Processing Of Chemosensory Information Research Articles

Electrophysiological and Motor Responses to Chemosensory Stimuli in Isolated Cephalopod Arms.

While there is behavioral and anatomical evidence that coleoid cephalopods use their arms to "taste" substances in the environment, the neurophysiology of chemosensation has been largely unexamined. The range and sensitivity of detectable chemosensory stimuli, and the processing of chemosensory information, are unknown. To begin to address these issues, we developed a technique for recording neurophysiological responses from isolated arms, allowing us to test responses to biologically relevant stimuli. We tested arms from both a pelagic species (Doryteuthis pealeii) and a benthic species (Octopus bimaculoides) by attaching a suction electrode to the axial nerve cord to record neural activity in response to chemical stimuli. Doryteuthis pealeii arms showed anecdotal responses to some stimuli but generally did not tolerate the preparation; tissue was nonviable within minutes ex vivo. Octopus bimaculoides arms were used successfully, with tissue remaining healthy and responsive for several hours. Arms responded strongly to fish skin extract, glycine, methionine, and conspecific skin extract but not to cephalopod ink or seawater controls. Motor responses were also observed in response to detected stimuli. These results suggest that chemosensory receptor cells on O. bimaculoides arms were able to detect environmentally relevant chemicals and drive local motor responses within the arm. Further exploration of potential chemical stimuli for O. bimaculoides arms, as well as investigations into the neural processing within the arm, could enhance our understanding of how this species uses its arms to explore its environment. While not successful in D. pealeii, this technique could be attempted with other cephalopod species, as comparative questions remain of interest.

Attending to the Chemical Senses.

Theorizing around the topic of attention and its role in human information processing largely emerged out of research on the so-called spatial senses: vision, audition, and to a lesser extent, touch. Thus far, the chemical senses have received far less research interest (or should that be attention) from those experimental psychologists and cognitive neuroscientists interested in the topic. Nevertheless, this review highlights the key role that attentional selection also plays in chemosensory information processing and awareness. Indeed, many of the same theoretical approaches/experimental paradigms that were originally developed in the context of the spatial senses, can be (and in some cases already have been) extended to provide a useful framework for thinking about the perception of taste/flavour. Furthermore, a number of those creative individuals interested in modifying the perception of taste/flavour by manipulating product-extrinsic cues (such as, for example, music in the case of sonic seasoning) are increasingly looking to attentional accounts in order to help explain the empirical phenomena that they are starting to uncover. However, separate from its role in explaining sonic seasoning, gaining a better understanding of the role of attentional distraction in modulating our eating/drinking behaviours really ought to be a topic of growing societal concern. This is because distracted diners (e.g., those who eat while watching TV, fiddling with a mobile device or smartphone, or even while driving) consume significantly more than those who mindfully pay attention to the sensations associated with eating and drinking.

Experience-Dependent Plasticity in Accessory Olfactory Bulb Interneurons following Male-Male Social Interaction.

Chemosensory information processing in the mouse accessory olfactory system guides the expression of social behavior. After salient chemosensory encounters, the accessory olfactory bulb (AOB) experiences changes in the balance of excitation and inhibition at reciprocal synapses between mitral cells (MCs) and local interneurons. The mechanisms underlying these changes remain controversial. Moreover, it remains unclear whether MC-interneuron plasticity is unique to specific behaviors, such as mating, or whether it is a more general feature of the AOB circuit. Here, we describe targeted electrophysiological studies of AOB inhibitory internal granule cells (IGCs), many of which upregulate the immediate-early gene Arc after male-male social experience. Following the resident-intruder paradigm, Arc-expressing IGCs in acute AOB slices from resident males displayed stronger excitation than nonexpressing neighbors when sensory inputs were stimulated. The increased excitability of Arc-expressing IGCs was not correlated with changes in the strength or number of excitatory synapses with MCs but was instead associated with increased intrinsic excitability and decreased HCN channel-mediated IH currents. Consistent with increased inhibition by IGCs, MCs responded to sensory input stimulation with decreased depolarization and spiking following resident-intruder encounters. These results reveal that nonmating behaviors drive AOB inhibitory plasticity and indicate that increased MC inhibition involves intrinsic excitability changes in Arc-expressing interneurons.SIGNIFICANCE STATEMENT The accessory olfactory bulb (AOB) is a site of experience-dependent plasticity between excitatory mitral cells (MCs) and inhibitory internal granule cells (IGCs), but the physiological mechanisms and behavioral conditions driving this plasticity remain unclear. Here, we report studies of AOB neuronal plasticity following male-male social chemosensory encounters. We show that the plasticity-associated immediate-early gene Arc is selectively expressed in IGCs from resident males following the resident-intruder assay. After behavior, Arc-expressing IGCs are more strongly excited by sensory input stimulation and MC activation is suppressed. Arc-expressing IGCs do not show increased excitatory synaptic drive but instead show increased intrinsic excitability. These data indicate that MC-IGC plasticity is induced after male-male social chemosensory encounters, resulting in enhanced MC suppression by Arc-expressing IGCs.

Heterogeneous effects of norepinephrine on spontaneous and stimulus-driven activity in the male accessory olfactory bulb.

Norepinephrine (NE) release has been linked to experience-dependent plasticity in many model systems and brain regions. Among these is the rodent accessory olfactory system (AOS), which is crucial for detecting and processing socially relevant environmental cues. The accessory olfactory bulb (AOB), the first site of chemosensory information processing in the AOS, receives dense centrifugal innervation by noradrenergic fibers originating in the locus coeruleus. Although NE release has been linked to behavioral plasticity through its actions in the AOB, the impacts of noradrenergic modulation on AOB information processing have not been thoroughly studied. We made extracellular single-unit recordings of AOB principal neurons in ex vivo preparations of the early AOS taken from adult male mice. We analyzed the impacts of bath-applied NE (10 μM) on spontaneous and stimulus-driven activity. In the presence of NE, we observed overall suppression of stimulus-driven neuronal activity with limited impact on spontaneous activity. NE-associated response suppression in the AOB came in two forms: one that was strong and immediate (21%) and one other that involved gradual, stimulus-dependent monotonic response suppression (47%). NE-associated changes in spontaneous activity were more modest, with an overall increase in spontaneous spike frequency observed in 25% of neurons. Neurons with increased spontaneous activity demonstrated a net decrease in chemosensory discriminability. These results reveal that noradrenergic signaling in the AOB causes cell-specific changes in chemosensory tuning, even among similar projection neurons.NEW & NOTEWORTHY Norepinephrine (NE) is released throughout the brain in many behavioral contexts, but its impacts on information processing are not well understood. We studied the impact of NE on chemosensory tuning in the mouse accessory olfactory bulb (AOB). Electrophysiological recordings from AOB neurons in ex vivo preparations revealed that NE, on balance, inhibited mitral cell responses to chemosensory cues. However, NE's effects were heterogeneous, indicating that NE signaling reshapes AOB output in a cell- and stimulus-specific manner.

Individual Neurons Confined to Distinct Antennal-Lobe Tracts in the Heliothine Moth: Morphological Characteristics and Global Projection Patterns.

To explore fundamental principles characterizing chemosensory information processing, we have identified antennal-lobe projection neurons in the heliothine moth, including several neuron types not previously described. Generally, odor information is conveyed from the primary olfactory center of the moth brain, the antennal lobe, to higher brain centers via projection neuron axons passing along several parallel pathways, of which the medial, mediolateral, and lateral antennal-lobe tract are considered the classical ones. Recent data have revealed the projections of the individual tracts more in detail demonstrating three main target regions in the protocerebrum; the calyces are innervated mainly by the medial tract, the superior intermediate protocerebrum by the lateral tract exclusively, and the lateral horn by all tracts. In the present study, we have identified, via iontophoretic intracellular staining combined with confocal microscopy, individual projection neurons confined to the tracts mentioned above, plus two additional ones. Further, using the visualization software AMIRA, we reconstructed the stained neurons and registered the models into a standard brain atlas, which allowed us to compare the termination areas of individual projection neurons both across and within distinct tracts. The data demonstrate a morphological diversity of the projection neurons within distinct tracts. Comparison of the output areas of the neurons confined to the three main tracts in the lateral horn showed overlapping terminal regions for the medial and mediolateral tracts; the lateral tract neurons, on the contrary, targeted mostly other output areas in the protocerebrum.

Chemosensory Information Processing between Keratinocytes and Trigeminal Neurons

Trigeminal fibers terminate within the facial mucosa and skin and transmit tactile, proprioceptive, chemical, and nociceptive sensations. Trigeminal sensations can arise from the direct stimulation of intraepithelial free nerve endings or indirectly through information transmission from adjacent cells at the peripheral innervation area. For mechanical and thermal cues, communication processes between skin cells and somatosensory neurons have already been suggested. High concentrations of most odors typically provoke trigeminal sensations in vivo but surprisingly fail to activate trigeminal neuron monocultures. This fact favors the hypothesis that epithelial cells may participate in chemodetection and subsequently transmit signals to neighboring trigeminal fibers. Keratinocytes, the major cell type of the epidermis, express various receptors that enable reactions to multiple environmental stimuli. Here, using a co-culture approach, we show for the first time that exposure to the odorant chemicals induces a chemical communication between human HaCaT keratinocytes and mouse trigeminal neurons. Moreover, a supernatant analysis of stimulated keratinocytes and subsequent blocking experiments with pyrodoxalphosphate-6-azophenyl-2',4'-disulfonate revealed that ATP serves as the mediating transmitter molecule released from skin cells after odor stimulation. We show that the ATP release resulting from Javanol® stimulation of keratinocytes was mediated by pannexins. Consequently, keratinocytes act as chemosensors linking the environment and the trigeminal system via ATP signaling.

Brain Response to the Luminal Nutrient Stimulation

Nutrient signals called postingestive effects are thought to be the sixth sensory signals (gut feeling). They modulate motivation to eat and subsequent food preference. Although the other five senses are well-characterized, there are still a lot of mysteries in the processing of luminal nutrient information in the brain. Development of functional MRI (fMRI) technique during the past 10 years enabled researchers to investigate brain function for the detection, perception, and processing of chemosensory information in human beings. We developed an awake rat fMRI technique to investigate whole brain function for the detection of ingested nutrients. Here we will review a recent study of luminal nutrient information in humans and in rodents.

Centrifugal telencephalic afferent connections to the main and accessory olfactory bulbs

Parallel to the olfactory system, most mammals possess an accessory olfactory or vomeronasal system. The olfactory and vomeronasal epithelia project to the main and accessory olfactory bulbs, which in turn project to adjacent areas of the telencephalon, respectively. New data indicate that projections arising from the main and accessory olfactory bulbs partially converge in the rostral telencephalon and are non-overlapping at caudal telencephalic levels. Therefore, the basal telencephalon should be reclassified in olfactory, vomeronasal, and mixed areas. On the other hand, it has been demonstrated that virtually all olfactory- and vomeronasal-recipient structures send reciprocal projections to the main and accessory olfactory bulbs, respectively. Further, non-chemosensory recipient structures also projects centrifugally to the olfactory bulbs. These feed-back projections appear to be essential modulating processing of chemosensory information. The present work aims at characterizing centrifugal projections to the main and accessory olfactory bulbs arising from olfactory, vomeronasal, mixed, and non-chemosensory recipient telencephalic areas. This issue has been addressed by using tracer injections in the rat and mouse brain. Tracer injections were delivered into the main and accessory olfactory bulbs as well as in olfactory, vomeronasal, mixed, and non-chemosensory recipient telencephalic structures. The results confirm that olfactory- and vomeronasal-recipient structures project to the main and accessory olfactory bulbs, respectively. Interestingly, olfactory (e.g., piriform cortex), vomeronasal (e.g., posteromedial cortical amygdala), mixed (e.g., the anterior medial amygdaloid nucleus), and non-chemosensory-recipient (e.g., the nucleus of the diagonal band) structures project to the main and to the accessory olfactory bulbs thus providing the possibility of simultaneous modulation and interaction of both systems at different stages of chemosensory processing.

Responses to olfactory and intranasal trigeminal stimuli: relation to the respiratory cycle

The aim of the study was to investigate whether the perception of intranasal chemosensory stimuli changes in relation to the respiratory cycle. We investigated 40 healthy subjects with normal olfactory function who participated in four sessions. The first session was used to adapt subjects to the experimental conditions, and, specifically, to train a certain breathing technique (velopharyngeal closure) which prevents intranasal respiratory air-flow. In each of the following three sessions one of three stimulants was tested, namely phenyl ethyl alcohol (PEA), hydrogen sulfide (H2S), or the trigeminal stimulant carbon dioxide (CO2). The sequence of testing the three stimulants was randomized across all participants. Sessions were separated by at least 1 day. Chemosensory event-related potentials (ERP) were recorded in response to 80 stimuli each. Following each stimulus subjects rated its intensity using a computerized visual analogue scale. Respiration was recorded using a probe in front of the subjects' mouth. While presentation of chemosensory stimuli was performed independent of the respiratory cycle, responses were averaged off-line according to the subjects' respiratory phase when the stimuli had been presented. Intensity of olfactory or trigeminal stimuli did not differ significantly in relation to the respiratory cycle. Olfactory ERP to phenylethyl alcohol were larger when stimuli were presented during inspiration. Similarly, responses to H2S tended to be larger when stimuli were presented during inspiratory phases. In addition, responses to CO2 were larger when stimuli were presented during inspiration. Differences in relation to the respiratory cycle were found specifically for early ERP components. It is important to note that the changes of chemosensory information processing were found in the absence of changes of intranasal airflow. These data indicate on an electrophysiological level that there is priming of both olfactory and trigeminally mediated sensations in relation to the respiratory cycle.

Conspecific and heterospecific alarm substance induces behavioral responses in piau fish Leporinus piau

In ostariophysan fish, the detection of alarm substance released from the skin of a conspecific or a sympatric heterospecific may elicit alarm reactions or antipredator behavioral responses. In this study, experiments were performed to characterize and quantify the behavioral response threshold of Leporinus piau, both individually and in schools, to growing dilutions of conspecific (CAS) and heterospecific skin extract (HAS). The predominant behavioral response to CAS stock stimulation was biphasic for fish held individually, with a brief initial period of rapid swimming followed by a longer period of immobility or reduced swimming activity. As the dilution of skin extract was increased, the occurrence and magnitude of the biphasic alarm response tended to decrease, replaced by a slowing of locomotion. Slowing was the most common antipredator behavior, observed in 62.5% of animals submitted to HAS stimulation. School cohesion, measured as proximity of fish to the center of the school, and swimming activity near the water surface significantly increased after exposure to CAS when compared with the control group exposed to distilled water. Histological analysis of the epidermis revealed the presence of Ostariophysi-like club cells. The presence of these cells and the behavioral responses to conspecific and heterospecific skin extract stimulation suggest the existence of a pheromone alarm system in L. piau similar to that in Ostariophysi, lending further support for the neural processing of chemosensory information in tropical freshwater fish.

In vivo vomeronasal stimulation reveals sensory encoding of conspecific and allospecific cues by the mouse accessory olfactory bulb

The rodent vomeronasal system plays a critical role in mediating pheromone-evoked social and sexual behaviors. Recent studies of the anatomical and molecular architecture of the vomeronasal organ (VNO) and of its synaptic target, the accessory olfactory bulb (AOB), have suggested that unique features underlie vomeronasal sensory processing. However, the neuronal representation of pheromonal information leading to specific behavioral and endocrine responses has remained largely unexplored due to the experimental difficulty of precise stimulus delivery to the VNO. To determine the basic rules of information processing in the vomeronasal system, we developed a unique preparation that allows controlled and repeated stimulus delivery to the VNO and combined this approach with multisite recordings of neuronal activity in the AOB. We found that urine, a well-characterized pheromone source in mammals, as well as saliva, activates AOB neurons in a manner that reliably encodes the donor animal's sexual and genetic status. We also identified a significant fraction of AOB neurons that respond robustly and selectively to predator cues, suggesting an expanded role for the vomeronasal system in both conspecific and interspecific recognition. Further analysis reveals that mixed stimuli from distinct sources evoke synergistic responses in AOB neurons, thereby supporting the notion of integrative processing of chemosensory information.

Mammalian olfactory receptors

Perception of chemical stimuli from the environment is essential to most animals; accordingly, they are equipped with a complex olfactory system capable of receiving a nearly unlimited number of odorous substances and pheromones. This enormous task is accomplished by olfactory sensory neurons (OSNs) arranged in several chemosensory compartments in the nose. The sensitive and selective responsiveness of OSNs to odorous molecules and pheromones is based on distinct receptors in their chemosensory membrane; consequently, olfactory receptors play a key role for a reliable recognition and an accurate processing of chemosensory information. They are therefore considered as key elements for an understanding of the principles and mechanisms underlying the sense of smell. The repertoire of olfactory receptors in mammals encompasses hundreds of different receptor types which are highly diverse and expressed in distinct subcompartments of the nose. Accordingly, they are categorized into several receptor families, including odorant receptors (ORs), vomeronasal receptors (V1Rs and V2Rs), trace amine-associated receptors (TAARs), formyl peptide receptors (FPRs), and the membrane guanylyl cyclase GC-D. This large and complex receptor repertoire is the basis for the enormous chemosensory capacity of the olfactory system.

Olfactory conditioning in the zebrafish ( Danio rerio)

The zebrafish olfactory system is an attractive model for studying neural processing of chemosensory information. Here we characterize zebrafish olfactory behaviors and their modification through learning, using an apparatus consisting of a circular flow-through tank that allows controlled administration of odorants. When exposed to the amino acids l-alanine and l-valine, naive zebrafish responded with appetitive swimming behavior, which we measured as the number of >90° turns made during 30 s observation periods. Such appetitive responses were not observed when naive zebrafish were exposed to an unnatural odorant, phenylethyl alcohol (PEA). Repeated pairing of amino acids or PEA (conditioned stimuli, CS) with food flakes (unconditioned stimuli; UCS) increased odorant-evoked appetitive swimming behavior in all fish tested. The zebrafish also learned to restrict this behavior to the vicinity of a feeding ring, through which UCS were administered. When both nares were temporarily occluded, conditioned fish failed to respond to odorants, confirming that these behaviors were mediated by olfaction. These results represent the first demonstration of a classically conditioned appetitive response to a behaviorally neutral odorant in fish. Furthermore, they complement recent demonstrations of conditional place preferences in fish. By virtue of its robustness and simplicity, this method will be a useful tool for future research into the biological basis of olfactory learning in zebrafish.

Acceleration of chemo-sensory information processing using transient features

A snapshot from the steady-state response of chemical sensors conveys, on average, more mature and relevant information regarding the analyte than a snapshot from the transient can provide. Nevertheless, time constraints in many applications make it infeasible to wait for and extract steady-state features. Substituting them by transient ones is the only viable solution to accelerate odor processing. Based on measurements recorded from metal-oxide sensors, we point to a correlation between a transient feature and the steady-state resistance that are observed in response to fixed analyte concentration. We utilize this correlation to expedite standard quantification and classification substantially while ensuring the performance that the steady-state feature can provide.

Changes in olfactory function in pregnancy and postpartum

Objective To track changes in olfactory performance during pregnancy and the postpartum. Methods In a prospective study the olfactory function of 38 pregnant women was assessed at about 12, 21, and 36 weeks of pregnancy and 7 weeks after delivery. A control group of 46 nonpregnant women were also asked to rate the intensity and hedonic tone of 10 “natural” odors. Results There was no difference in olfactory performance between the women in the first trimester of pregnancy and the controls, but at approximately 36 weeks of pregnancy the pregnant women experienced a decreased odor threshold compared with the nonpregnant controls, and this decrease was still present after delivery. There was no significant difference between the groups in capacity for odor discrimination or odor identification. “Objective” decreases were observed even though pregnant women rated their olfactory sensitivity significantly higher than the controls. Although the 2 groups did not differ with regard to intensity ratings of the 10 “natural” odors, there were differences with regard to the hedonic ratings. Conclusion Pregnancy is accompanied by changes in olfactory performance. Changes in hedonic odor ratings indicate a potential embryo-protective mechanism. The discrepancy between “objective” and “subjective” olfactory function may relate to changes in the cognitive processing of chemosensory information during pregnancy.

Effects of age on feeding behavior and chemosensory processing in the pond snail, Lymnaea stagnalis

This study used behavioral and electrophysiological techniques to examine age-related changes in the feeding behavior and chemosensory processing in the pond snail, Lymnaea stagnalis. Increasing age was associated with a 50% decrease in long-term food consumption. Analysis of short-term sucrose-evoked feeding bouts showed an age-related increase in the number of animals that failed to respond to the stimulus. Of the animals that did respond increasing age was associated with a decrease in the number of sucrose-evoked bites and a increase in the duration of the swallow phase. These changes were observed with both 0.01 and 0.05 M sucrose stimuli but were not seen when 0.1 M sucrose was used as the stimulus. Electrophysiological analysis of the chemosensory pathway in semi-intact lip-CNS preparations failed to demonstrate a significant change in the neuronal information entering the cerebral ganglia from the lips via the median lip nerve, but did demonstrate an age-related deficit in the neuronal output from the cerebral ganglia. This deficit was also dependent on the sucrose concentration and mirrored the concentration-dependent changes in feeding behavior. In summary, aging appeared to affect central but not peripheral processing of chemosensory information and suggests that this deficit contributes to the age-related changes in feeding behavior.

Neurobiology of the Gustatory-Salivary Reflex

All neural information resulting from chemical stimulation of taste buds in the oral cavity, pharynx and larynx travels via the facial (VII), glossopharyngeal (IX) and vagus (X) nerves to terminate in the nucleus of the solitary tract (NST) in the brainstem. The NST is responsible for initial processing and distribution of chemosensory information. At higher relays in the central nervous system the processes of detection, discrimination and affective responses occur resulting in the sensation we call taste and the behavioral reactions to that sensation. In addition, the NST connects to efferent motor systems involved in oral facial motor reflexes and systems controlling the initiation and flow of saliva. Thus, the NST plays a pivotal role in the neural processing of chemosensory information derived from stimulation of taste buds. Beginning in 1961 (Pfaffmann et al., 1961) a large number of investigators in different laboratories have examined the NST using anatomical, and neurophysiological techniques. The topographical projections of the VII, IX and Xth nerves conveying sensory information to the NST have been determined using different methods in several species (Torvik, 1956; Norgren, 1981; Whitehead and Frank, 1983; Hamilton and Norgren, 1984). The morphology of the NST has been studied and the neuronal architecture defined (Whitehead, 1988). Neurons in the NST have been described as belonging to three major anatomical types—multipolar, elongate and ovoid (Whitehead, 1988; Lasiter and Kachele, 1988; King and Bradley, 1994; Mistretta and Labyak, 1994), and using immunocytochemistry the presence of GABA and other neuropeptides has been described (Lasiter and Kachele, 1988; Barry et al., 1993).

Central localization of plasticity involved in appetitive conditioning inLymnaea

Learning to associate a conditioned (CS) and unconditioned stimulus (US) results in changes in the processing of CS information. Here, we address directly the question whether chemical appetitive conditioning of Lymnaea feeding behavior involves changes in the peripheral and/or central processing of the CS by using extracellular recording techniques to monitor neuronal activity at two stages of the sensory processing pathway. Our data show that appetitive conditioning does not affect significantly the overall CS response of afferent nerves connecting chemosensory structures in the lips and tentacles to the central nervous system (CNS). In contrast, neuronal output from the cerebral ganglia, which represent the first central processing stage for chemosensory information, is enhanced significantly in response to the CS after appetitive conditioning. This demonstrates that chemical appetitive conditioning in Lymnaea affects the central, but not the peripheral processing of chemosensory information. It also identifies the cerebral ganglia of Lymnaea as an important site for neuronal plasticity and forms the basis for detailed cellular studies of neuronal plasticity.

Neuropeptide Y in the olfactory system, forebrain and pituitary of the teleost, Clarias batrachus

Distribution of neuropeptide Y (NPY)-like immunoreactivity in the forebrain of catfish Clarias batrachus was examined with immunocytochemistry. Conspicuous immunoreactivity was seen in the olfactory receptor neurons (ORNs), their projections in the olfactory nerve, fascicles of the olfactory nerve layer in the periphery of bulb and in the medial olfactory tracts as they extend to the telencephalic lobes. Ablation of the olfactory organ resulted in loss of immunoreactivity in the olfactory nerve layer of the bulb and also in the fascicles of the medial olfactory tracts. This evidence suggests that NPY may serve as a neurotransmitter in the ORNs and convey chemosensory information to the olfactory bulb, and also to the telencephalon over the extrabulbar projections. In addition, network of beaded immunoreactive fibers was noticed throughout the olfactory bulb, which did not respond to ablation experiment. These fibers may represent centrifugal innervation of the bulb. Strong immunoreactivity was encountered in some ganglion cells of nervus terminalis. Immunoreactive fibers and terminal fields were widely distributed in the telencephalon. Several neurons of nucleus entopeduncularis were moderately immunoreactive; and a small population of neurons in nucleus preopticus periventricularis was also labeled. Immunoreactive terminal fields were particularly conspicuous in the preoptic, the tuberal areas, and the periventricular zone around the third ventricle and inferior lobes. NPY immunoreactive cells and fibers were detected in all the lobes of the pituitary gland. Present results describing the localization of NPY in the forebrain of C. batrachus are in concurrence with the pattern of the immunoreactivity encountered in other teleosts. However, NPY in olfactory system of C. batrachus is a novel feature that suggests a role for the peptide in processing of chemosensory information.

Appetitive and Aversive Olfactory Learning in Humans Studied Using Event-Related Functional Magnetic Resonance Imaging

We combined event-related functional magnetic resonance imaging (fMRI) with olfactory classical conditioning to differentiate the neural responses evoked during appetitive and aversive olfactory learning. Three neutral faces [the conditioned stimuli (CS+)] were repetitively paired with pleasant, neutral, or unpleasant odors [the unconditioned stimuli (UCS)] in a partial reinforcement schedule. A fourth face was never paired to odor [the nonconditioned stimulus (CS-)]. Learning-related neural activity, comparing unpaired (face only) CS+ stimuli with CS-, showed valence-independent activations in rostral and caudal orbitofrontal cortex (OFC). Medial OFC responded to the appetitive (app) CS+, whereas lateral OFC responded to the aversive (av) CS+. Within nucleus accumbens, neural responses showed divergent activation profiles that increased with time in response to the appCS+ but decreased in response to the avCS+. In posterior amygdala, responses were elicited by the appCS+, which habituated over time. In temporal piriform cortex, neural responses were evoked by the avCS+, which progressively increased with time. These results highlight regional and temporal dissociations during olfactory learning and imply that emotionally salient odors can engender cross-modal associative learning. Moreover, the findings suggest that the role of human primary (piriform) and secondary olfactory cortices transcends their function as mere intermediaries of chemosensory information processing.