Odor Representations Research Articles

Odor representation and coding by the mitral/tufted cells in the olfactory bulb.

The olfactory bulb (OB) is the first relay station in the olfactory system and functions as a crucial hub. It can represent odor information precisely and accurately in an ever-changing environment. As the only output neurons in the OB, mitral/tufted cells encode information such as odor identity and concentration. Recently, the neural strategies and mechanisms underlying odor representation and encoding in the OB have been investigated extensively. Here we review the main progress on this topic. We first review the neurons and circuits involved in odor representation, including the different cell types in the OB and the neural circuits within and beyond the OB. We will then discuss how two different coding strategies-spatial coding and temporal coding-work in the rodent OB. Finally, we discuss potential future directions for this research topic. Overall, this review provides a comprehensive description of our current understanding of how odor information is represented and encoded by mitral/tufted cells in the OB.

Single-neuron representations of odours in the human brain

Olfaction is a fundamental sensory modality that guides animal and human behaviour1,2. However, the underlying neural processes of human olfaction are still poorly understood at the fundamental—that is, the single-neuron—level. Here we report recordings of single-neuron activity in the piriform cortex and medial temporal lobe in awake humans performing an odour rating and identification task. We identified odour-modulated neurons within the piriform cortex, amygdala, entorhinal cortex and hippocampus. In each of these regions, neuronal firing accurately encodes odour identity. Notably, repeated odour presentations reduce response firing rates, demonstrating central repetition suppression and habituation. Different medial temporal lobe regions have distinct roles in odour processing, with amygdala neurons encoding subjective odour valence, and hippocampal neurons predicting behavioural odour identification performance. Whereas piriform neurons preferably encode chemical odour identity, hippocampal activity reflects subjective odour perception. Critically, we identify that piriform cortex neurons reliably encode odour-related images, supporting a multimodal role of the human piriform cortex. We also observe marked cross-modal coding of both odours and images, especially in the amygdala and piriform cortex. Moreover, we identify neurons that respond to semantically coherent odour and image information, demonstrating conceptual coding schemes in olfaction. Our results bridge the long-standing gap between animal models and non-invasive human studies and advance our understanding of odour processing in the human brain by identifying neuronal odour-coding principles, regional functional differences and cross-modal integration.

Recent odor experience selectively modulates olfactory sensitivity across the glomerular output in the mouse olfactory bulb.

Although animals can reliably locate and recognize odorants embedded in complex environments, the neural circuits for accomplishing these tasks remain incompletely understood. Adaptation is likely to be important as it could allow neurons in a brain area to adjust to the broader sensory environment. Adaptive processes must be flexible enough to allow the brain to make dynamic adjustments, while maintaining sufficient stability so that organisms do not forget important olfactory associations. Processing within the mouse olfactory bulb is likely involved in generating adaptation, although there are conflicting models of how it transforms the glomerular output of the mouse olfactory bulb. Here we performed 2-photon Ca2+ imaging from mitral/tufted glomeruli in awake mice to determine the time course of recovery from adaptation, and whether it acts broadly or selectively across the glomerular population. Individual glomerular responses, as well as the overall population odor representation was similar across imaging sessions. However, odor-concentration pairings presented with interstimulus intervals upwards of 30-s evoked heterogeneous adaptation that was concentration-dependent. We demonstrate that this form of adaptation is unrelated to variations in respiration, and olfactory receptor neuron glomerular measurements indicate that it is unlikely to be inherited from the periphery. Our results indicate that the olfactory bulb output can reliably transmit stable odor representations, but recent odor experiences can selectively shape neural responsiveness for upwards of 30 seconds. We propose that neural circuits that allow for non-uniform adaptation across mitral/tufted glomerular could be important for making dynamic adjustments in complex odor environments.

Consistent social odor representation across 7 languages: the Social Odor Scale translation and validation.

The Social Odor Scale (SOS) is a 12-item questionnaire initially developed and validated in Italian and German to investigate self-reported awareness of social odors, which are odors emanating from the human body that convey diverse information and evoke various emotional responses. The scale includes a total score and 3 subscales representing social odors in the respective categories: romantic partner, familiar, and strangers. Here, we aimed to (i) replicate the validation of the Italian and German versions of the SOS, (ii) translate and validate the SOS into multiple additional languages (French, English, Dutch, Swedish, Chinese), and (iii) explore whether the factor structure of each translated version aligns with the original versions. Confirmatory Factor Analysis (CFA) supported the scale's structure, yielding a good fit across all languages. Notable differences in SOS mean scores were observed among the different languages: Swedish participants exhibited lower social odor awareness compared to the other groups, whereas Chinese participants reported higher social odor awareness compared to Dutch and Swedish participants. Furthermore, SOS scores correlated with respondents' geographical location, with higher (i.e. northern) latitudes linked to lower social odor awareness. These results corroborate the SOS as a valid and reliable instrument, especially for the SOS total score and the Familiar and Partner factors, emphasizing the influence of individual and geographic factors on social odor awareness.

The primacy model and the structure of olfactory space.

Understanding sensory processing relies on the establishment of a consistent relationship between the stimulus space, its neural representation, and perceptual quality. In olfaction, the difficulty in establishing these links lies partly in the complexity of the underlying odor input space and perceptual responses. Based on the recently proposed primacy model for concentration invariant odor identity representation and a few assumptions, we have developed a theoretical framework for mapping the odor input space to the response properties of olfactory receptors. We analyze a geometrical structure containing odor representations in a multidimensional space of receptor affinities and describe its low-dimensional implementation, the primacy hull. We propose the implications of the primacy hull for the structure of feedforward connectivity in early olfactory networks. We test the predictions of our theory by comparing the existing receptor-ligand affinity and connectivity data obtained in the fruit fly olfactory system. We find that the Kenyon cells of the insect mushroom body integrate inputs from the high-affinity (primacy) sets of olfactory receptors in agreement with the primacy theory.

Non-spatial hippocampal behavioral timescale synaptic plasticity during working memory is gated by entorhinal inputs.

Behavioral timescale synaptic plasticity (BTSP) is a form of synaptic potentiation where the occurrence of a single large plateau potential in CA1 hippocampal neurons leads to the formation of reliable place fields during spatial learning tasks. We asked whether BTSP could also be a plasticity mechanism for generation of non-spatial responses in the hippocampus and what roles the medial and lateral entorhinal cortex (MEC and LEC) play in driving non-spatial BTSP. By performing simultaneous calcium imaging of dorsal CA1 neurons and chemogenetic inhibition of LEC or MEC while mice performed an olfactory working memory task, we discovered BTSP-like events which formed stable odor-specific fields. Critically, the success rate of calcium events generating a significant odor-field increased with event amplitude, and large events exhibited asymmetrical formation with the newly formed odor-fields preceding the timepoint of their induction event. We found that MEC and LEC play distinct roles in modulating BTSP: MEC inhibition reduced the frequency of large calcium events, while LEC inhibition reduced the success rate of odor-field generation. Using two-photon calcium imaging of LEC and MEC temporammonic axons projecting to CA1, we found that LEC projections to CA1 were strongly odor selective even early in task learning, while MEC projection odor-selectivity increased with task learning but remained weaker than LEC. Finally, we found that LEC and MEC inhibition both slowed representational drift of odor representations in CA1 across 48 hours. Altogether, odor-specific information from LEC and strong odor-timed activity from MEC are crucial for driving BTSP in CA1, which is a synaptic plasticity mechanism for generation of both spatial and non-spatial responses in the hippocampus that may play a role in explaining representational drift and one-shot learning of non-spatial information.

Dense and persistent odor representations in the olfactory bulb of awake mice.

Recording and analysis of neural activity is often biased toward detecting sparse subsets of highly active neurons, masking important signals carried in low magnitude and variable responses. To investigate the contribution of seemingly noisy activity to odor encoding, we used mesoscale calcium imaging from mice of both sexes to record odor responses from the dorsal surface of bilateral olfactory bulbs (OBs). The outer layer of the mouse OB is comprised of dendrites organized into discrete "glomeruli", which are defined by odor receptor-specific sensory neuron input. We extracted activity from a large population of glomeruli and used logistic regression to classify odors from individual trials with high accuracy. We then used add-in and drop-out analyses to determine subsets of glomeruli necessary and sufficient for odor classification. Classifiers successfully predicted odor identity even after excluding sparse, highly active glomeruli, indicating that odor information is redundantly represented across a large population of glomeruli. Additionally, we found that Random Forest feature selection informed by Gini Inequality (RFGI) reliably ranked glomeruli by their contribution to overall odor classification. RFGI provided a measure of "feature importance" for each glomerulus that correlated with intuitive features like response magnitude. Finally, in agreement with previous work, we found that odor information persists in glomerular activity after odor offset. Together, our findings support a model of olfactory bulb odor coding where sparse activity is sufficient for odor identification, but information is widely, redundantly available across a large population of glomeruli, with each glomerulus representing information about more than one odor.Significance statement This study leverages meso-scale imaging and machine learning to investigate how odor information is first represented in the brain. Typically, recordings of neuronal activity focus on active individual cells, potentially overlooking broader variations in neuronal responses across populations. Our results demonstrate that a considerable amount of olfactory information is redundantly distributed across a large proportion of olfactory bulb glomeruli. Even after excluding a majority of glomeruli, odor identification remained possible. These findings indicate that, although a few glomeruli are sufficient for odor recognition, an abundance of additional information is represented across a broad population. Understanding how the brain manages redundant olfactory information will shed light on its adaptive mechanisms for navigating diverse real-world circumstances and responding to fluctuating internal states.

Adult-neurogenesis allows for representational stability and flexibility in early olfactory system.

In the early olfactory system, adult-neurogenesis, a process of neuronal replacement results in the continuous reorganization of synaptic connections and network architecture throughout the animal's life. This poses a critical challenge: How does the olfactory system maintain stable representations of odors and therefore allow for stable sensory perceptions amidst this ongoing circuit instability? Utilizing a detailed spiking network model of early olfactory circuits, we uncovered dual roles for adult-neurogenesis: one that both supports representational stability to faithfully encode odor information and also one that facilitates plasticity to allow for learning and adaptation. In the main olfactory bulb, adult-neurogenesis affects neural codes in individual mitral and tufted cells but preserves odor representations at the neuronal population level. By contrast, in the olfactory piriform cortex, both individual cell responses and overall population dynamics undergo progressive changes due to adult-neurogenesis. This leads to representational drift, a gradual alteration in sensory perception. Both processes are dynamic and depend on experience such that repeated exposure to specific odors reduces the drift due to adult-neurogenesis; thus, when the odor environment is stable over the course of adult-neurogenesis, it is neurogenesis that actually allows the representations to remain stable in piriform cortex; when those olfactory environments change, adult-neurogenesis allows the cortical representations to track environmental change. Whereas perceptual stability and plasticity due to learning are often thought of as two distinct, often contradictory processing in neuronal coding, we find that adult-neurogenesis serves as a shared mechanism for both. In this regard, the quixotic presence of adult-neurogenesis in the mammalian olfactory bulb that has been the focus of considerable debate in chemosensory neuroscience may be the mechanistic underpinning behind an array of complex computations.

Unveiling the odor representation in the inner brain of Drosophila through compressed sensing

The olfaction begins when odorant molecules stimulate a finite number of receptor types in the olfactory neurons. According to the recent connectomics data of the olfactory system, the olfactory neural circuit is multilayered and there is an overall dimensionality reduction from the input to output layer. Compressed sensing (CS) is an algorithm that enables the recovery of high-dimensional signals from the data compressed in a lower dimension when the representation of such signals is sufficiently sparse. By analyzing the recent connectomics data, we find that the organization of the olfactory system effectively satisfies the necessary conditions for CS to work. The neural activity profile of projection neurons (PNs) can be faithfully recovered from a low-dimensional response profile of mushroom body output neurons (MBONs), which can be reconstructed using the electrophysiological recordings of a wide range of odorants. By leveraging the residuals calculated between the measured and the predicted MBON responses, we visualize the perceptual odor space using the residual spectrum and discuss the differentiability of an odor from others. Our study highlights the sparse coding of odor to the receptor space as an essential component for odor identifiability, clarifying the principles underlying the concentration-dependent odor percept. Further, simultaneous exposure of the olfactory system to many different odorants saturates the neural activity profile of PNs, significantly degrading the capacity of signal recovery and resulting in a perceptual state analogous to “olfactory white.” Our study applying CS to the real connectomics data encompassing the neural circuitry from PNs to MBONs via Kenyon cells provides quantitative insights into the neural signal processing and odor representation in the inner brain of . Published by the American Physical Society 2024

Ring-shaped odor coding in the antennal lobe of migratory locusts

The representation of odors in the locust antennal lobe with its >2,000 glomeruli has long remained a perplexing puzzle. We employed the CRISPR-Cas9 system to generate transgenic locusts expressing the genetically encoded calcium indicator GCaMP in olfactory sensory neurons. Using two-photon functional imaging, we mapped the spatial activation patterns representing a wide range of ecologically relevant odors across all six developmental stages. Our findings reveal a functionally ring-shaped organization of the antennal lobe composed of specific glomerular clusters. This configuration establishes an odor-specific chemotopic representation by encoding different chemical classes and ecologically distinct odors in the form of glomerular rings. The ring-shaped glomerular arrangement, which we confirm by selective targeting of OR70a-expressing sensory neurons, occurs throughout development, and the odor-coding pattern within the glomerular population is consistent across developmental stages. Mechanistically, this unconventional spatial olfactory code reflects the locust-specific and multiplexed glomerular innervation pattern of the antennal lobe.

Pheromone representation in the ant antennal lobe changes with age

While the neural basis of age-related decline has been extensively studied,1,2,3 less is known about changes in neural function during the pre-senescent stages of adulthood. Adult neural plasticity is likely a key factor in social insect age polyethism, where individuals perform different tasks as they age and divide labor in an age-dependent manner.4,5,6,7,8,9 Primarily, workers transition from nursing to foraging tasks,5,10 become more aggressive, and more readily display alarm behavior11,12,13,14,15,16 as they get older. While it is unknown how these behavioral dynamics are neurally regulated, they could partially be generated by altered salience of behaviorally relevant stimuli.4,6,7 Here, we investigated how odor coding in the antennal lobe (AL) changes with age in the context of alarm pheromone communication in the clonal raider ant (Ooceraea biroi).17 Similar to other social insects,11,12,16 older ants responded more rapidly to alarm pheromones, the chemical signals for danger. Using whole-AL calcium imaging,18 we then mapped odor representations for five general odorants and two alarm pheromones in young and old ants. Alarm pheromones were represented sparsely at all ages. However, alarm pheromone responses within individual glomeruli changed with age, either increasing or decreasing. Only two glomeruli became sensitized to alarm pheromones with age, while at the same time becoming desensitized to general odorants. Our results suggest that the heightened response to alarm pheromones in older ants occurs via increased sensitivity in these two core glomeruli, illustrating the importance of sensory modulation in social insect division of labor and age-associated behavioral plasticity.

Cholecystokinin-expressing superficial tufted cells modulate odour representation in the olfactory bulb and olfactory behaviours.

In mammals, odour information within the olfactory bulb (OB) is processed by complex neural circuits before being ultimately represented in the action potential activity of mitral/tufted cells (M/Ts). Cholecystokinin-expressing (CCK+) superficial tufted cells (sTCs) are a subset of tufted cells that potentially contribute to olfactory processing in the OB by orchestrating M/T activity. However, the exact role of CCK+ sTCs in modulating odour processing and olfactory function in vivo is largely unknown. Here, we demonstrate that manipulating CCK+ sTCs can generate perception and induce place avoidance. Optogenetic activation/inactivation of CCK+ sTCs exerted strong but differing effects on spontaneous and odour-evoked M/T firing. Furthermore, inactivation of CCK+ sTCs disrupted M/T odour encoding and impaired olfactory detection and odour discrimination. These results establish the role of CCK+ sTCs in odour representation and olfactory behaviours. KEY POINTS: Mice could perceive the activity of CCK+ sTCs and show place avoidance to CCK+ sTC inactivation. Optical activation of CCK+ sTCs increased the percentage of cells with odour response but reduced the odour-evoked response in M/Ts in awake mice. Optical inactivation of CCK+ sTCs greatly decreased spontaneous firing and odour-evoked response in M/Ts. Inactivation of CCK+ sTCs impairs the odour decoding performance of M/Ts and disrupts odour detection and discrimination behaviours in mice. These results indicate that CCK+ sTCs participate in modulating the odour representation and maintaining normal olfactory-related behaviours.

Analysis of Legal Protection of Non-traditional Trademarks from a Global PerspectiveTaking Scent Trademarks as an Example

With the constant advancement of the economy and technology, especially the development of high-tech technology, the traditional trademark can no longer meet the needs of social and enterprise development. From traditional visual trademarks to three-dimensional trademarks, sound marks, and even scent marks, non-traditional trademarks are playing an increasingly important role in transactions. Therefore, the legal protection of non-traditional trademarks is particularly important. By collecting and reading the relevant references and materials, this paper adopts the methods of literature analysis, comparison method, and case analysis to analyze of Legal protection of non-traditional trademarks from a global perspectivetaking scent trademarks as an example. The paper points out the current situation in the protection of non-traditional trademarks, taking scent marks as an example, and puts forward some feasible suggestions, such as: Reference the "staff " representation method, Promoting scientific and technological progress, and Scent branding prompts in the product. It can be concluded that the registration and protection systems of scent trademarks in various countries are not perfect, they can be shown in the strict requirements for registrable trademarks regarding the representation of odor, the distinctiveness of trademark registration, and the functional of marks. Countries should pay more attention to them. The study suggests that it is necessary to improve the scent trademark legislation, which is conducive to better protection of intellectual property rights, encourage innovation of enterprises, and safeguard the interests of enterprises.

Semantic context-dependent neural representations of odors in the human piriform cortex revealed by 7T MRI.

Olfactory perception depends not only on olfactory inputs but also on semantic context. Although multi-voxel activity patterns of the piriform cortex, a part of the primary olfactory cortex, have been shown to represent odor perception, it remains unclear whether semantic contexts modulate odor representation in this region. Here, we investigated whether multi-voxel activity patterns in the piriform cortex change when semantic context modulates odor perception and, if so, whether the modulated areas communicate with brain regions involved in semantic and memory processing beyond the piriform cortex. We also explored regional differences within the piriform cortex, which are influenced by olfactory input and semantic context. We used 2 × 2 combinations of word labels and odorants that were perceived as congruent and measured piriform activity with a 1-mm isotropic resolution using 7T MRI. We found that identical odorants labeled with different words were perceived differently. This labeling effect was observed in multi-voxel activity patterns in the piriform cortex, as the searchlight decoding analysis distinguished identical odors with different labels for half of the examined stimulus pairs. Significant functional connectivity was observed between parts of the piriform cortex that were modulated by labels and regions associated with semantic and memory processing. While the piriform multi-voxel patterns evoked by different olfactory inputs were also distinguishable, the decoding accuracy was significant for only one stimulus pair, preventing definitive conclusions regarding the locational differences between areas influenced by word labels and olfactory inputs. These results suggest that multi-voxel patterns of piriform activity can be modulated by semantic context, possibly due to communication between the piriform cortex and the semantic and memory regions.

Vasopressin differentially modulates the excitability of rat olfactory bulb neuron subtypes.

Vasopressin (VP) plays a crucial role in social memory even at the level of the olfactory bulb (OB), where OB VP cells are activated during social interactions. However, it remains unclear how VP modulates olfactory processing to enable enhanced discrimination of very similar odors, e.g., rat body odors. Thus far, it has been shown that VP reduces firing rates in mitral cells (MCs) during odor presentation in vivo and decreases the amplitudes of olfactory nerve-evoked excitatory postsynaptic potentials (ON-evoked EPSPs) in external tufted cells in vitro. We performed whole-cell patch-clamp recordings and population Ca2+ imaging on acute rat OB slices. We recorded ON-evoked EPSPs as well as spontaneous inhibitory postsynaptic currents (IPSCs) from two types of projection neurons: middle tufted cells (mTCs) and MCs. VP bath application reduced the amplitudes of ON-evoked EPSPs and the frequencies of spontaneous IPSCs in mTCs but did not change those in MCs. Therefore, we analyzed ON-evoked EPSPs in inhibitory interneurons, i.e., periglomerular cells (PGCs) and granule cells (GCs), to search for the origin of increased inhibition in mTCs. However, VP did not increase the amplitudes of evoked EPSPs in either type of interneurons. We next performed two-photon population Ca2+ imaging in the glomerular layer and the superficial GC layer of responses to stronger ON stimulation than during patch-clamp experiments that should evoke action potentials in the measured cells. We observed that VP application increased ON-evoked Ca2+ influx in juxtaglomerular cells and GC somata. Thus, our findings indicate inhibition by VP on projection neurons via strong ON input-mediated inhibitory interneuron activity. This neural modulation could improve representation of odors, hence, better discriminability of similar odors, e.g., conspecific body odors.

Non-canonical odor representation and learning in Dipteran brains

For decades, the vinegar fly Drosophila melanogaster has provided a window into the structure and function of the insect olfactory system, revealing a remarkable organizational correspondence between insects and vertebrates. In both clades, olfactory sensory neurons expressing the same class of sensory receptor proteins exclusively target a dedicated neuropil known as a glomerulus. Here, we review recent evidence from Drosophila and other Dipteran taxa that challenges this canonical view, showing widespread co-expression of olfactory sensory proteins within the same neurons. We discuss the potential consequences of co-expression for early sensory processing in the insect olfactory system. As well, we situate these findings within the broader framework of olfactory learning, highlighting recent findings that suggest a wider importance of the antennal lobe than has been previously appreciated.

Odor representations from the two nostrils are temporally segregated in human piriform cortex

The human olfactory system has two discrete channels of sensory input, arising from olfactory epithelia housed in the left and right nostrils. Here, we asked whether the primary olfactory cortex (piriform cortex [PC]) encodes odor information arising from the two nostrils as integrated or distinct stimuli. We recorded intracranial electroencephalogram (iEEG) signals directly from PC while human subjects participated in an odor identification task where odors were delivered to the left, right, or both nostrils. We analyzed the time course of odor identity coding using machine-learning approaches and found that uni-nostril odor inputs to the ipsilateral nostril are encoded ∼480-ms faster than odor inputs to the contralateral nostril on average. During naturalistic bi-nostril odor sampling, odor information emerged in two temporally segregated epochs, with the first epoch corresponding to the ipsilateral and the second epoch corresponding to the contralateral odor representations. These findings reveal that PC maintains distinct representations of odor input from each nostril through temporal segregation, highlighting an olfactory coding scheme at the cortical level that can parse odor information across nostrils within the course of a single inhalation.

Functional odor map heterogeneity is based on multifaceted glomerular connectivity in larval Xenopus olfactory bulb

Glomeruli are the functional units of the vertebrate olfactory bulb (OB) connecting olfactory receptor neuron (ORN) axons and mitral/tufted cell (MTC) dendrites. In amphibians, these two circuit elements regularly branch and innervate multiple, spatially distinct glomeruli. Using functional multiphoton-microscopy and single-cell tracing, we investigate the impact of this wiring on glomerular module organization and odor representations on multiple levels of the Xenopus laevis OB network. The glomerular odor map to amino acid odorants is neither stereotypic between animals nor chemotopically organized. Among the morphologically heterogeneous group of uni- and multi-glomerular MTCs, MTCs can selectively innervate glomeruli formed by axonal branches of individual ORNs. We conclude that odor map heterogeneity is caused by the coexistence of different intermingled glomerular modules. This demonstrates that organization of the amphibian main olfactory system is not strictly based on uni-glomerular connectivity.

Gain modulation and odor concentration invariance in early olfactory networks.

The broad receptive field of the olfactory receptors constitutes the basis of a combinatorial code that allows animals to detect and discriminate many more odorants than the actual number of receptor types that they express. One drawback is that high odor concentrations recruit lower affinity receptors which can lead to the perception of qualitatively different odors. Here we addressed the contribution that signal-processing in the antennal lobe makes to reduce concentration dependence in odor representation. By means of calcium imaging and pharmacological approach we describe the contribution that GABA receptors play in terms of the amplitude and temporal profiles of the signals that convey odor information from the antennal lobes to higher brain centers. We found that GABA reduces the amplitude of odor elicited signals and the number of glomeruli that are recruited in an odor-concentration-dependent manner. Blocking GABA receptors decreases the correlation among glomerular activity patterns elicited by different concentrations of the same odor. In addition, we built a realistic mathematical model of the antennal lobe that was used to test the viability of the proposed mechanisms and to evaluate the processing properties of the AL network under conditions that cannot be achieved in physiology experiments. Interestingly, even though based on a rather simple topology and cell interactions solely mediated by GABAergic lateral inhibitions, the AL model reproduced key features of the AL response upon different odor concentrations and provides plausible solutions for concentration invariant recognition of odors by artificial sensors.

Social odor discrimination and its enhancement by associative learning in the hippocampal CA2 region

Although the hippocampus is crucial for social memory, how social sensory information is combined with contextual information to form episodic social memories remains unknown. Here, we investigated the mechanisms for social sensory information processing using two-photon calcium imaging from hippocampal CA2 pyramidal neurons (PNs)-which are crucial for social memory-in awake head-fixed mice exposed to social and non-social odors. We found that CA2 PNs represent social odors of individual conspecifics and that these representations are refined during associative social odor-reward learning to enhance the discrimination of rewarded compared with unrewarded odors. Moreover, the structure of the CA2 PN population activity enables CA2 to generalize along categories of rewarded versus unrewarded and social versus non-social odor stimuli. Finally, we found that CA2 is important for learning social but not non-social odor-reward associations. These properties of CA2 odor representations provide a likely substrate for the encoding of episodic social memory.