Septal Organ Research Articles

Photoneuroendocrine, circadian and seasonal systems: from photoneuroendocrinology to circadian biology and medicine.

This contribution highlights the scientific development of two intertwined disciplines, photoneuroendocrinology and circadian biology. Photoneuroendocrinology has focused on nonvisual photoreceptors that translate light stimuli into neuroendocrine signals and serve rhythm entrainment. Nonvisual photoreceptors first described in the pineal complex and brain of nonmammalian species are luminance detectors. In the pineal, they control the formation of melatonin, the highly conserved hormone of darkness which is synthesized night by night. Pinealocytes endowed with both photoreceptive and neuroendocrine capacities function as "photoneuroendocrine cells." In adult mammals, nonvisual photoreceptors controlling pineal melatonin biosynthesis and pupillary reflexes are absent from the pineal and brain and occur only in the inner layer of the retina. Encephalic photoreceptors regulate seasonal rhythms, such as the reproductive cycle. They are concentrated in circumventricular organs, the lateral septal organ and the paraventricular organ, and represent cerebrospinal fluid contacting neurons. Nonvisual photoreceptors employ different photopigments such as melanopsin, pinopsin, parapinopsin, neuropsin, and vertebrate ancient opsin. After identification of clock genes and molecular clockwork, circadian biology became cutting-edge research with a focus on rhythm generation. Molecular clockworks tick in every nucleated cell and, as shown in mammals, they drive the expression of more than 3000 genes and are of overall importance for regulation of cell proliferation and metabolism. The mammalian circadian system is hierarchically organized; the central rhythm generator is located in the suprachiasmatic nuclei which entrain peripheral circadian oscillators via multiple neuronal and neuroendocrine pathways. Disrupted molecular clockworks may cause various diseases, and investigations of this interplay will establish a new discipline: circadian medicine.

Interpretive review: Semiochemicals in domestic pigs and dogs.

This interpretive review includes discussion of the available scientific literature with interpretations by the authors. The broad field of semiochemicals can be confusing to scientists and consumers. This review attempts to summarize the known scientific studies for pig and dog semiochemicals while at the same time attempting to refine our use of terminology. The specific objectives of this interpretive review are to summarize and interpret much of the key scientific literature (but not the lay literature) on semiochemicals in pigs and dogs to include (1) definitions of semiochemicals and related molecules including pheromones, (2) to briefly summarize olfactory organs, and (3) and to examine the scientific literature for semiochemical mechanisms and applications in dogs and pigs (two domesticated species with known olfactory acuity). Dogs and pigs have olfactory features that are similar in that they both lack certain olfactory organs (Grueneberg ganglion and Septal Organ) and they have a small vomeronasal organ (VNO) without some major receptors that are found in other species. The primary olfactory organs for both pigs and dogs are the main olfactory epithelium and perhaps the trigeminal nerve. Several examples of pheromones activating the brain via the MOE or Trigeminal nerve rather than the VNO challenge the concept that the VNO is the site of pheromone sensing. We believe it is not appropriate to label something a pheromone when evidence is not available to show that it is a pheromone. We offer definitions for the terms semiochemicals, pheromones, interomones and others and then determine if the evidence is sufficient to call certain semiochemicals a pheromone. Here we review mixed, largely negative, scientific reports of the efficacy of some products labeled as "pheromones" that are more appropriately called semiochemicals. Interomones can have a more powerful effect on dog behavior and physiology than semiochemicals marketed as pheromones. Because marketing of semiochemicals is far ahead of the science, bringing some logic and uniformity to the field will benefit animals and hopefully cause less consumer confusion. Semiochemicals have the potential to offer powerful solutions to behavioral problems using more naturally occurring molecules.

Bcl11b is required for proper odorant receptor expression in the mouse septal organ.

Individual olfactory sensory neurons (OSNs) in the mouse main olfactory epithelium express a single odorant receptor (OR) gene from the repertoire of either class I or class II ORs. The transcription factor Bcl11b determines the OR class to be expressed in OSNs. The septal organ (SO), a small neuroepithelium located at the ventral base of the nasal septum, is considered as an olfactory subsystem because it expresses a specific subset of ORs. However, the mechanisms underlying the generation and differentiation of SO-OSN remain unknown. In the present study, we show that the generation and differentiation of SO-OSN employ the same genetic pathway as in the OSN lineage, which is initiated by the neuronal fate determinant factor Ascl1. Additionally, the key role of Bcl11b in the SO is demonstrated by the abnormal phenotypes of Bcl11b-deficient mice: significant reduction in the expression of OR genes and in the number of mature SO-OSNs. Although SO-OSNs are specified to express a subset of class II OR genes in wild-type mice, the Bcl11b deletion led to the expression of class I OR genes, while the expression of class II OR genes was significantly decreased, with one exception of Olfr15. These results indicate that Bcl11b is necessary for proper OR expression in SO-OSNs.

Unique nasal septal island in dromedary camels may play a role in pain perception: microscopic studies.

The septal organs are islands or patches of sensory epithelium, located in the ventral parts of the nasal septum and innervated by the olfactory nerve. The septal island in dromedaries (Camelus dromedarius) was unusually located in the rostro-dorsal part of the nasal septum, where the ethmoidal branch of the trigeminal nerve provides innervation to the island mucosa. Therefore, the objectives of this study were to reveal the microscopic and ultrastructure of this island and to explain the probable functions. Twelve septal islands from 12 healthy male camels were used. Unlike the olfactory epithelium, which has a pseudostratified structure, the island neuroepithelium had a true neural lamination. Furthermore, in electron micrographs, the receptor, bipolar, and basal cells were connected with an orderly, organized network of cell-cell communication, which had some spine synapses. This network substituted the absence of supporting cells, maintained the shape of the tissue, and held the cells together. Moreover, the receptor cells were not similar to any of the different types of olfactory sensory neurons. Instead, they possessed the apical domain that might be specialized for the detection of chemical stimuli. Interestingly, a resident population of immune cells, namely mast cells and macrophages, was observed. The probable functions were discussed based on the cellular context and architecture. The nasal septal island in dromedaries may have a role in pain perception. The receptor cells most probably work as nociceptive cells that interact with the resident immune cells to coordinate pain signaling with immune response.

Coding of pheromones by vomeronasal receptors.

Communication between individuals is critical for species survival, reproduction, and expansion. Most terrestrial species, with the exception of humans who predominantly use vision and phonation to create their social network, rely on the detection and decoding of olfactory signals, which are widely known as pheromones. These chemosensory cues originate from bodily fluids, causing attractive or avoidance behaviors in subjects of the same species. Intraspecific pheromone signaling is then crucial to identify sex, social ranking, individuality, and health status, thus establishing hierarchies and finalizing the most efficient reproductive strategies. Indeed, all these features require fine tuning of the olfactory systems to detect molecules containing this information. To cope with this complexity of signals, tetrapods have developed dedicated olfactory subsystems that refer to distinct peripheral sensory detectors, called the main olfactory and the vomeronasal organ, and two minor structures, namely the septal organ of Masera and the Grueneberg ganglion. Among these, the vomeronasal organ plays the most remarkable role in pheromone coding by mediating several behavioral outcomes that are critical for species conservation and amplification. In rodents, this organ is organized into two segregated neuronal subsets that express different receptor families. To some extent, this dichotomic organization is preserved in higher projection areas of the central nervous system, suggesting, at first glance, distinct functions for these two neuronal pathways. Here, I will specifically focus on this issue and discuss the role of vomeronasal receptors in mediating important innate behavioral effects through the recognition of pheromones and other biological chemosignals.

The functional relevance of olfactory marker protein in the vertebrate olfactory system: a never-ending story.

Olfactory marker protein (OMP) was first described as a protein expressed in olfactory receptor neurons (ORNs) in the nasal cavity. In particular, OMP, a small cytoplasmic protein, marks mature ORNs and is also expressed in the neurons of other nasal chemosensory systems: the vomeronasal organ, the septal organ of Masera, and the Grueneberg ganglion. While its expression pattern was more easily established, OMP's function remained relatively vague. To date, most of the work to understand OMP's role has been done using mice lacking OMP. This mostly phenomenological work has shown that OMP is involved in sharpening the odorant response profile and in quickening odorant response kinetics of ORNs and that it contributes to targeting of ORN axons to the olfactory bulb to refine the glomerular response map. Increasing evidence shows that OMP acts at the early stages of olfactory transduction by modulating the kinetics of cAMP, the second messenger of olfactory transduction. However, how this occurs at a mechanistic level is not understood, and it might also not be the only mechanism underlying all the changes observed in mice lacking OMP. Recently, OMP has been detected outside the nose, including the brain and other organs. Although no obvious logic has become apparent regarding the underlying commonality between nasal and extranasal expression of OMP, a broader approach to diverse cellular systems might help unravel OMP's functions and mechanisms of action inside and outside the nose.

Chapter 4 - Anatomy of the olfactory mucosa

The classic notion that humans are microsmatic animals was born from comparative anatomy studies showing the reduction in the size of both the olfactory bulbs and the limbic brain relative to the whole brain. However, the human olfactory system contains a number of neurons comparable to that of most other mammals, and humans have exquisite olfactory abilities. Major advances in molecular and genetic research have resulted in the identification of extremely large gene families that express receptors for sensing odors. Such advances have led to a renaissance of studies focused on both human and nonhuman aspects of olfactory physiology and function. Evidence that olfactory dysfunction is among the earliest signs of a number of neurodegenerative and neuropsychiatric disorders has led to considerable interest in the use of olfactory epithelial biopsies for potentially identifying such disorders. Moreover, the unique features of the olfactory ensheathing cells have made the olfactory mucosa a promising and unexpected source of cells for treating spinal cord injuries and other neural injuries in which cell guidance is critical. The olfactory system of humans and other primates differs in many ways from that of other species. In this chapter we provide an overview of the anatomy of not only the human olfactory mucosa but of mucosae from a range of mammals from which more detailed information is available. Basic information regarding the general organization of the olfactory mucosa, including its receptor cells and the large number of other cell types critical for their maintenance and function, is provided. Cross-species comparisons are made when appropriate. The polemic issue of the human vomeronasal organ in both the adult and fetus is discussed, along with recent findings regarding olfactory subsystems within the nose of a number of mammals (e.g., the septal organ and Grüneberg ganglion).

Structural organization of the olfactory organ in an amphihaline migratory fish Hilsa, Tenualosa ilisha.

The histological as well as ultramicroscopic structures of olfactory system of an amphihaline migratory fish hilsa Tenualosa ilisha, were studied. The sexually matured riverine fish were collected from a common breeding habitat-the Hooghly, a tributary of river Ganga, West Bengal, India. This study revealed that the riverine hilsa has larger olfactory bulb compared to marine hilsa with the olfactory lobes well exposed through nostrils. The olfactory lamellae (OL) are 40-45 in number and posses three distinct layers of sensory cells across each lamellae, namely, outer receptor cells (RC), middle sensory cells, and inner basal cells (BC). Besides the above arrangement, the sensory part of olfactory epithelium (OE) also bears rich microvillous cells exposed to the surface of the OE. The sensory and non-sensory surfaces on OL are distinguishable, with clear dendritic cells on sensory epithelium and solitary chemosensory cells on non sensory OE. Abundance of both types of cells in the OE is an indication of its chemoattraction ability towards molecules of amino acid origin. The feature of having abundant, dense, and large dendritic knobs on the surface of OE describes resemblance to the typical morphology of the chemosensory septal organs neuron. The expression of four G protein subunits, like Gαs/olf, Gαq, Gαo, and Gαi-3 in OE indicate that its olfaction is a functional attributes of two olfactory systems, namely main olfactory system and Vomaronasal Olfactory System. Expression of ACIII and PLCβ2 in OE further confirms two signaling pathways involved in odorant reception in hilsa. RESEARCH HIGHLIGHTS: The olfactory bulb in the amphihaline migratory fish hilsa is big in size, with 40-45 lamellae. Its sensory areas showed multilayered cellular features with prominent sensory as well as microvillous cells, whereas non-sensory area possesses solitary chemosensory cells. The expression of four G protein subunits, Gαs/olf, Gαq, Gαo, and Gαi-3 in olfactory epithelium indicates that its olfaction is a functional attributes of two olfactory systems, namely main olfactory system and vomaronasal olfactory system.

The Cytoarchitecture of the Telencephalon of Betta Splendens Regan 1910 (Perciformes: Anabantoidei) with a Stereological Approach to the Supracommissural and Postcommissural Nuclei.

Teleostean fish brains are useful models to study cellular and functional specializations along the phylogenesis. The Betta splendens Regan 1910 (Siamese fighting fish; Perciformes:Anabantoidei) is known for its aggressive display, courtship behavior, nest building, and offspring care. Here, we present novel and detailed data about the cytoarchitecture of the olfactory bulb and the telencephalic hemispheres of this fish. The hematoxylin-eosin and Nissl techniques served to identify brain nuclei (n = 19 males and n = 21 females) and for the stereological evaluation of the numerical density of cells and the proportion of neurons and glial cells in the ventral telencephalon supracommissural (Vs) and postcommissural (Vp) nuclei of adult males and females. These nuclei are putative homologs of the sexually dimorphic medial amygdala in mammals. The olfactory bulb of Betta splendens consists of 5 concentrically arranged layers plus ganglion cells of the terminal nerves. The dorsal telencephalon consists of 16 different cell groups. The ventral telencephalon has 8 nuclei, plus the lateral septal organ and the nuclei of the preoptic area forming an anatomical continuum. The rostrocaudal extent of the Vs and Vp is not different between sexes. In both nuclei, the proportion of neurons to glial cells is approximately 2:1 and the density of neurons and glial cells is not different between sexes. These morphological findings can subserve future research on the brain function of the Betta splendens and the search for neural sex differences in other central areas of this same species, in other teleost species, or yet in other related vertebrate group. Anat Rec, 00:000-000, 2017. © 2017 Wiley Periodicals, Inc. Anat Rec, 301:88-110, 2018. © 2017 Wiley Periodicals, Inc.

Revisiting the Vomeronasal System From an Integrated Perspective.

"Olfactory subsystems" is a relatively new terminology to refer to the different regions of the nasal cavity featuring olfactory sensory neurons. In mice, the olfactory chemical cues are detected in four well delimited areas: the main olfactory epithelium, the septal organ, Grüneberg's ganglion, and the sensory epithelium of the vomeronasal organ. Nevertheless, such distribution is by no means exhibited by all mammals. In microsmatic mammals -humans included- the only existing olfactory subsystem is the main olfactory epithelium. This raises the question of whether the lack of certain olfactory structures in those species implies that they are unable to identify certain olfactory signals, or on the contrary, their main olfactory epithelium assumes such role. It would be interesting to determine, in the context of biomedical research, if the sense of smell in humans is fully or partially endowed with the wide range of functions assigned to the vomeronasal system in mice. If it is not, presumptive implications of the lack of such functions should be addressed in human health and well-being. Anat Rec, 299:1488-1491, 2016. © 2016 Wiley Periodicals, Inc.

Changes in brain peptides associated with reproduction and energy homeostasis in photosensitive and photorefractory migratory redheaded buntings

Present study examined the expression of brain peptides associated with the reproduction and energy homeostasis (GnRH/GnIH, NPY/VIP), and assessed their possible functional association in the photosensitive (non-breeding, pre-breeding), photostimulated (breeding) and photorefractory (post-breeding) migratory redheaded buntings (Emberiza bruniceps), using double-labeled immunohistochemistry. Particularly, we measured immunoreactive (-ir) cell numbers, per cent cell area and cell optical density (OD) in the preoptic area (GnRH-I), midbrain (GnRH-II), paraventricular nucleus (GnIH), dorsomedial hypothalamus, DMH and infundibular complex, INc (NPY and VIP), and lateral septal organ (VIP) of buntings kept under natural photoperiods at the wintering latitude (26°55′N). There was a significant seasonal difference in GnRH-I, not GnRH-II, with reduced -ir cells in the photosensitive and photorefractory buntings, and notably with increased cell OD between the refractory and non-breeding states with no increase in testis size. Also, increased cell OD of GnIH neurons in non-breeding state indicated its role in the maintenance of small testes during the post-refractory period. Overall, seasonal changes in GnRH-I and GnIH were found consistent with their suggested roles in reproductive regulation of absolute photorefractory birds. Further, there was a significant seasonal change in cell OD of NPY neurons in DMH, not the INc. In contrast, VIP immunoreactivity was seasonally altered, with a significantly higher VIP-ir cells in breeding than the pre-breeding state. Finally, close proximity between perikarya with fibres suggested functional interactions between the GnRH and GnIH, and NPY and VIP. Thus, seasonal plasticity of brain peptides is perhaps the part of neural regulation of seasonal reproduction and associated energy homeostasis in migratory songbirds.

Properties of an optogenetic model for olfactory stimulation.

In olfactory research it is difficult to deliver stimuli with defined intensity and duration to olfactory sensory neurons. Expression of channelrhodopsin 2 (ChR2) in olfactory sensory neurons provides a means to activate these neurons with light flashes. Appropriate mouse models are available. The present study explores the suitability of an established olfactory marker protein (OMP)/ChR2-yellow fluorescent protein (YFP) mouse model for ex vivo experimentation. Expression of ChR2 in sensory neurons of the main olfactory epithelium, the septal organ and vomeronasal organ is characterized. Expression pattern of ChR2 in olfactory receptor neurons and the properties of light responses indicate that light stimulation does not impact on signal transduction in the chemosensory cilia. Light-induced electro-olfactograms are characterized with light flashes of different intensities, durations and frequencies. The impact of light-induced afferent stimulation on the olfactory bulb is examined with respect to response amplitude, polarity and low-pass filtering. For the examination of sensory processing, it is helpful to deliver stimuli in precisely defined temporal and spatial patterns with accurate control of stimulus intensity. This is challenging in experiments with the mammalian olfactory system because airborne odorants have to be transported into the intricate sensory structures of the nose and must dissolve in mucus to be detected by sensory neurons. Defined and reproducible activity can be generated in olfactory sensory neurons that express the light-gated ion channel channelrhodopsin 2 (ChR2). The neurons can be stimulated by light flashes in a controlled fashion by this optogenetic approach. Here we examined the application of an olfactory marker protein (OMP)/ChR2-yellow fluorescent protein (YFP) model for ex vivo exploration of the olfactory epithelium and the olfactory bulb of the mouse. We studied the expression patterns of ChR2 in the main olfactory system, the vomeronasal system, and the septal organ, and we found that ChR2 is absent from the sensory cilia of olfactory sensory neurons. In the olfactory epithelium, we characterized light-induced electro-olfactograms with respect to peripheral encoding of stimulus intensity, stimulus duration and stimulus frequency. In acute slices of the olfactory bulb, we identified specific aspects of the ChR2-induced input signal, concerning its dynamic range, its low-pass filter property and its response to prolonged stimulation. Our study describes the performance of the OMP/ChR2-YFP model for ex vivo experimentation on the peripheral olfactory system and documents its versatility and its limitations for olfactory research.

Distribution of vasotocin- and vasoactive intestinal peptide-like immunoreactivity in the brain of blue tit (Cyanistes coeruleus).

Blue tits (Cyanistes coeruleus) are songbirds, used as model animals in numerous studies covering a wide field of research. Nevertheless, the distribution of neuropeptides in the brain of this avian species remains largely unknown. Here we present some of the first results on distribution of Vasotocine (AVT) and Vasoactive intestinal peptide (VIP) in the brain of males and females of this songbird species, using immunohistochemistry mapping. The bulk of AVT-like cells are found in the hypothalamic supraoptic, paraventricular and suprachiasmatic nuclei, bed nucleus of the stria terminalis, and along the lateral forebrain bundle. Most AVT-like fibers course toward the median eminence, some reaching the arcopallium, and lateral septum. Further terminal fields occur in the dorsal thalamus, ventral tegmental area and pretectal area. Most VIP-like cells are in the lateral septal organ and arcuate nucleus. VIP-like fibers are distributed extensively in the hypothalamus, preoptic area, lateral septum, diagonal band of Broca. They are also found in the bed nucleus of the stria terminalis, amygdaloid nucleus of taenia, robust nucleus of the arcopallium, caudo-ventral hyperpallium, nucleus accumbens and the brainstem. Taken together, these results suggest that both AVT and VIP immunoreactive structures show similar distribution to other avian species, emphasizing evolutionary conservatism in the history of vertebrates. The current study may enable future investigation into the localization of AVT and VIP, in relation to behavioral and ecological traits in the brain of tit species.

A Binary Genetic Approach to Characterize TRPM5 Cells in Mice.

Transient receptor potential channel subfamily M member 5 (TRPM5) is an important downstream signaling component in a subset of taste receptor cells making it a potential target for taste modulation. Interestingly, TRPM5 has been detected in extra-oral tissues; however, the function of extra-gustatory TRPM5-expressing cells is less well understood. To facilitate visualization and manipulation of TRPM5-expressing cells in mice, we generated a Cre knock-in TRPM5 allele by homologous recombination. We then used the novel TRPM5-IRES-Cre mouse strain to report TRPM5 expression by activating a τGFP transgene. To confirm faithful coexpression of τGFP and TRPM5 we generated and validated a new anti-TRPM5 antiserum enabling us to analyze acute TRPM5 protein expression. τGFP cells were found in taste bud cells of the vallate, foliate, and fungiform papillae as well as in the palate. We also detected TRPM5 expression in several other tissues such as in the septal organ of Masera. Interestingly, in the olfactory epithelium of adult mice acute TRPM5 expression was detected in only one (short microvillar cells) of two cell populations previously reported to express TRPM5. The TRPM5-IC mouse strain described here represents a novel genetic tool and will facilitate the study and tissue-specific manipulation of TRPM5-expressing cells in vivo.

Seasonal plasticity in the peptide neuronal systems: potential roles of gonadotrophin-releasing hormone, gonadotrophin-inhibiting hormone, neuropeptide Y and vasoactive intestinal peptide in the regulation of the reproductive axis in subtropical Indian weaver birds.

Two experiments examined the expression of gonadotrophin-releasing and inhibiting hormones (GnRH-I, GnRH-II and GnIH), neuropeptide Y (NPY) and vasoactive intestinal peptide (VIP) in subtropical Indian weaver birds, which demonstrate relative photorefractoriness. Experiment 1 measured peptide expression levels in the form of immunoreactive (-IR) cells, percentage cell area and cell optical density in the preoptic area (GnRH-I), midbrain (GnRH-II), paraventricular nucleus (GnIH), mediobasal hypothalamus [dorsomedial hypothalamus (DMH), infundibular complex (INc), NPY and VIP] and lateral septal organ (VIP) during the progressive, breeding, regressive and nonbreeding phases of the annual reproductive cycle. GnRH-I was decreased in the nonbreeding and VIP was increased in INc in the breeding and regressive states. GnRH-II and NPY levels did not differ between the testicular phases. Double-labelled immunohistochemistry (IHC) revealed a close association between the GnRH/GnIH, GnRH/NPY, GnRH/VIP and GnIH/NPY peptide systems, implicating them interacting and playing roles in the reproductive regulation in weaver birds. Experiment 2 further measured these peptide levels in the middle of day and night in weaver birds that were maintained under short days (8 : 16 h light /dark cycle; photosensitive), exposed to ten long days (16 : 8 h light /dark cycle; photostimulated) or maintained for approximately 2 years on a 16 : 8 h light /dark cycle (photorefractory). Reproductively immature testes in these groups precluded the possible effect of an enhanced gonadal feedback on the hypothalamic peptide expression. There were group differences in the GnRH-I (not GnRH-II), GnIH, NPY and VIP immunoreactivity, albeit with variations in immunoreactivity measures in the present study. These results, which are consistent with those reported in birds with relative photorefractoriness, show the distribution and possibly a complex interaction of key neuropeptides in the regulation of the annual reproductive cycle in Indian weaver birds.

Exploring avian deep-brain photoreceptors and their role in activating the neuroendocrine regulation of gonadal development.

In the eyes of mammals, specialized photoreceptors called intrinsically photosensitive retinal ganglion cells (ipRGC) have been identified that sense photoperiodic or daylight exposure, providing them over time with seasonal information. Detectors of photoperiods are critical in vertebrates, particularly for timing the onset of reproduction each year. In birds, the eyes do not appear to monitor photoperiodic information; rather, neurons within at least 4 different brain structures have been proposed to function in this capacity. Specialized neurons, called deep brain photoreceptors (DBP), have been found in the septum and 3 hypothalamic areas. Within each of the 4 brain loci, one or more of 3 unique photopigments, including melanopsin, neuropsin, and vertebrate ancient opsin, have been identified. An experiment was designed to characterize electrophysiological responses of neurons proposed to be avian DBP following light stimulation. A second study used immature chicks raised under short-day photoperiods and transferred to long day lengths. Gene expression of photopigments was then determined in 3 septal-hypothalamic regions. Preliminary electrophysiological data obtained from patch-clamping neurons in brain slices have shown that bipolar neurons in the lateral septal organ responded to photostimulation comparable with mammalian ipRGC, particularly by showing depolarization and a delayed, slow response to directed light stimulation. Utilizing real-time reverse-transcription PCR, it was found that all 3 photopigments showed significantly increased gene expression in the septal-hypothalamic regions in chicks on the third day after being transferred to long-day photoperiods. Each dissected region contained structures previously proposed to have DBP. The highly significant increased gene expression for all 3 photopigments on the third, long-day photoperiod in brain regions proposed to contain 4 structures with DBP suggests that all 3 types of DBP (melanopsin, neuropsin, and vertebrate ancient opsin) in more than one neural site in the septal-hypothalamic area are involved in reproductive function. The neural response to light of at least 2 of the proposed DBP in the septal/hypothalamic region resembles the primitive, functional, sensory ipRGC well characterized in mammals.

The nasal cavity and its olfactory sensory territories.

The nasal cavity and its olfactory sensory territories.

Possible functional role of olfactory subsystems in monitoring inhalation and exhalation.

In parallel with progress in understanding the canonical main olfactory system, a recent advance has accelerated elucidation of the structural organization and functional properties of the olfactory subsystems. These include the ganglion of Grueneberg, septal organ, and specific subsets of olfactory sensory neurons in the main olfactory epithelium. The emerging concept of the olfactory subsystems is that each subsystem expresses distinct classesofchemoreceptorsandsignaltransduction molecules, and thereby detects distinct categories of odor or pheromone molecules. The chemical signals detected by the subsystems are sent via their axons to subsystem-specific domains in the glomerular map of the main olfactory bulb (Mori and Sakano, 2011).

Dog and mouse: toward a balanced view of the mammalian olfactory system.

Although the most intensively studied mammalian olfactory system is that of the mouse, in which olfactory chemical cues of one kind or another are detected in four different nasal areas [the main olfactory epithelium (MOE), the septal organ (SO), Grüneberg's ganglion, and the sensory epithelium of the vomeronasal organ (VNO)], the extraordinarily sensitive olfactory system of the dog is also an important model that is increasingly used, for example in genomic studies of species evolution. Here we describe the topography and extent of the main olfactory and vomeronasal sensory epithelia of the dog, and we report finding no structures equivalent to the Grüneberg ganglion and SO of the mouse. Since we examined adults, newborns, and fetuses we conclude that these latter structures are absent in dogs, possibly as the result of regression or involution. The absence of a vomeronasal component based on VR2 receptors suggests that the VNO may be undergoing a similar involutionary process.

The nasal cavity of the sheep and its olfactory sensory epithelium.

Macro and microdissection methods, conventional histology and immunohistochemical procedures were used to investigate the nasal cavity and turbinate complex in fetal and adult sheep, with special attention to the ethmoturbinates, the vestibular mucosa, and the septal mucosa posterior to the vomeronasal organ. The ectoturbinates, which are variable in number and size, emerge and develop later than the endoturbinates. The olfactory sensory epithelium is composed of basal cells, neurons, and sustentacular cells organized in strata, but numerous different types are distinguishable on the basis of their thickness and other properties; all variants are present on the more developed turbinates, endoturbinates II and III. Mature neurons and olfactory nerve bundles express olfactory marker protein. We found no structure with the characteristics that in mouse define the septal organ or the ganglion of Grüneberg. Our results thus suggest that in sheep olfactory sensory neurons are exclusively concentrated in the main olfactory epithelium and (to a lesser extent) in the vomeronasal organ.