Septal Olfactory Organ Research Articles

Phylogenic Studies on the Olfactory System in Vertebrates

ABSTRACTThe olfactory receptor organs and their primary centers are classified into several types. The receptor organs are divided into fish-type olfactory epithelium (OE), mammal-type OE, middle chamber epithelium (MCE), lower chamber epithelium (LCE), recess epithelium, septal olfactory organ of Masera (SO), mammal-type vomeronasal organ (VNO) and snake-type VNO. The fish-type OE is observed in flatfish and lungfish, while the mammal-type OE is observed in amphibians, reptiles, birds and mammals. The MCE and LCE are unique to Xenopus and turtles, respectively. The recess epithelium is unique to lungfish. The SO is observed only in mammals. The mammal-type VNO is widely observed in amphibians, lizards and mammals, while the snake-type VNO is unique to snakes. The VNO itself is absent in turtles and birds. The mammal-type OE, MCE, LCE and recess epithelium seem to be descendants of the fish-type OE that is derived from the putative primitive OE. The VNO may be derived from the recess epithelium or fish-type OE and differentiate into the mammal-type VNO and snake-type VNO. The primary olfactory centers are divided into mammal-type main olfactory bulbs (MOB), fish-type MOB and mammal-type accessory olfactory bulbs (AOB). The mammal-type MOB first appears in amphibians and succeeds to reptiles, birds and mammals. The fish-type MOB, which is unique to fish, may be the ancestor of the mammal-type MOB. The mammal-type AOB is observed in amphibians, lizards, snakes and mammals and may be the remnant of the fish-type MOB.

Structure of Masera's Septal Olfactory Organ in Cat (Felis silvestris f. catus) - Light Microscopy in Selected Stages of Ontogeny

The septal organ /SO/ (Masera's organ /MO/) is a chemoreceptor presently considered one of three types of olfactory organs (along with the principal olfactory region and vomeronasal organ). Notwithstanding the septal organ having been first described by Rodolfo Masera in 1943, little is known of the properties of sensory neurons or of its functional significance in chemoreception. Until now the septal organ has been described only in laboratory rodents and some marsupials. This work refers to its existence in the domestic cat (Felis silvestris f. catus). The septal organ can be identified at the end of embryonic period - 27 or 28 days of ontogenesis in cats (the 6th developmental stage of Štěrba) - coincident with formation of the principal olfactory region in nasal cavity. At 45 days of ontogenesis (the 9th developmental stage of Štěrba), this septal olfactory organ is of circular or oval shape, 120 μm in diameter, in ventral part of septum nasi, lying caudally to the opening of ductus incisivus. The structure of the epithelium of septal olfactory organ is clearly distinct from the respiratory epithelium of the nasal cavity. It varies in thickness, cellular composition, as well as free surface appearance, and even lack the typical structure of sensory epithelium, in this developmental period. Nerve bundles and glandular acini are lacking in the lamina propria mucosae of the septal organ and in the adjacent tissues. Glands appear as the single non-luminized cords of epithelia extending from the surface. The adjacent respiratory epithelium contains numerous goblet cells.

Immunohistochemical studies on the differential maturation of three types of olfactory organs in the rats.

Differential maturation of three types of olfactory organs, the olfactory epithelium (OE), the vomeronasal organ (VNO) and the septal olfactory organ of Masera (MO), was examined immunohistochemically in embryonic and newborn rats by the use of antiprotein gene product 9.5 (PGP 9.5) serum. These olfactory organs were derived in common from the olfactory placode as neuroepithelia. In the OE, PGP 9.5-immunopositive olfactory cells first appeared at 13 days of gestation. The OE maturated completely, and showed the same cytological features as in the adult at 20 days of gestation. The MO first appeared as a dense mass of PGP 9.5-immunopositive sensory cells on the most ventrocaudal part of the nasal septum at 15 days of gestation and was evidently isolated from the OE by the decrease of immunopositive cells in the intercalated epithelium between the OE and the MO at 20 days of gestation. However, even at 7 days after birth, the MO did not complete its development and contained sensory cells aggregating in the mass. The VNO was separated from the nasal cavity at 13 days of gestation as a tubular structure of a neuroepithelium including PGP 9.5-immunopositive sensory cells. These cells gradually increased in number in the sensory epithelium of the VNO and extended their dendritic processes to the free surface at 7 days after birth. These findings clarified the differential maturation of these olfactory organs. That is, the OE completes its development before birth, while the MO and VNO after birth.

Analysis of the possible alerting function of the septal organ in rats: A lesional and behavioral study

The septal olfactory organ is a small patch of sensory epithelium located on the septal wall at the entry of the nasopharynx. There is a general consensus that by sampling olfactory stimuli during periods of rest, this organ may have an alerting function. To verify this hypothesis, we have lesioned by electrocoagulation the septal organ of male rats and recorded by polygraphy their awakening reaction in response to biologically meaningful (trimethyl-thiazoline, dimethyl sulfite, food) and meaningless (geraniol, eucalyptol) odorants. The awakening reactions of both lesioned and intact rats in response to these odorants were studied according to 3 parameters, frequency, latency and duration of awakening, and were analyzed using three-way analyses of variance. Data show that no significant difference in the awakening reactions was observed between control and lesioned animals. In all cases, the biologically meaningful odors presented the highest awakening influence. In addition, two of these odors (trimethyl-thiazoline and dimethyl sulfite) elicited a later habituation in comparison to biologically meaningless odors. From our results, it could be inferred that the hypothesis regarding an alerting function that would be specific to the septal organ, appears no longer current.

Lectin-binding patterns of the olfactory receptors (olfactory epithelium, vomeronasal organ and septal olfactory organ of Masera) in the rat

Lectin-binding patterns of the three olfactory receptors, olfactory epithelium (OE), vomeronasal organ (VNO) and septal olfactory organ of Masera (MO), and their associated glands, Bowman's gland (BG) of the OE, Jacobson's gland (JG) of the VNO, and anonymous gland (MG) of the MO were examined in the rat with 21 biotinylated lectins. On the free surface of the receptors, all lectins showed moderate to intense staining in at least one of receptors, although PHA-E in the VNO, BSL II and PNA in the MO, and DBA, BSL I, VVA, SJA, PSA and LCA in both the OE and the MO showed no staining. In the soma of receptor neurons, 16 lectins showed staining in at least one of receptors, although SJA showed no staining in both the OE and the MO, and BSL I showed staining in a small number of cells in these receptors. In the associated glands, 15 lectins showed staining in at least one of them, although PNA in the JG, LEL in the MG, and s-WGA and DBA in both the BG and the MG showed no staining. These findings suggest that the VNO takes charge of an olfactory discrimination different from that of the OE and MO, although the latter two take charge of that similar to each other.

Immunohistochemical demonstration of protein gene product 9.5 (PGP 9.5) in the primary olfactory system of the rat

Immunohistochemical localization of protein gene product 9.5 (PGP 9.5) was examined in the primary olfactory system of the rat. Receptor cells were intensely immunopositive in the olfactory epithelium, vomeronasal organ and septal olfactory organ of Masera. The mitral cells and some of the short-axon cells of the main olfactory bulb (MOB) were also intensely immunopositive, while the tufted cells of the MOB and the mitral/tufted cells of the accessory olfactory bulb varied in immunoreactivity.

Fine structure of the septal olfactory organ of Masera and its associated gland in the golden hamster.

Fine structures of the septal olfactory organ of Masera (MO) and its associated gland, a kind of olfactory glands, were examined in the golden hamster in comparison with those of the olfactory epithelium (OE) and vomeronasal organ (VNO) and their associated olfactory glands. Bipolar neurons of the MO were divided into two types according to their apical morphology, one similar to the receptors of the OE and the other to those of the vomeronasal sensory epithelium (VSE). The former was dominant and interpreted as main receptors of the MO. The other was less in number, but also regarded as a kind of olfactory receptors. The ultrastructural features of supporting cells of the MO were similar to those of the OE rather than to those of the VSE, while those of basal cells were almost in common in the OE, VSE and MO. The associated gland of the MO was positive to both PAS and alcian blue as the Bowman's gland of the OE. The fine structure of the associated gland of the MO was also similar to that of Bowman's gland. The present findings on the fine structure of the MO and its associated gland strongly suggest that the MO fulfills an olfactory function similar to that of the OE.

Fine structure of the vomeronasal and septal olfactory epithelia and of glandular structures.

The vomeronasal and septal olfactory organs are two neurosensory structures in the mammalian nasal septum which are poorly understood relative to the main olfactory system. The vomeronasal organ is a paired, blind-ending tubular structure that opens rostrally into the nasal cavity in some species and into the incisive ducts in others. When present in mammals, the septal olfactory organ is an island of olfactory mucosa positioned such that it is in the primary air pathway in the caudal portion of the nasal cavity. Mammalian nasal glands, with a diverse histochemical and ultrastructural morphology, secrete a variety of substances onto the mucosal surface. One of these substances, odorant binding protein, localized in bovine nasal glands and lateral nasal glands of rodents, may be important in the capture and conveyance of odorant molecules to olfactory receptors. The objectives of this paper are to present original data while reviewing the literature on the ultrastructure of vomeronasal and septal olfactory neuroepithelia, and of vomeronasal, bovine nasal, and lateral nasal glands. Nasal tissues from pigs, calves, and hamsters were prepared for electron microscopy. Neurosensory epithelia of the porcine vomeronasal organ and the hamster septal olfactory organ are similar to that described for the vomeronasal and septal olfactory organs of other mammals. Bovine nasal and rodent lateral nasal glands consist of subregions which differ morphologically; the most abundant acinar cell type in the bovine nasal gland contains lightly electron dense secretory granules while that of the rodent lateral nasal gland contains both small electron dense and large, electron lucent granules. The porcine vomeronasal gland contains numerous small, dense granules of a diverse morphology.

Calcitonin-gene-related-peptide-immunoreactive innervation of the rat head with emphasis on specialized sensory structures.

The distribution of calcitonin-gene-related peptide-like immunoreactivity (CGRP-IR) was studied in sections of decalcified rat head and selected whole-mount preparations in order to address the complex peptidergic innervation patterns in peripheral cephalic specialized zones and to examine neuronal ganglia in situ. Labeled neuron somata in trigeminal, glossopharyngeal, and vagal ganglia comprised a large proportion of small to medium size type B ganglion cells. Parasympathetic ganglia (ciliary, otic, sphenopalatine, submandibular) revealed a small population of labeled somata and numerous perisomatic IR axons, whereas sympathetic ganglion cells (superior cervical) were devoid of label though richly innervated by perisomatic IR axons. The gustatory geniculate ganglion contained only a few labeled neurons and axons. Coarse peripheral CGRP-IR axons were traced to skeletal muscle motor end plates (e.g., lingual, tensor tympani, etc.), and thin sensory axons most densely innervated the cornea, iris, general integument, all mucosal epithelia lining the tympanic, nasal, sinus and oropharyngeal cavities, and the cerebral meninges. Blood vessels, glands, ducts, and their orifices were often heavily innervated, and specific specializations and exceptions are discussed. Distinctive patterns of IR innervation characterized the various specialized sensory systems, including 1) cochlear and vestibular hair cells; 2) lingual, palatal, oropharyngeal, and laryngoepiglottal taste buds; 3) main olfactory epithelium and axons projecting to glomeruli in specific sectors of main olfactory bulb; 4) septal-olfactory organ; 5) vomeronasal organ; and 6) the nervus terminalis system. Secretory epithelia (ciliary body, choroid plexus, and stria vascularis) were notably lacking in CGRP-IR. Despite the multiplicity of functionally distinct CGRP neuronal and axonal populations, certain generalizations merit consideration. The extensive innervation of chemosensory nasal and oral epithelia may contribute to specific chemical sensitivities (e.g., relating to olfactory and gustatory senses) as well as evoking "nociceptive" responses to chemical irritants as part of a "common chemical sense." An efferent role for some of these peptidergic afferent axons may also be inferred from their specific distributions. Sites involved in regulating access to and sensitivity of sense organs to external stimuli (e.g., cochlear and vestibular hair cells, taste bud orifices, and main olfactory epithelium) are heavily innervated. Other IR axons are in position to exert control over airflow through nasal turbinates, glandular secretion, blood circulation, and duct transport systems.(ABSTRACT TRUNCATED AT 400 WORDS)

Masera's organ responds to odorants

Electroolfactogram (EOG) recordings from the rat septal olfactory organ (SO) provide the first demonstration of its broad-range chemosensitivity, and clearly establish this structure as a functioning component of the mammalian intranasal chemosensory system. SO sensitivity to lower concentrations of at least one common test odorant (pentyl acetate) exceeds that at sites located on the septal portion of the main olfactory neuroepithelium. Signals from the SO, as first proposed, thus could have an alerting function and provide information relebant to odor stimulus assessment.

Freeze-fracture study of the plasma membranes of the septal olfactory organ of Masera

The olfactory border and the apical cell contacts of the organ of Masera (MO) of the mouse were investigated by freeze-fracture electron microscopy. The olfactory border is mainly composed of the terminals of receptor and supporting cells. Cells with thick microvillus-like projections, though less frequent than the other two cell types, also contribute to the border. Olfactory knobs show transitions between those displaying numerous cilia and those characterized by few or no cilia. The olfactory cilia have a typical necklace of 6-9 rows of particles. The eruption of developing cilia seems to be preceded by the formation of circular arrays of particles. The density of intramembranous particles (IMP) per micron2 in P- and E-faces of the ciliary membranes is 1095 +/- 190 and 205 +/- 65, respectively. In the microvilli of supporting cells, the density of IMP per micron2 is 1800 +/- 270 for the P-face and 570 +/- 135 for the E-face. At the base of the supporting cell microvilli, rod-shaped particles are observed. The lateral plasma membranes of these cells bear orthogonal arrays of particles. In the apical region of the MO neuroepithelium, extensive zonulae occludentes are present which seal the intercellular cleft. The zonulae occludentes between supporting and receptor cells are composed of 5-13 junctional strands, usually arranged in an elongate network. Zonulae occludentes between supporting cells are, in addition to the elongate network, also arranged in a mesh-like pattern. Gap junctions, both associated with the zonulae occludentes and independent of them, are occasionally found between supporting cells. The results obtained indicate that important similarities exist between the neuroepithelium of the MO and the olfactory epithelium proper, whereas remarkable differences exist between the MO and the vomeronasal neuroepithelium.

Demonstration of terminalis, olfactory, trigeminal and perivascular nerves in the rat nasal septum.

The innervation of the nasal septum and around the olfactory bulb has been investigated in rats by means of whole-mount preparations and histological sections. Silver staining, OsO4 staining, PAS staining, cholinesterase reaction and fluorescence for catecholamine-containing nerves were used. The nervus terminalis forms on the medial side of the olfactory bulb a ganglionated plexus, from which branches are given off which course peripherally with the vomeronasal nerves. From a dorsal part of the terminalis nerve plexus an anterior branch is given off which runs along the anterior ethmoidal nerve to the nasal vestibule where it connects with a group of ganglia. The peripheral branches of the nerve run from here along two epithelial cristae formed histologically like dermal papillae. Ventrally in the respiratory region at the junction of the nasal cavity and the nasopharynx is a 1 times 2 mm area with olfactory epithelium, glands of Bowman and an independent innervation from the olfactory bulb. This is the so-called septal olfactory organ. Trigeminal nerves form a plexus in the respiratory region and in the vestibule, but do not supply the olfactory region. Catecholamine-containing and cholinesterase-positive nerves run along the meningeal arteries on the cribriform plate and accompany their branches to the vascular plexus in the olfactory and respiratory regions. Double innervation is found not only of this vascular plexus but of the venous sinuses in the swell bodies of the vestibule. The glands of the nose are not surrounded by catecholamine-containing nerves.

Septal olfactory organ in Peromyscus

1. 1. Compound light and scanning electron microscopic examination of the septal olfactory organ revealed a mucosal structure similar to that of the olfactory recess. 2. 2. Structures observed in scanning electron micrographs were interpreted as olfactory cilia, secretory droplets and duct orifices of olfactory glands. 3. 3. The area of epithelium occupied by the septal organ was highly variable both between individual mice and between contralateral septal surfaces of the same animal.