Role In Olfactory Processing Research Articles

Association of Exercise with Better Olfactory Performance and Higher Functional Connectivity Between the Olfactory Cortex and the Prefrontal Cortex: A Resting-State Functional Near-Infrared Spectroscopy Study.

Background: Olfactory deterioration is suggested to be a predictor of some neurodegenerative diseases. Recent studies indicate that physical exercise has a positive relationship with olfactory performance, and a subregion in the prefrontal cortex (PFC) may play an important role in olfactory processing. The PFC is not only related to olfactory function but it also engages in complex functions such as cognition and emotional processing. Methodology: Our study compared the functional connectivity between the olfactory cortex and the PFC in healthy individuals who exercised regularly and healthy persons who did not. Those who exercised more than three times/week for at least 30 min each time were considered the exercise group, and those who did not meet this exercise criteria were considered the nonexercise group. We also assessed their odor threshold. Participants were aged 55 years or older, and the two groups were balanced for age, sex, body mass index, and educational level. Results: We found that compared with individuals who did not exercise, exercisers had a significantly lower threshold for detecting odors. In addition, the olfactory cortex had stronger connectivity with the PFC in exercisers than in nonexercisers. More specifically, when the PFC was grouped into three subregions, namely, the ventrolateral prefrontal cortex (VLPFC), dorsolateral prefrontal cortex (DLPFC), and frontopolar cortex (FPA), Pearson correlation analysis revealed stronger connectivity between the VLPFC and the orbitofrontal cortex (OFC), between the OFC and the FPA, and between the left and right OFC hemispheres in the exercisers. In addition, Granger causality indicated higher directional connectivity from the DLPFC to the OFC in exercisers than in nonexercisers. Conclusion: Our findings indicated that the exercise group not only had better olfactory performance but also had stronger functional connectivity between the olfactory cortex and the PFC than nonexercise group.

Uncovering the Chemosensory System of a Subterranean Termite, Odontotermes formosanus (Shiraki) (Isoptera: Termitidae): Revealing the Chemosensory Genes and Gene Expression Patterns.

Termites are eusocial insects. Chemical signals between colony members are crucial to the smooth running of colony operations, but little is known about their olfactory system and the roles played by various chemosensory genes in this process. Chemosensory genes are involved in basic olfactory perception in insects. Odontotermes formosanus (Shiraki) is one of the most damaging pests to agricultural crops, forests, and human-made structures. To better understand the olfactory system and the genes involved in olfactory processing in O. formosanus, we produced a transcriptome of worker termites. In this study, we identified 13 OforOBPs, 1 OforCSP, 15 OforORs, 9 OforGRs, and 4 OforSNMPs. Multiple sequence alignments were used in the phylogenetic study, which included data from other termite species and a wide variety of insect species. Moreover, we also investigated the mRNA expression levels using qRT-PCR. The significantly high expression levels of OforCSP1, OforOBP2, OforOR1, and OforSNMP1 suggest that these genes may play important roles in olfactory processing in termite social behavior, including caste differentiation, nestmate and non-nestmate discrimination, and the performance of colony operations among members. Our research establishes a foundation for future molecular-level functional studies of chemosensory genes in O. formosanus, which might lead to the identification of novel targets for termite integrated pest management.

The potential for clinical application of automatic quantification of olfactory bulb volume in MRI scans using convolutional neural networks

The olfactory bulbs (OBs) play a key role in olfactory processing; their volume is important for diagnosis, prognosis and treatment of patients with olfactory loss. Until now, measurements of OB volumes have been limited to quantification of manually segmented OBs, which is a cumbersome task and makes evaluation of OB volumes in large scale clinical studies infeasible. Hence, the aim of this study was to evaluate the potential of our previously developed automatic OB segmentation method for application in clinical practice and to relate the results to clinical outcome measures.To evaluate utilization potential of the automatic segmentation method, three data sets containing MR scans of patients with olfactory loss were included. Dataset 1 (N = 66) and 3 (N = 181) were collected at the Smell and Taste Center in Ede (NL) on a 3 T scanner; dataset 2 (N = 42) was collected at the Smell and Taste Clinic in Dresden (DE) on a 1.5 T scanner. To define the reference standard, manual annotation of the OBs was performed in Dataset 1 and 2. OBs were segmented with a method that employs two consecutive convolutional neural networks (CNNs) that the first localize the OBs in an MRI scan and subsequently segment them.In Dataset 1 and 2, the method accurately segmented the OBs, resulting in a Dice coefficient above 0.7 and average symmetrical surface distance below 0.3 mm. Volumes determined from manual and automatic segmentations showed a strong correlation (Dataset 1: r = 0.79, p < 0.001; Dataset 2: r = 0.72, p = 0.004). In addition, the method was able to recognize the absence of an OB. In Dataset 3, OB volumes computed from automatic segmentations obtained with our method were related to clinical outcome measures, i.e. duration and etiology of olfactory loss, and olfactory ability. We found that OB volume was significantly related to age of the patient, duration and etiology of olfactory loss, and olfactory ability (F(5, 172) = 11.348, p < 0.001, R2 = 0.248).In conclusion, the results demonstrate that automatic segmentation of the OBs and subsequent computation of their volumes in MRI scans can be performed accurately and can be applied in clinical and research population studies. Automatic evaluation may lead to more insight in the role of OB volume in diagnosis, prognosis and treatment of olfactory loss.

The microbiome–nose–brain axis in health and disease

Growing evidence implicates the bacterial populations in the nose as an important factor for personal and global health. Here, we provide a brief overview of the nasal microbiome and speculate on its potential roles in olfactory processing and neurodegeneration, with a particular focus on Parkinson's disease (PD).

Olfactory bulb volume and cortical thickness evolve during sommelier training

Brain plasticity is essential for experts to acquire the abilities they need. Sommeliers are olfaction experts who display differences in olfactory regions in the brain that correlate with greater olfactory abilities. While most studies on this topic are cross‐sectional, we used a longitudinal design and invited 17 sommelier students at the start and end of their training then to compare them to 17 control students to study the effects of training‐related brain plasticity. After a year and a half, 5 sommelier students and 4 control students dropped out, leading to 12 sommelier students versus 13 controls. We used magnetic resonance imaging to measure cortical thickness and olfactory bulb volume, as this structure plays a crucial role in olfactory processing. We used the Sniffin' Sticks test to evaluate olfactory performance. During training, olfactory bulb volume increased in sommelier students while there was no significant change in the control group. We also observed that thickness of right entorhinal cortex increased, and cortical thickness decreased in other cerebral regions. Our olfactory tests did not reveal any significant changes in sommelier students. In conclusion, this is the first longitudinal study to report an increase in olfactory bulb volume in olfaction experts in line with the notion of effects of ecological training‐related brain plasticity. The mixed results about cortical thickness might be explained by a “overproduction‐pruning” model of brain plasticity, according to which the effects of training‐related plasticity are non‐linear and simultaneously involve different processes.

Insula and Olfaction: A Literature Review and Case Report

(1) Background: It is well established that the insula is involved in olfaction, though its specific role in olfactory processing remains uncertain. In this paper, we first review the current literature on the insula and olfaction. Then, we describe the case of a 56-year-old man with a left insular cavernoma that caused olfactory disturbances. (2) Results: Structural neuroimaging studies suggest that insular gray matter volume is related to olfactory function, and functional neuroimaging shows that various types of stimuli lead to either lateralized or bilateral insular activations. Studies using electro-cortical stimulation reveal a specific region of the insular cortex, around the central insular sulcus, that could be related to unpleasant odor processing. Previous cases of insular lesions leading to olfactory disturbances suggest that left-sided insular lesions may more frequently lead to olfactory changes. In our patient with a left insular cavernoma, odors that were previously perceived as pleasant started smelling unpleasant and were hard to distinguish. Despite these subjective complaints, olfactory function assessed with the Sniffin’ Sticks test was normal. (3) Conclusions: Current tests may not be sensitive to all types of olfactory impairments associated with insular damage, and further studies should be conducted to develop olfactory tests assessing the hedonic appreciation of odors.

CB1 Receptors in the Anterior Piriform Cortex Control Odor Preference Memory

The retrieval of odor-related memories shapes animal behavior. The anterior piriform cortex (aPC) is the largest part of the olfactory cortex, and it plays important roles in olfactory processing and memory. However, it is still unclear whether specific cellular mechanisms in the aPC control olfactory memory, depending on the appetitive or aversive nature of the stimuli involved. Cannabinoid-type 1 (CB1) receptors are present in the aPC (aPC-CB1), but their potential impact on olfactory memory was never explored. Here, we used a combination of behavioral, genetic, anatomical, and electrophysiological approaches to characterize the functions of aPC-CB1 receptors in the regulation of appetitive and aversive olfactory memory. Pharmacological blockade or genetic deletion of aPC-CB1 receptors specifically impaired the retrieval of conditioned odor preference (COP). Interestingly, expression of conditioned odor aversion (COA) was unaffected by local CB1 receptor blockade, indicating that the role of aPC endocannabinoidsignaling is selective for retrieval of appetitive memory. Anatomical investigations revealed thatCB1 receptors are highly expressed on aPC GABAergic interneurons, and exvivo electrophysiological recordings showed that their pharmacological activation reduces miniature inhibitory post-synaptic currents (mIPSCs) onto aPC semilunar (SL), but not pyramidal principal neurons. COP retrieval, but not COA, was associated with a specific CB1-receptor-dependent decrease of mIPSCs in SL cells. Altogether, these data indicate that aPC-CB1 receptor-dependent mechanisms physiologically control the retrieval of olfactory memory, depending on odor valence and engaging modulation of local inhibitory transmission.

The Functional Role of Olfactory Bulb Granule Cell Subtypes Derived From Embryonic and Postnatal Neurogenesis.

It has been shown in a variety of mammalian species that sensory experience can regulate the development of various structures, including the retina, cortex, hippocampus, and olfactory bulb (OB). In the mammalian OB, the development of dendrites in excitatory projection neurons, such as mitral and tufted cells, is well known to be dependent on odor experience. Odor experience is also involved in the development of another OB population, a subset of inhibitory interneurons that are generated in the ventricular-subventricular zone throughout life and differentiate into granule cells (GCs) and periglomerular cells. However, the roles that each type of interneuron plays in the control of olfactory behaviors are incompletely understood. We recently found that among the various types of OB interneurons, a subtype of GCs expressing the oncofetal trophoblast glycoprotein 5T4 gene is required for odor detection and discrimination behaviors. Our results suggest that embryonic-born OB interneurons, including 5T4-positive GCs, play a crucial role in fundamental olfactory responses such as simple odor detection and discrimination behaviors. By contrast, postnatal- and adult-born OB interneurons are important in the learning of more complicated olfactory behaviors. Here, we highlight the subtypes of OB GCs, and discuss their roles in olfactory processing and behavior, with a particular focus on the relative contributions of embryonically and postnatally generated subsets of GCs in rodents.

Olfactory stimulation induces cerebellar vermis activation during sexual learning in male rats

The cerebellum is a complex structure mainly recognized for its participation in motor activity and balance, and less understood for its role in olfactory processing. Herein, we assessed Fos immunoreactivity (Fos-IR) in the cerebellar vermis following exposure to different odors during sexual training in male rats. Males were allowed to copulate for either one, three or five sessions. One day after the corresponding session they were exposed during 60 min to woodshaving that was either: clean (Control), sprayed with almond scent (Alm) or from cages of sexually receptive females (RF). The vermis of the cerebellum was removed, cut in sagittal sections and analyzed for Fos-IR to infer activation. Our results showed that the cerebellum responded with more Fos-IR in the Alm and RF groups as compared to Control. More copulatory sessions resulted in more odor-induced Fos-IR, especially in the RF group. Accordingly, we discuss possible mechanisms on how the cerebellum mediates processing of both unconditioned and conditioned odors, and how sexual experience accelerates such process.

Recovery of glomerular morphology in the olfactory bulb of young mice after disruption caused by continuous odorant exposure

Olfactory glomeruli are the first synaptic site of the olfactory system and are formed by the convergence of axons of the same type of sensory neurons onto the olfactory bulbs of the brain. Although the anatomical organization of glomeruli is conserved across species, their particular role in olfactory processing remains uncertain. We studied the composition and maintenance of glomeruli by means of a genetic model, mI7-IRES-tauGFP knock-in young mice, where the cytoskeleton of sensory neurons expressing the mI7 olfactory receptor is tagged with green fluorescent protein. Animals were continuously exposed to heptaldehyde, a cognate ligand of the mI7 receptor, from postnatal days 5–10. We hypothesized that continuous odorant exposure will induce changes in glomerular morphology, and that this can be recovered if the normal odorant environment is reestablished within the early postnatal period. We assessed changes in the distribution of mI7 axons in glomerular morphology, as well as possible changes in the number of the mI7 olfactory sensory neurons. Following odorant exposure the well-defined convergence of mI7 fibers into a single glomerulus was disrupted, producing numerous neighboring glomeruli partially innervated by mI7 fibers. After the normal odor environment was reestablished the number of glomeruli partially innervated by mI7 fibers decreased significantly. Moreover, we found that multiple supernumerary mI7 glomeruli were formed. Our results confirm the significant role of sensory input in glomerular formation and maintenance. Additionally, we show that the developing olfactory system actively maintains glomerular morphology, suggesting the importance of this for olfactory processing.

Intraglomerular lateral inhibition promotes spike timing variability in principal neurons of the olfactory bulb.

The activity of mitral and tufted cells, the principal neurons of the olfactory bulb, is modulated by several classes of interneurons. Among them, diverse periglomerular (PG) cell types interact with the apical dendrites of mitral and tufted cells inside glomeruli at the first stage of olfactory processing. We used paired recording in olfactory bulb slices and two-photon targeted patch-clamp recording in vivo to characterize the properties and connections of a genetically identified population of PG cells expressing enhanced yellow fluorescent protein (EYFP) under the control of the Kv3.1 potassium channel promoter. Kv3.1-EYFP(+) PG cells are axonless and monoglomerular neurons that constitute ∼30% of all PG cells and include calbindin-expressing neurons. They respond to an olfactory nerve stimulation with a short barrage of excitatory inputs mediated by mitral, tufted, and external tufted cells, and, in turn, they indiscriminately release GABA onto principal neurons. They are activated by even the weakest olfactory nerve input or by the discharge of a single principal neuron in slices and at each respiration cycle in anesthetized mice. They participate in a fast-onset intraglomerular lateral inhibition between principal neurons from the same glomerulus, a circuit that reduces the firing rate and promotes spike timing variability in mitral cells. Recordings in other PG cell subtypes suggest that this pathway predominates in generating glomerular inhibition. Intraglomerular lateral inhibition may play a key role in olfactory processing by reducing the similarity of principal cells discharge in response to the same incoming input.

Oxytocin, social cognition and psychiatry.

Oxytocin (OT) has an ancient role in modulating sensing and responding to social stimuli, from nematodes to man. OT regulates not only mammalian labor and nursing, but also maternal behavior. There is increasing evidence that OT influences human parenting and mediates the impact of parenting on infant socio-emotional development (Rilling and Young, 2014). Preclinical studies suggest that OT increases the salience and rewarding value of social stimuli. OT acts in the rodent amygdala to enhance the salience of social olfactory cues. thereby facilitating social recognition, in the striatum to mediate social reward and in the hippocampus to enhance signal to noise neurotransmission. These fundamental processes likely contribute to more complex OT-mediated behaviors, including social bonding. The effects of OT on social information processing in rodents make it an enticing pharmacological target for enhancing social cognition. However, two issues introduce skepticism for translating the compelling preclinical observations into effective pharmacotherapies to improve social functioning in psychiatric disorders, including autism and schizophrenia: (1) rodents use olfaction as the primary social perception modality, while primates rely more on visual and auditory social perception; (2) little is known regarding the pharmacokinetics of current OT administration methods or the impact of chronic OT treatment. Recent studies from our laboratory address these issues. A common polymorphism in the human OT receptor (OXTR) gene predicts face recognition skills in families with a child with autism. This effect was present in all family members in two independent populations, yet there was no evidence of an association with autism diagnosis (Skuse et al, 2014). This study supports a role for the OT system in human visual social information processing analogous to its role in olfactory processing in rodents. Nonhuman primates are useful for exploring the mechanisms of intranasal OT (IN-OT) administration. We showed that OT administered nasally by a pediatric nebulizer modestly elevates OT in the cerebrospinal fluid of anesthetized macaques (Modi et al, 2014). Importantly, intranasal OT also robustly elevated plasma OT for an extended period of time. Thus IN-OT may increase brain OT signaling, but peripheral mechanisms should be considered. Comparative studies of brain OXTR distribution in primates reveal the potential mechanisms by which OT modulates social information processing (Freeman et al, 2014a, 2014b). In all primate species examined, OXTRs are concentrated in cholinergic regions involved in visual and auditory processing, including the nucleus basalis of Meynert, which coordinates neural activity in the amygdala and cortex, thereby modulating attention to visual cues. IN-OT may enhance some aspects of social cognition through the mechanisms described above, but the efficacy may be limited by brain penetration. Stimulating endogenous central OT release pharmacologically is a viable alternative for increasing OT neural signaling. Melanocortin receptor agonists stimulate OT release from hypothalamic slices, potentiate OT release in the ventral striatum, and enhance OT-dependent behavior in prairie voles (unpublished data). Neonatal melanocortin receptor activation acutely activates OT neurons, and daily treatment for the first week of life enhances adult social bonding in prairie voles (Barrett et al, 2014). Thus, the OT system remains an attractive target for clinically enhancing social cognition, and alternative pharmacological strategies for enhancing OT neurotransmission should be explored.

Presynaptic Inhibition of Olfactory Sensory Neurons: New Mechanisms and Potential Functions

Presynaptic inhibition is the suppression of neurotransmitter release from a neuron by inhibitory input onto its presynaptic terminal. In the olfactory system, the primary sensory afferents from the olfactory neuroepithelium to the brain's olfactory bulb are strongly modulated by a presynaptic inhibition that has been studied extensively in brain slices and in vivo. In rodents, this inhibition is mediated by γ-amino butyric acid (GABA) and dopamine released from bulbar interneurons. The specialized GABAergic circuit is now well understood to include a specific subset of GAD65-expressing periglomerular interneurons that stimulate presynaptic GABAB receptors to reduce presynaptic calcium conductance. This inhibition is organized to permit the selective modulation of neurotransmitter release from specific populations of olfactory sensory neurons based on their odorant receptor expression, includes specialized microcircuits to create a tonically active inhibition and a separate feedback inhibition evoked by sensory input, and can be modulated by centrifugal projections from other brain regions. Olfactory nerve output can also be modulated by dopaminergic circuitry, but this literature is more difficult to interpret. Presynaptic inhibition of olfactory afferents may extend their dynamic range but could also create state-dependent or odorant-specific sensory filters on primary sensory representations. New directions exploring this circuit's role in olfactory processing are discussed.

Diverse populations of intrinsic cholinergic interneurons in the mouse olfactory bulb

Cholinergic activities affect olfactory bulb (OB) information processing and associated learning and memory. However, the presence of intrinsic cholinergic interneurons in the OB remains controversial. As a result, morphological and functional properties of these cells are largely undetermined. We characterized cholinergic interneurons using transgenic mice that selectively mark choline acetyltransferase (ChAT)-expressing cells and immunolabeling. We found a significant number of intrinsic cholinergic interneurons in the OB. These interneurons reside primarily in the glomerular layer (GL) and external plexiform layer (EPL) and exhibit diverse distribution patterns of nerve processes, indicating functional heterogeneity. Further, we found these neurons express ChAT and vesicular acetylcholine transporter (VAChT), but do not immunoreact to glutamatergic, GABAergic or dopaminergic markers and are distinct from calretinin-expressing interneurons. Interestingly, the cholinergic population partially overlaps with the calbindin D28K-expressing interneuron population, revealing the neurotransmitter identity of this sub-population. Additionally, we quantitatively determined the density of VAChT labeled cholinergic nerve fibers in various layers of the OB, as well as the intensity of VAChT immunoreactivity within the GL, suggesting primary sites of cholinergic actions. Taken together, our results provide clear evidence showing the presence of a significant number of cholinergic interneurons and that these morphologically and distributionally diverse interneurons make up complex local cholinergic networks in the OB. Thus, our results suggest that olfactory information processing is modulated by dual cholinergic systems of local interneuron networks and centrifugal projections.

Discrimination Training with Multimodal Stimuli Changes Activity in the Mushroom Body of the Hawkmoth Manduca sexta

BackgroundThe mushroom bodies of the insect brain play an important role in olfactory processing, associative learning and memory. The mushroom bodies show odor-specific spatial patterns of activity and are also influenced by visual stimuli.Methodology/Principal FindingsFunctional imaging was used to investigate changes in the in vivo responses of the mushroom body of the hawkmoth Manduca sexta during multimodal discrimination training. A visual and an odour stimulus were presented either together or individually. Initially, mushroom body activation patterns were identical to the odour stimulus and the multimodal stimulus. After training, however, the mushroom body response to the rewarded multimodal stimulus was significantly lower than the response to the unrewarded unimodal odour stimulus, indicating that the coding of the stimuli had changed as a result of training. The opposite pattern was seen when only the unimodal odour stimulus was rewarded. In this case, the mushroom body was more strongly activated by the multimodal stimuli after training. When no stimuli were rewarded, the mushroom body activity decreased for both the multimodal and unimodal odour stimuli. There was no measurable response to the unimodal visual stimulus in any of the experiments. These results can be explained using a connectionist model where the mushroom body is assumed to be excited by olfactory stimulus components, and suppressed by multimodal configurations.ConclusionsDiscrimination training with multimodal stimuli consisting of visual and odour cues leads to stimulus specific changes in the in vivo responses of the mushroom body of the hawkmoth.

Cholinergic Modulation of Neuronal Excitability in the Accessory Olfactory Bulb

The accessory olfactory bulb (AOB), the first relay of chemosensory information in the Vomeronasal system, receives extensive cholinergic innervation from the basal forebrain. Cholinergic modulation of neuronal activity in the olfactory bulb has been hypothesized to play an important role in olfactory processing; however, little is known about the cellular actions of acetylcholine (ACh) within the AOB. Here using in vitro slice preparation, we show that muscarinic acetylcholine receptor (mAChR) activation increases neuronal excitability of granule and mitral/tufted cells (GCs and MCs) in the AOB. Activation of mAChRs increased excitability of GCs by three distinct mechanisms: induction of a long-lasting depolarization, activation of a slow afterdepolarization (sADP), and an increase in excitatory glutamatergic input due to MC depolarization. The depolarization and sADP were elicited by the selective agonist 4-[[[(3-chlorophenyl)amino]carbonyl]oxy]-N,N,N-trimethyl-2-butyn-1-aminium chloride (100 μM) and blocked by low concentrations of pirenzepine (300 nM), indicating that they result from activation of M1-like mAChRs. In contrast, cholinergic stimulation increased the excitability of MCs via recruitment of nicotinic AChRs (nAChRs) and M1-like mAChRs. Submaximal activation of these receptors, however, decreased the excitability of MCs. Surprisingly, we found that unlike GCs in the main olfactory bulb, GCs in the AOB are excited by mAChR activation in young postnatal neurons, suggesting marked differences in cholinergic regulation of development between these two regions of the olfactory bulb.

A diversity of putative carboxylesterases are expressed in the antennae of the noctuid moth Spodoptera littoralis

Recent studies have suggested that pheromone-degrading enzymes belonging to the carboxylesterase family could play a role in the dynamics of the olfactory response to acetate sex pheromones in insects. Bioinformatic analyses of a male antennal expressed sequence tag library allowed the identification of 19 putative esterase genes expressed in the antennae of the moth Spodoptera littoralis. Phylogenetic analysis revealed that these genes belong to different insect esterase clades, defined by their putative cellular localization and substrate preferences. Interestingly, two of the 19 genes appeared to be antennal specific, suggesting a specific role in olfactory processing. This high esterase diversity suggested that the antennae are the location for intense esterase-based metabolism, against potentially a large range of exogenous and endogenous molecules.

Specific spatio-temporal activities in the cerebral ganglion of Incilaria fruhstorferi in response to superior and inferior tentacle nerve stimulation

In terrestrial gastropod mollusks (slugs and snails), olfaction is the dominant sensory modality guiding various kinds of behavior. Anatomical studies indicate that olfactory information is processed in the brain (the cerebral ganglion) in two lobes in particular: the procerebrum (PC) and the metacerebrum (MtC). This implies that olfactory functions emerge from simultaneous and cooperative processing in the PC and the MtC. However, no previous physiological study has investigated the activity in these two lobes simultaneously. In the present study, the activity evoked by electrical stimulation of the olfactory nerves, the superior and inferior tentacle nerves, was recorded optically from the whole cerebral ganglion of the terrestrial slug, Incilaria fruhstorferi. The results indicated that the evoked activity in the PC and the MtC showed two specific spatio-temporal patterns. First, when either set of nerves was stimulated, the activity of the medial neuropilar region of the MtC (the mMtC) always preceded the activity in the PC. Second, stimulation of the superior tentacle nerves activated the medial and lateral halves of the mMtC almost evenly, whereas stimulation of the inferior tentacle nerves activated the lateral half of the mMtC more strongly than the medial half. These results suggest that the activated region of the mMtC plays an important role in olfactory processing, especially with respect to the functional differences between the superior and inferior tentacles.

Tritocerebral tract input to the insect mushroom bodies

Insect mushroom bodies, best known for their role in olfactory processing, also receive sensory input from other modalities. In crickets and grasshoppers, a tritocerebral tract containing afferents from palp mechanosensory and gustatory centers innervates the accessory calyx. The accessory calyx is uniquely composed of Class III Kenyon cells, and was shown by immunohistochemistry to be present sporadically across several insect orders. Neuronal tracers applied to the source of tritocerebral tract axons in several species of insects demonstrated that tritocerebral tract innervation of the mushroom bodies targeted the accessory calyx when present, the primary calyces when an accessory calyx was not present, or both. These results suggest that tritocerebral tract input to the mushroom bodies is likely ubiquitous, reflecting the importance of gustation for insect behavior. The scattered phylogenetic distribution of Class III Kenyon cells is also proposed to represent an example of generative homology, in which the developmental program for forming a structure is retained in all members of a lineage, but the program is not “run” in all branches.

Role of GABAergic Inhibition in Shaping Odor-Evoked Spatiotemporal Patterns in theDrosophilaAntennal Lobe

Drosophila olfactory receptor neurons project to the antennal lobe, the insect analog of the mammalian olfactory bulb. GABAergic synaptic inhibition is thought to play a critical role in olfactory processing in the antennal lobe and olfactory bulb. However, the properties of GABAergic neurons and the cellular effects of GABA have not been described in Drosophila, an important model organism for olfaction research. We have used whole-cell patch-clamp recording, pharmacology, immunohistochemistry, and genetic markers to investigate how GABAergic inhibition affects olfactory processing in the Drosophila antennal lobe. We show that many axonless local neurons (LNs) in the adult antennal lobe are GABAergic. GABA hyperpolarizes antennal lobe projection neurons (PNs) via two distinct conductances, blocked by a GABAA- and GABAB-type antagonist, respectively. Whereas GABAA receptors shape PN odor responses during the early phase of odor responses, GABAB receptors mediate odor-evoked inhibition on longer time scales. The patterns of odor-evoked GABAB-mediated inhibition differ across glomeruli and across odors. Finally, we show that LNs display broad but diverse morphologies and odor preferences, suggesting a cellular basis for odor- and glomerulus-dependent patterns of inhibition. Together, these results are consistent with a model in which odors elicit stimulus-specific spatial patterns of GABA release, and as a result, GABAergic inhibition increases the degree of difference between the neural representations of different odors.