229 Slide Molecular Genetic Approaches toChemoreceptionNEURON-SPECIFIC ODOR RECEPTOR GENE CHOICE INDROSOPHILARay A. 1 , Van Der Goes Van Naters W. 1 , Carlson J. 1 1 MCDB, YaleUniversity, New Haven, CTWe have uncovered a combinatorial code of cis-regulatory elementscritical to the process of odor receptor (Or) gene choice in Drosophila.The Or gene family in Drosophila consists of 60 members. Individualmembers are expressed in specific functional classes of olfactoryreceptor neurons (ORNs) of either the antenna or the maxillary palp.We have identified both positive and negative cis-regulatory elementsthat act together to dictate the organ-specific expression of individualreceptor genes in the maxillary palp. In Drosophila a special subset ofORNs expresses not one but two Or genes. We have identified twodistinct mechanisms by which such co-expression can be achieved.First, we demonstrate that some co-expressed Or genes share specificsequence motifs in their flanking DNA that can direct expression in acertain neuronal type. Second, for some of the tightly linked Or genes,we have found evidence that the downstream gene can share the samepromoter as the upstream one by virtue of alternative splicing of themRNA. The majority of ORNs, however, express a single Or gene. Wehave used comparative genomics to identify evolutionarily conservedcis-acting neuron-specific elements that act in specifying the expressionof these classes of palp ORNs. In vivo analysis of these elements hasled to a model for Or gene choice. We find that within an olfactoryorgan a given Or gene has the ability to express in a few different ORNclasses; however, transcriptional repressors restrict expression to theappropriate neuronal class. Finally, we have identified transcriptionfactors that are required for this highly regulated process of Or genechoice.230 Slide Molecular Genetic Approaches toChemoreceptionALTERING OLFACTORY NEURON IDENTITY WITHECTOPIC EXPRESSION OF G-PROTEIN COUPLEDRECEPTORSChesler A. 1 , Le Pichon C. 2 , Peterlin Z.A. 3 , Matthews G. 4 , Zou D. 2 ,Firestein S. 2 1 Biology, Columbia University, New York, NY; 2 ColumbiaUniversity, New York, NY; 3 Biological Sciences, Columbia University,New York, NY; 4 Neurobiology and Behavior, Columbia University, NewYork, NYIn the mammalian olfactory epithelium (OE), each olfactory sensoryneuron (OSN) is thought to express only one allele of a single olfactoryreceptor (OR) gene. It has been proposed that the expressed OR proteinrepresses activation of additional OR genes through feedback inhibition.The resulting singular OR expression effectively determines the identityof the OSN by defining its odorant response properties and influencingits axonal projection. The mechanisms by which ORs influence OSNidentity are unknown. We have taken a gain-of-function approachusing ultrasound-guided injection of the embryonic OE with retroviralvectors to study the role of ORs in the development of OSN identity.We find that the ectopic expression of functional GPCRs, either ORI7or beta-2-Adrenergic Receptor, perturb odorant response profiles andaxonal sorting without a noticeable effect on endogenous ORexpression. Additionally, expression of an amino-terminally mutatedORI7 that encodes a full-length and properly trafficked OR does notconfer octanal responsiveness or perturb axonal sorting. Our worksupports a model wherein modulation of G-protein signaling, at least inpart, encodes OSN identity. In conclusion, embryonic expression ofGPCRs using retroviral vectors offers a new gain-of-function approachfor studying the role of ORs in olfactory development. SupportContributed By: NIDCD.231 Slide Molecular Genetic Approaches toChemoreceptionUBIQUITOUS EXPRESSION OF AN ODORANT RECEPTORIMPAIRS PROPER AXON TARGETING TO THE OLFACTORYBULBVidaltamayo R. 1 , Reed R.R. 1 1 Molecular Biology and Genetics,Howard Hughes Medical Institute, Baltimore, MDIn order to study the effects of widespread expression of an odorantreceptor (OR) in the olfactory epithelium, we have generated atransgenic mouse line in which we placed the coding region of the mI7OR gene into the transcription factor O/E3 locus. Expression of thistranscription factor occurs early in olfactory neuron development andpersists in mature olfactory neurons in the adult mouse. In the O/E3-mI7 line, we detect widespread expression of mI7 OR, encompassing allfour zones of the olfactory epithelium, from birth (postnatal day 0) up toadulthood (8 week-old mice), both at the mRNA, by in situhybridization, and protein level, by immunofluorescence.Electroolfactogram recordings show a 100-fold increase in the doseresponse to the mI7 ligand, heptaldehyde (-4.59 ± 1.17 mV at 1 x 10 -6M 7-al in O/E3-mI7 and -5.35 mV ± 0.25 mV at 1 X 10 -4 M 7-al in wtmice, P < 0.05, Student´s T-test, n = 4). The olfactory neurons fromthese animals express high levels of mI7 protein and provide a usefulmodel for examining adaptation and OR protein biochemistry. Althoughubiquitous expression of mI7 does not abolish the expression ofendogenous ORs (M72, mOR28), it dramatically affects the targeting ofM72-expressing axons. Moreover, stereotyped alterations in theposition of convergent axons to generate glomeruli suggest a model forsorting axons and glomerular formation. Supported by Howard HughesMedical Institute and NIH232 <strong>Symposium</strong> Changing the Development of Tasteand OlfactionFLAVOR PROGRAMMING DURING INFANCYMennella J.A. 1 1 Monell Chemical Senses Center, Philadelphia, PAA major factor that has inhibited progress in understanding the originof human flavor preferences is the absence of a robust, experimentalparadigm. In addition to demonstrating that experience with flavors inamniotic fluid, mothers´ milk and formulas contribute to individualdifferences in flavor preferences, we have identified a naturallyoccurring flavor variation that can be exploited to study this important,yet under-investigated, research area. Using as a model system a classof infant formulas that are hydrolyzed-protein based and havedistinctive flavors which are unpalatable to older-aged infants andadults, we found that early exposure to this formula (Nutramigen)resulted in a complete shift in hedonic tone to the flavor of this formulafrom one of absolute distaste to eager acceptance. The effects of earlyexposure were particularly persistent, leading to heightened preferencesfor sour tastes, as well as for the taste and aroma of the formula andsimilarly flavored foods, several years after the child´s last exposure.These finding provided evidence of the clearest example of a sensitiveperiod in the development of responses to flavors in humans thus faridentified. This research is supported by NIH Grant HD37119.58
233 <strong>Symposium</strong> Changing the Development of Tasteand OlfactionEARLY ENVIRONMENTAL EVENTS SHAPE THENEUROBIOLOGICAL <strong>DEVELOPMENT</strong> OF THE GUSTATORYSYSTEMHill D.L. 1 , Mangold J. 1 1 Psychology, University of Virginia,Charlottesville, VAImpressive morphological, physiological, and behavioral changescharacterize the postnatal development of the rat gustatory system.While much has been learned by studying developmental processes incontrol rats, complementary experiments using early dietary sodiumrestriction have been of great value in learning how the taste system isorganized. For example, recent findings from our laboratorydemonstrate that the dramatic diet-related differences in theorganization and size of gustatory afferent terminal fields in the ratbrainstem are due to a reorganization of terminal fields with age incontrol rats and a lack of an age-related reorganization in experimentalrats. Due to large differences in primary afferent taste responsesbetween groups, this finding suggests that activity-dependent “pruning”of axonal arbors may occur in controls but not in early sodiumrestrictedrats. This symposium presentation will provide a summary ofearly developmental diet-related changes in peripheral taste functionand changes in central structure and function. An emphasis will beplaced on more recent experiments in which we have examined theinterrelationships among the terminal fields of three gustatory nerves inthe rat NTS during normal development and how various dietarymanipulations instituted during embryonic development differentiallyalters this development. Supported by NIH grant R01 DC00407.235 <strong>Symposium</strong> Changing the Development of Tasteand OlfactionNEURAL NETWORKS INVOLVED IN OLFACTORY FEARCONDITIONING IN RATS FROM INFANCY TO ADULTHOODMouly A. 1 1 Institut des Sciences Cognitives, CNRS-Université Lyon1,Bron, Rhone, FranceIn a recent study, we investigated whether synaptic changes could bedetected at different levels of the olfactory pathways following an odorfear conditioning learning. For this, evoked potentials induced in fourrecording sites (anterior piriform cortex, posterior piriform cortex,cortical and basolateral nuclei of the amygdala) in response to electricalstimulation of the olfactory bulb were collected before odor-shocktraining (baseline) and during the retention test. The data showed thatlearning was accompanied by a lasting increase in signal amplitude incortical amygdala, and a transient increase in posterior piriform cortexand basolateral amygdala, suggesting a differential involvement of thesestructures in recognition of the learned odor. Paradoxically earlyolfactory fear conditioning causes odor preferences in rat pups during atemporally defined sensitive period. We assessed the effects of thisearly experience on adult olfactory fear conditioning. Specifically,infant rats were trained daily from 8-12-days old in an odor fearconditioning paradigm. In adulthood, these animals were trained againin the same paradigm and tested for freezing to the presentation of odoralone. In parallel, amygdala activation following acquisition wasassessed using 2-DG mapping.The data showed that, compared tocontrol animals, early trained animals presented lower levels of freezingand no detectable amygdala participation in the adult fear conditioningparadigm. This suggests that early neonatal experience has lastingconsequences on both the behavior and the neural network observed inadulthood in odor fear learning.234 <strong>Symposium</strong> Changing the Development of Tasteand OlfactionONTOGENETIC EMERGENCE OF LEARNED/NATURALFEAR AND <strong>DEVELOPMENT</strong> OF THE ODOR PATHWAY TOTHE AMYGDALASullivan R.M. 1 1 Department of Zoology, University of Oklahoma,Norman, OKInfant rats do not express fear to natural predator odors, nor do theylearn to avoid odors from olfactory fear conditioning until postnatal day10. This is coincident with the emergence of walking suggesting fearemerges when the risk of predation increases. The ability to express fearappears to be due to the functional emergence of the amygdala at PN10in both natural fear and conditioned fear. Fear to predator odor andlearned fear can be either retarded or advanced by manipulatingcorticosteroid (CORT) levels. Moreover, CORT is capable of changingthe neural pathway of the odor and presumably underlies the behavioralchange. Specifically, before the emergence of fearful behaviors (learned& unlearned), odor are relayed from the bulb to the anterior piriformcortex. For the expression of fear, either in older pups or younger pupsgiven CORT, the olfactory circuit is bulb, posterior piriform andamygdala, although a direct bulb-amygdala pathway also exists. Wewill present data suggesting odor-shock fear conditioning in 14-day-oldpups produces an odor aversion with amygdala participation when themother is absent but an odor preference with amygdala nonparticipationwhen the mother is present. The ability of maternal presence to alterolfactory fear learning appears due to the mother´s regulation of pups´CORT with maternal odor and tactile stimulation maintaining pups lowCORT levels, even during stressful presentations of 0.5-mA shock.This work is supported by NIH-NICHD, NSF-IBN and OCAST.236 <strong>Symposium</strong> Changing the Development of Tasteand OlfactionNEURAL ANALYSIS OF PREDATOR ODOR-INDUCED FEARAND EMOTIONAL MEMORYTakahashi L.K. 1 1 Psychology, University of Hawaii, Honolulu, HIInfant and adult rodents exposed to predator odor exhibit diversefear-related responses including freezing, avoidance, and stresshormone secretion. Brain structures, with prominent binding of stresshormones, such as the hippocampus and the amygdala likely play keyroles in the development and modulation of fear behavior. However,very little information is known on the neural basis of emotionallearning and memory associated with predator odor. We are especiallyinterested in the amygdala, a brain region implicated in fearconditioning using aversive electric shock, in mediating predator odorinducedemotional behavior. Our predator odor studies in adult ratshave shown that basolateral amygdalar (BLA) lesions impair theelicitation of cat odor-induced unconditioned and conditioned fearresponses. In addition, the medial amygdala (MeA), which is associatedwith aggressive and sexual behavior, plays an important role inmodulating cat odor-induced fear. MeA lesions were as effective asBLA lesions in attenuating predator odor-induced unconditioned andconditioned behavior. The specific roles of the BLA and MeA inpredator odor fear consolidation and memory retrieval were furtherevaluated using temporary brain inactivation methods. Muscimolinjections made immediately after exposure to cat odor produced asubsequent impairment in conditioned fear behavior but only wheninjected in the BLA. In contrast, the retrieval of conditioned fearbehavior was attenuated but only when lidocaine was injected into theMeA just prior to the conditioned fear test. Our novel resultsunderscore the unique participation of the BLA and MeA inconsolidation and retrieval processes underlying predator odoremotional memory. Supported by NS39406.59
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