanations seems to be achievable. So, even though <strong>the</strong><br />

orthodox <strong>and</strong> hardened Neo-Darw<strong>in</strong>ian views on directed<br />

mutation has been revised <strong>and</strong> updated, <strong>the</strong> revision of <strong>the</strong><br />

Darw<strong>in</strong>ian view has not touched certa<strong>in</strong> residual <strong>and</strong><br />

crucial elements constitutive of some mean<strong>in</strong>g of<br />

“r<strong>and</strong>omness”, which make an appeal to <strong>the</strong> Lamarckian<br />

language of <strong>in</strong>tentionality <strong>in</strong>appropriate. This alternative<br />

seems to illustrate <strong>the</strong> sensible view shared by most<br />

practitioners.<br />

However, I th<strong>in</strong>k that this epistemological move<br />

cannot shed any light on <strong>the</strong> ontological issue perta<strong>in</strong><strong>in</strong>g to<br />

<strong>the</strong> nature of mutation, <strong>and</strong> on <strong>the</strong> basic question concern<strong>in</strong>g<br />

<strong>the</strong> existence of genu<strong>in</strong>e Lamarckian processes.<br />

What could a genu<strong>in</strong>e Lamarckian phenomenon be like?<br />

Genu<strong>in</strong>e <strong>in</strong>tentionality would perhaps <strong>in</strong>volve <strong>the</strong><br />

preferential environmental <strong>in</strong>duction of useful <strong>and</strong><br />

beneficial mutations, which is not happen<strong>in</strong>g, or <strong>the</strong><br />

target<strong>in</strong>g of specific genes, which is still quite controversial.<br />

But an equally promis<strong>in</strong>g avenue would be to consider<br />

genu<strong>in</strong>e <strong>in</strong>tentionality as <strong>in</strong>volv<strong>in</strong>g <strong>the</strong> choice of starved<br />

<strong>and</strong> stresses cells to enter <strong>the</strong> hypermutation state. I th<strong>in</strong>k<br />

that at this juncture lies <strong>the</strong> best chance to v<strong>in</strong>dicate some<br />

genu<strong>in</strong>e Lamarckian credentials. If <strong>the</strong> hypermutation state<br />

is triggered by some as yet unspecified <strong>and</strong> unknown<br />

capacity of <strong>the</strong> organism, <strong>the</strong>n Lamarckism could be<br />

v<strong>in</strong>dicated. Or better it would be if we were unable to f<strong>in</strong>d a<br />

mechanistic basis for <strong>the</strong> cell response.<br />

Intentionality does not seem to be needed to provide<br />

explanations that save <strong>the</strong> known phenomena of adaptive<br />

mutagenesis. Still, this could be simply appearance. The<br />

crucial question rema<strong>in</strong>s: how do we expla<strong>in</strong> <strong>the</strong> fact that<br />

only a subpopulation of bacteria hypermutates? It seems<br />

that some element <strong>in</strong> <strong>the</strong> environment triggers such a<br />

state, or perhaps it is a cell that chooses to do so? Where<br />

is <strong>the</strong> evidence <strong>in</strong> favour of <strong>the</strong> first “mechanical”<br />

hypo<strong>the</strong>sis? And is <strong>the</strong> first hypo<strong>the</strong>sis simpler than <strong>the</strong><br />

second? And why is it simpler to hypo<strong>the</strong>sise that <strong>the</strong><br />

bacterium senses an environmental signal ra<strong>the</strong>r than<br />

hypo<strong>the</strong>sis<strong>in</strong>g that it chooses to hypermutate? In <strong>the</strong> first<br />

case we need a sensory mechanism, <strong>in</strong> <strong>the</strong> second just<br />

agency of some k<strong>in</strong>d. Fur<strong>the</strong>rmore, even a sensory<br />

mechanism could be sufficient to phagocyte a slice of <strong>the</strong><br />

<strong>in</strong>tentional cake.<br />

The position of <strong>the</strong> reductionist <strong>and</strong> materialist rests<br />

on <strong>the</strong> adoption of <strong>the</strong> argument from parsimony. But<br />

voiced have been raised to argue aga<strong>in</strong>st this adoption of<br />

Ockham’s razor, <strong>and</strong> aga<strong>in</strong>st a more general ontological<br />

conservatism (Sober 2001), especially when <strong>the</strong><br />

<strong>in</strong>tentionality hypo<strong>the</strong>sis is ontologically as economical as<br />

<strong>the</strong> alternatives. The issue is that as <strong>the</strong>re are no<br />

arguments <strong>in</strong> favour of <strong>the</strong> reification of <strong>in</strong>tentionality as a<br />

dist<strong>in</strong>ct <strong>and</strong> emergent quality, so <strong>the</strong>re are no arguments<br />

aga<strong>in</strong>st.<br />

So it could be that <strong>the</strong> more one digs <strong>in</strong>to <strong>the</strong><br />

subject <strong>the</strong> more <strong>the</strong> Lamarckian language of choice<br />

makes sense. In this sense it is not enough to po<strong>in</strong>t out<br />

that polymerase IV <strong>and</strong> <strong>the</strong> down regulation of MMR are<br />

<strong>in</strong>volved <strong>in</strong> <strong>the</strong> process of hypermutation, that is, an errorenhanc<strong>in</strong>g<br />

<strong>and</strong> a less efficient error-edit<strong>in</strong>g process. The<br />

question rema<strong>in</strong>s of what triggers <strong>the</strong> state, of whe<strong>the</strong>r it is<br />

<strong>the</strong> bacterium that “decides” to hypermutate.<br />

Also note that <strong>in</strong> many ways a panpsychist<br />

hypo<strong>the</strong>sis, with <strong>the</strong> postulation of all its monads, is not<br />

clearly more complex <strong>and</strong> <strong>the</strong>refore amenable to be<br />

chopped off with one stroke of Ockham’s razor. Even<br />

bacterial chemotaxis is nowadays described somehow <strong>in</strong><br />

terms of cells’ <strong>in</strong>tentionality (cells are analogised to<br />

nanobra<strong>in</strong>s). The panpsychist perspective accord<strong>in</strong>g to<br />

which <strong>in</strong>tentionality is everywhere rema<strong>in</strong>s simpler that any<br />

emergentist view. If mentality <strong>and</strong> <strong>in</strong>tentionality are basic<br />

<strong>and</strong> primitive, <strong>the</strong>n ascriptions of mentality are, despite <strong>the</strong><br />

risk of anthropomorphism, always justifiable. The idea that<br />

<strong>the</strong>re exists a cont<strong>in</strong>uum of levels of <strong>in</strong>tentionality, ra<strong>the</strong>r<br />

than an abrupt emergence of someth<strong>in</strong>g totally new,<br />

rema<strong>in</strong>s appeal<strong>in</strong>g.<br />

The great appeal of <strong>the</strong> Darw<strong>in</strong>ian account (e.g.<br />

based on a bl<strong>in</strong>d variation <strong>and</strong> selective retention formula)<br />

is that it offers a very general mode of explanation, <strong>and</strong> a<br />

clear recipe to dispense of epiphenomenal <strong>in</strong>tentionality<br />

(Hull 1988). The active agent easily disappears from<br />

selectionist accounts. But can we get rid of agency so<br />

363

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