The early development of the Jacobson's organ in the cat (Felis silvestris)

The histological study of the nose of an embryo of the cat (Felis silvestris 10 mm) presents a simple cavity which caudally communicates with the oral cavity by a narrow cleft, the primitive choana. This aperture results from the former rupture of the membrana bucconasalis of which the present material still contains remnants. Compared to the simple nasal cavity, the organ of Jacobson, situated at the base of the broad fetal nasal septum, shows a progressive development. From this one might conclude, according to ontogenetical rules, an early functional importance of the accessory olfactory organ. The currently fetal nose under investigation presents the organ as a primary open groove which increases in depth from rostral to caudal. Only caudally, a short section of the organ is already formed into a tube. From the functional point of view, the epithelial lining of this organ is still undifferentiated, but its dorsal part is clearly thickened. It seems that this indicates an early arrangement of the later olfactory epithelium. Beside this it is striking that in early fetal life, the organ of Jacobson extends to a remarkable length within the nasal cavity. Obviously the organ occupies, early in ontogeny, its area inside the little developed nose. Finally, the striking resemblance of the topography of the organ of Jacobson between early embryos of mammals and those of fetal and even some adult reptiles is discussed.     

The nasopalatine ducts and associated structures in the rhesus monkey (Macaca mulatta): topography, prenatal development, function, and phylogeny

Morphological and developmental characteristics of the rhesus monkey nasopalatine duct system and associated primary palatal structures are described along with functional and phylogenetic considerations. Examination of five adult palates and coronal sections of 13 fetal palates together with dissections of a sixth adult specimen and of a 119-day-old fetal palate reveal that the lateral lobes of the tripartate incisive papilla cover clefts leading into the ducts. The ducts pierce the bony palate to enter the nasal fossae in proximity to the incisive suture. The ontogenetic stability of the duct path reflects the retention of ancient duct and primitive choanae relationships and functionally maintains an optimal oral odorant-to-receptor channel. Sixteen timed pregnancy specimens (35-100 days) provided histological material for documenting rostral nasopalatal development. Duct primordia, identified at 35 days, had by 40 days formed the medial duct walls (conjoined septum-papilla from the primary medial palatal component), the lateral duct walls (maxillary processes), and the rostral walls (fused maxillary-intermaxillary components). The caudal walls derive from the fusion of palatal shelves with the papilla (45 days), thus distinguishing primary and secondary fusion modes. Duct epithelial maturation occurs between 70 and 100 days. The absence of a vomeronasal system is attributed to reduction of olfaction in reproductive behavior, while the presence of the coevolved nasopalatine ducts is linked to the persistence of epiglottal-velar valving. The ducts serve as oral food-odor conduits in otherwise functionally separated respiratory and digestive tracts.     

Late embryonic development of the vomeronasal complex of the cat (Felis silvestris)

An examination of 2 feline embryos in different stages of development (overall length 60 and 115 mm respectively) reveals a well developed vomeronasal complex in each case. Jacobson's Organs embedded within the paraseptal cartilage form long blind tubes at the base of the septum nasi. The cartilage is caudally tub-shaped and embraces rostrally completely the organ over a considerable length. In this manner a long, nearly tunnel-like tube is formed which represents a modified form of the original outer bar and which has not been described so far in cats. It stretches rostro-ventrally across the branching region of the paraseptal cartilage as far as the mouth of Jacobson's Organ. The dorsal branch of the cartilago paraseptalis on the other hand forms a vertically oriented strip which connects to the lamina transversalis anterior. The ductus nasopalatinus passing through the palate is laterally supported by a cartilago ductus nasopalatini which rostrally to the mouth of Jacobson's Organ forms a unified element with the ventral branch of the cartilago paraseptalis. In the case of the younger cat embryo, this cartilago ductus nasopalatini is yet weakly developed. The ductus nasopalatini of the embryos studied are in an amazingly retarded state of development. The ductus, which are blocked in the early stages of the embryonic development during secondary palate formation, form predominantly solid strands of epithelium. By dissolving the cemented epithelium, the ductus are open. But even in the case of the older embryo of the cat, this process is not completed yet. The short duct connecting Jacobson's Organ with the ductus nasopalatinus is also still closed in both embryos. Such cemented sections of epithelium of the younger embryo reveals an interesting relation between the ductus nasopalatinus and the ductus nasolacrimalis which so far has not been pointed out for mammals. From the point of view of phylogenetics, the locally specialized vomeronasal complex of cats exhibits all the criteria of a progressive development of characteristics.     

The histological structure and histochemistry of the mucosa of the nasal conchae in geese,Anser anser

We investigated the histological structure and histochemistry of the nasal conchae of geese and compared these structures with those of other avian species. The rostral, middle and caudal conchae were dissected from the nasal cavity of eight geese, fixed in Carnoy’s solution and embedded in paraffin. The entrance of the rostral concha was lined by keratinized stratified squamous epithelium, which toward the middle concha was replaced by modified keratinized squamous epithelium, the deep layer of which opened into tubular glandular structures containing secretory epithelium on crypt-like invaginations. The lamina propria of the rostral concha contained numerous Grandry’s and Herbst corpuscles, which are pressure-sensitive receptors peculiar to waterfowl. The lamina propria of the middle concha contained solitary lymphoid follicles and lymphocyte infiltrations. The cartilaginous component of the middle concha was highly convoluted and resembled a spiral of two and a half scrolls, which were lined by pseudostratified columnar epithelium. We observed that unlike mammals, this epithelium contained mostly intraepithelial alveolar glands rather than goblet cells. The caudal concha was similar to the middle concha, but less convoluted. It was lined by olfactory epithelium and its lamina propria contained serous Bowman’s glands as well as olfactory nerve fibers. Histochemical examination demonstrated that while none of the conchae contained sulfated mucins, except for the cartilage, the intraepithelial glands of the rostral and middle conchae contained mostly carboxylated acidic mucin and some neutral mucin, and were thus of the mixed type. The outermost scroll of the spiral of the middle concha contained some periodate-Schiff stained mucins. Of the glands of the mucosa of the middle concha, the deep tubuloalveolar glands in the convex parts of the scrolls contained primarily acidic mucins, while the shallow intraepithelial alveolar glands in the concave parts of the scrolls contained primarily neutral mucins. Our findings indicate that the rostral and caudal conchae primarily have a sensory function and the middle concha participates in mucosal defense.     

Comparative anatomical studies of the vomeronasal complex and the rostral palate of various mammals. I

The anatomy of the vomeronasal complex and, in connection with this, the structures of the rostral palate were studied in different species of mammals, namely members of the order Marsupialia, Scandentia, Insectivora, Primates, Rodentia, and Lagomorpha. The following results were obtained: The organs of Jacobson of all forms studied are well-developed. The organ of Jacobson is situated at the base of the nasal septum and opens rostrally, always closely connected to the nasopalatine duct. Even in rodents, lagomorphs and Solenodon, where the openings of the organs are positioned rostral to the ductus, both systems are nevertheless connected by means of special furrows. Accordingly the organs of Jacobson are functionally much more closely related to the oral cavity than to the nasal cavity, which they actually belong to. This can be emphasized by the peculiar structures of the rostral palate inclosing the papilla palatina and with it the oral openings of the nasopalatine ducts. In all species studied, the anterior part of the upper jaw presents a very interesting situation because the median furrow of the rhinarium communicates directly or indirectly with the sulcus papillae palatinae, thus forming a very distinct system of grooves which preserves a connection between the nasopalatine ducts and the preoral surroundings. In rodents, lagomorphs, and Solenodon, we find in this part of the palate a special situation because of their unusually arranged incisors, which are not separated by a diastema. However, also in these cases, there are distinct connecting passages between the papilla palatina and the extraoral surroundings. The conditions found in Ratufa bicolor and in early stages of the rat demonstrate that the extraordinary topography of the rostral palate in rodents is a secondary formation by means of ontogeny and phylogeny. Cebus apella, a platyrrhine simian, shows already a clear reduction of palatal structures compared to those found in prosimians. In Setifer setosus and Echinops telfairi, we find the papilla palatina and with it the oral openings of the nasopalatine ducts overgrown by a bipartite caudal branch of the rhinarium. The neonate Setifer allows us to reconstruct the mechanism of this overgrowing procedure. We find a similar situation in Erinaceus, where the papilla palatina remains uncovered, however. Because of contradictory bibliographical data, some elements of the vomeronasal complex in mammals needed to be carefully analysed in regard to structure and nomenclature: in many species the paraseptal cartilage bifurcates rostrally into a dorsal and a ventral branch.(ABSTRACT TRUNCATED AT 400 WORDS)     

Transitional epithelial zone of the bovine nasal mucosa

To determine the extent and ultrastructure of epithelium lining the transitional nasal mucosa of the neonate, gnotobiotic calf tissues were prepared for scanning and transmission electron microscopy. Stratified cuboid epithelium of the rostral 40% of the nasal cavity contained few ciliated cells; the next caudal 10-15%, although ciliated, had extensive nonciliated areas. The predominant type of surface cell was nonciliated, had short microvilli, and contained a multilobate nucleus and numerous pinocytotic vesicles. In some areas the surface of these cells presented a cobblestone appearance. Basal cells contained numerous bundles of filaments, ribosomes, and basal vesicles. Caudally, nonciliated columnar cells included a cell type similar to the more rostral cuboid cell, as well as brush cells and immature secretory and ciliated cells. Goblet cells were infrequently observed. Intraepithelial nerve terminals were abundant. Other intraepithelial cells, often difficult to identify owing to varying characteristics, included lymphocytes. Based upon comparisons of this neonatal epithelium with mature epithelium, observed in earlier studies of other mammalian species, the transitional mucosa is believed normally to occupy an extensive area of the nasal cavity.     

Peripheral projections of nervus terminalis LHRH-containing neurons in the tiger salamander,Ambystoma tigrinum

The peripheral projections of the nervus terminalis (NT) have been difficult to examine due to the weak immunoreactivity of the processes to various antibodies. We performed two experimental manipulations in the tiger salamander in an attempt to increase the luteinizing hormone-releasing hormone-immunoreactive (LHRH-ir) labelling in the peripheral processes of the NT: 1) the NT was sectioned centrally, or 2) a 100 mg melatonin pellet was embedded subcutaneously for 3 days prior to sacriffice. Following these manipulations, animals were sacrifficed and tissue was processed with standard immunocytochemical techniques for the analysis of the distribution of LHRH-ir processes. In the nasal cavity, LHRH-ir fibers were observed projecting 1) into the rostral olfactory epithelium, 2) to Bowman's glands in the lamina propria of the rostromedial olfactory mucosa and ventrolateral mucosa between the main nasal cavity and Jacobson's organ, 3) into the naris constrictor muscle, and 4) along the palatine nerves and ganglia. These lesion and hormone manipulations have enabled the detection of peripheral projections of the NT not observed previously with immunocytochemical procedures alone. The wide distribution of LHRH-ir NT processes in the nasal cavity and cranium suggests that this nerve may influence many different cranial structures during appropriate pheromonal or neuroendocrine events.     

Functional Architecture of the Vomeronasal Organ of the Frog (Genus Rana)

Abstract The vomeronasal organ in the frog, genus Rana, is composed of three interconnected cavities; superior, middle and inferior, which are separated from and anterior to the principal olfactory cavity. The superior cavity is found just underneath the external naris and forms a vestibule both for the principal olfactory organ and the vomeronasal organ. The vomeronasal sensory epithelium is located in the medial region of the inferior cavity and contains ciliated cells and microvillous receptor cells. Inspection of microscopic sections of frogs that had been swimming in fluorescent colorants revealed fluorescence on the surface of the vomeronasal organ, but not on that of the olfactory organ. Observations in vivo show that water enters via the external naris by two fissures, one on each side of the movable nasal lid, passes the middle cavity to flow via the sensory epithelium of the inferior cavity. The design of the frog nose makes it possible for this amphibious animal to sample the chemical composition of its environment; above water the frog can inhale air and expose its olfactory organ to volatile substances; in water the vomeronasal organ samples water-borne substances. These new findings are discussed in relation to the air/water interface and the position of the amphibians in the evolution of terrestrial vertebrates.     

Retinoic acid alters EGF receptor expression during palatogenesis

Various growth factors are necessary for normal embryonic development and EGF receptors are present in developing palatal shelves of embryonic/fetal mice at least from day 12 of gestation. The medial epithelium of the palatal shelf undergoes a series of developmental events which do not occur in the oral and nasal epithelia. In utero and in organ culture, the control palatal medial epithelium shows a developmental decline in EGF receptors, demonstrated both by a decrease in the binding of antibody to EGF receptors and a decrease in the binding of 125I-EGF; decreases which are not observed in cells of the adjacent oral or nasal epithelium. During this period, medial cells cease DNA synthesis and undergo programmed cell death. Medial epithelial cells exposed to all-trans-retinoic acid continue to express EGF receptors, bind EGF, proliferate, fail to undergo programmed cell death and exhibit a morphology typical of nasal cells. The data suggest that this disturbance by retinoic acid of EGF receptor localization and subsequent alterations in differentiation of the epithelial cells plays a role in the retinoic-acid-mediated induction of cleft palate.     

Evolution of the Nasal Structure in the Lower Tetrapods

The gross structure of the nasal cavities and the distributionof the various types of epithelium lining them are describedbriefly; each living order of amphibians and reptiles possessesa characteristic and distinctive pattern. In most groups thereare two sensory areas, one lined by olfactory epithelium withnerve libers leading to the main olfactory bulb and the otherby vomeronasal epithelium with fibers to the accessory bulb.All amniotes except turtles have the vomeronasal epitheliumin a ventromedial outpocketing of the nose, the Jacobson's organ,and have one or more conchae projecting into the nasal cavityfrom the lateral wall. Although urodeles and turtles possessthe simplest nasal structure, it is not possible to show thatthey are primitive or to define a basic pattern for either amphibiansor reptiles; all the living orders are specialized and the nasalanatomy of extinct orders is unknown. Thus it is impossible,at present, to give a convincing picture of the course of nasalevolution in the lower tetrapods.     

Morphological observations in normal primary palate and cleft lip embryos in the Kyoto collection

Normal developmental events during human primary palate formation and alterations associated with cleft lip remain poorly defined. The purpose of this study was to analyze serially sectioned human embryos to identify morphological changes during normal palatal closure and alterations associated with failure of palatal formation. Normal and cleft embryos from the histological collection at the Congenital Anomaly Research Center at the University of Kyoto were studied and photographed for detailed evaluation. Seven serially sectioned cleft lip embryos of stages shortly after primary palate formation (Streeter-O'Rahilly stages 19, 20, and 22) with unilateral or bilateral clefts with varying degrees of clefting were studied. In the normal Kyoto embryos, initial nasal fin (epithelial seam) formation was observed between the medial nasal process and the lateral nasal and maxillary processes at stage 17. During stages 18 and 19, the nasal fin epithelium was replaced by an enlarging mesenchymal bridge, as the maxillary processes united with the medial nasal processes to form the primary palate. The most prominent features observed in the cleft embryos were a reduced thickness of mesenchymal bridging between the medial nasal and maxillary processes, with an excessive amount of epithelium at the junctions between these processes. With ingrowth of the maxillary processes, greater cell dispersion and apparent extracellular matrix accumulation were observed in the medial nasal region. During closure of the primary palate, terminal branches of the maxillary nerve crossed the mesenchymal bridge to the medial nasal region. The partial clefts had reduced maxillary ingrowth and smaller union areas with the medial nasal process. Detailed studies of experimental animal models are required to identify regional growth required for contact between the facial prominences, to clarify the mechanisms of mesenchymal ingrowth and epithelial displacement during palatal formation, and to identify local and/or general factors causing alterations that lead to primary palatal clefting.     

Histological and carbohydrate histochemical studies of the nasal mucosa of sheep in Saudi Arabia with special reference to its glandular tissue

The histology and carbohydrate histochemistry of the nasal mucosa with attention to glandular tissue had been studied in 7 heads of sheep. Tissues were taken from vestibular region, septum at level of the alar fold, rostral portion of nasal conchae, caudal portion of nasal conchae, middle portion of septum and ethmoidal conchae region. Stratified squamous nonkeratinized epithelium was observed covering the vestibular region. The propria-submucosa of the nasal vestibule was richly permeated with glands having affinity for PAS and non-alcianophilic. The post-vestibular portion of the nasal cavity was lined by transitional epithelium and caudal to it, stratified columnar nonciliated epithelium was noticed. The respiratory epithelium covered the caudal half of the nasal conchae and the major portion of the septum as well as the recesses of the ethmoidal conchae. The glands associated with the respiratory mucosa were thick, coiled and tubular, containing both nonalcianophilic PAS positive and alcianophilic PAS positive cells. The olfactory mucosa covered the ethmoidal conchae and showed predominant serous glands. The results were discussed with that given for other mammals and in regard to the respiratory functions of the nasal mucosa.     

Données histologiques sur les fosses nasales et leurs annexes chezCrocodylus niloticus Laurenti etCaiman crocodilus (Linnaeus) (Reptilia,Crocodylidae)

Summary The vestibulum is very short and lined by stratified squamous epithelium which contains many alveolar cells. The cavum nasi proprium is exceedingly complex, with three conchal formations and a series of six recesses and sinuses. Olfactory epithelium lines the whole dorsal or dorso-medial half of the cavum, but not the deep sinuses. Non sensory respiratory epithelium lines the ventral or ventro-lateral half of the cavum, all the caviconchal recess, the posterolateral recess, the postturbinal sinus, the postconchal cavity, and the nasopharyngeal duct. In olfactory epithelium the proportion of sensory cells is about 61 % inCrocodylus and 59 % inCaiman; the ratio of sensory cells to supporting cells is about 2.6/1 in the former and 1.8/1 in the latter. Bowman's glands are sero-mucous and normally developed. As in other reptiles, the respiratory epithelium is composed with mucous and ciliated cells; but, in ventrolateral part of the cavum, there are also sero-mucous cells forming small multicellular glands. The hypertrophied lachrymal duct constitutes a very large naso-lachrymal not previously described gland. The lumen is lined by mucous and ciliated cells, the collet of each branched tubular gland by mucous cells and the glandular tubes by sero-mucous cells. Adult crocodilians lack a Jacobson's organ and there are no vomeronasal sensory epithelium in the cavum.In reptiles, aquatic way of life generally involves regression in olfactory epithelium, while Jacobson's organ (or, in Testudines, vomeronasal epithelium) persists and occasionally increases. In crocodilians, things are exactly reverse. After comparing with other Tetrapods, it seems likely that Jacobson's organ has been lost by terrestrial ancestor of crocodilians and birds. Now, only crocodilians posses olfactory epithelium, and naso-lachrymal gland gives them a supplementary protection, necessary in semiaquatic environment.     

Distribution of epithelia in the nasal cavity of piglets

The epithelial distribution in the nasal cavity of piglets was studied by serial transverse sections. The epithelial distribution in the nasal cavity of healthy piglets varied according to the age of the animal. The transitional epithelium, which contained goblet cells but no ciliated cells, occupied a smaller proportion of the nasal cavity in the newborn piglets than in the 4-week-old piglets. The ciliated epithelium extended more rostrally in the newborn piglets and covered the non-mineralized rostral portion of the nasal ventral concha. At 28 days of age, the rostral cartilaginous concha is overlaid by the transitional epithelium, the respiratory epithelium covering the mineralized nasal ventral concha. The variations in the epithelial distribution according to age are discussed with regard to the greater susceptibility of newborn piglets to bacterial infection.     

Fine structure of the lateral-line organ of the common eel,Anguilla japonica

Summary The sensory epithelium of the lateral line organ of the common eel consists of two types of cells, (sensory and supporting). The sensory cell bears a kinocilium together with about 40 to 60 stereocilia on its surface. The kinocilium is situated either at rostral or at caudal margin of this cilial group. Such polarity of the cilial group of one cell is inverse to that of an adjacent cell.Two types of crystal-like inclusions exist in the sensory cells, consisting of granules 100 Å in diameter. Granules in one type are arranged regularly whereas those in the other rather irregularly.Two types of nerve endings exist at the base of sensory cells: one is predominant in number and contains few vesicles, accompanied by a dense spherical body surrounded by small vesicles in the sensory cell and the other is rare in number and contains many vesicles, accompanied by a small flat sac just beneath the plasma membrane of the sensory cell.The supporting cells contain numerous mitochondria, a well developed Golgi apparatus and rough-surfaced endoplasmic reticulum, and surround a sensory cell completely. Physiologic significance of some of these components is discussed.     

The septal organ of the rat during postnatal development

The septal organ of Masera (SO) is a small, isolated patch of olfactory epithelium, located in the ventral part of the nasal septum. We investigated in this systematic study the postnatal development of the SO in histological sections of rats at various ages from the day of birth (P1) to P666. The SO-area increases to a maximum at P66-P105, just as the animals reach sexual maturity, and decreases thereafter, significantly however only in males, indicating a limited neurogenetic capacity for regeneration. In contrast, the main olfactory epithelium area continues to expand beyond P300. The modified respiratory epithelium ('zwischen epithelium') separating the SO and the main olfactory epithelium contains a few olfactory neurons up to age P66. Its length increases postnatally so that the SO becomes more ventral to the OE. Although the position of the SO relative to other anatomical landmarks changes with development it is consistently located just posterior to the opening of the nasopalatine duct (NPAL). Thus, a possible function of the SO is in sensing chemicals in fluids entering the mouth by licking and then delivered to the nasal cavity via the NPAL; therefore the SO may be involved in social/sexual behavior as is the vomeronasal organ (VNO). We suggest that the SO is a separate accessory olfactory organ with properties somewhat different from both OE and VNO and may exist only in species where the NPAL does not open into the VNO.     

Palatogenesis: morphogenetic and molecular mechanisms of secondary palate development

Mammalian palatogenesis is a highly regulated morphogenetic process during which the embryonic primary and secondary palatal shelves develop as outgrowths from the medial nasal and maxillary prominences, respectively, remodel and fuse to form the intact roof of the oral cavity. The complexity of control of palatogenesis is reflected by the common occurrence of cleft palate in humans. Although the embryology of the palate has long been studied, the past decade has brought substantial new knowledge of the genetic control of secondary palate development. Here, we review major advances in the understanding of the morphogenetic and molecular mechanisms controlling palatal shelf growth, elevation, adhesion and fusion, and palatal bone formation.     

绵羊犁鼻器在繁殖中的作用

许多动物的尿液中含有重要的外激素信息,其生理意义多种多样。雌性尿液中多有表示本身性别及性状况的化学信息存在,行为学调查表明,犁鼻器的主要功能在于识别尿液气味。公羊正是依据这些化学信息识别并选择配偶的。通常公羊嗅到母羊,尤其是发情母羊尿液时,表现伸颈、抬头、卷缩上唇的特有行为型式,称之为卷唇行为或性嗅反射。这个行为是与犁鼻器的功能相联系的。它的作用在于关闭外鼻孔,堵闭会厌,使吸进的空气进入犁鼻器内,犁鼻器两侧的肌肉运动,腔内静脉窦的膨胀和收缩,又促进空气的流入和排出,如此使携带化学信息的载体不断进入犁鼻器官,成为绵羊感受化学信息的方式之一。 犁鼻器的作用可为公羊及早地、准确地选择发情的配偶提供信息;同时又能唤起公羊本身的性行为,刺激雌、雄发情及性活动的同步,对保证繁殖的成功具有一定的意义。说明绵羊犁鼻器作为化学感受器在化学通讯中特别是对繁殖行为具有重要作用。     

Differential antigen expression during metamorphosis in the tripartite olfactory system of the African clawed frog, Xenopus laevis

In the adult African clawed frog, Xenopus laevis, olfactory epithelium is housed in three separate nasal cavities: the principal cavity, the middle cavity, and the vomeronasal organ. The sensory epithelium in each of these cavities has distinct cellular features, and presumed physiological and behavioral functions, which arise during metamorphosis. Most notably, the middle cavity is formed de novo, and the principal cavity is transformed from a larval sensory epithelium with water exposure to an adult olfactory epithelium with air exposure. To understand the cellular nature of this plasticity more clearly, we characterized the staining patterns generated in the olfactory system of X. laevis with a new monoclonal antibody, anti-E7. The olfactory epithelium is first stained with anti-E7 during late embryonic development. Transection of the olfactory nerves during metamorphosis eliminates all staining and indicates that the staining is associated with mature or nearly mature olfactory receptor neurons. The antibody diffusely stains the vomeronasal organ throughout development and in adults. In the larval principal cavity, the olfactory receptor neurons are brightly stained, but this cellular staining is lost after metamorphosis. The mucus from Bowman's glands in the principal cavity, however, is intensely stained in adults. The middle cavity, throughout development and in adulthood, has the same staining characteristics as the larval principal cavity. Thus, the E7 antibody can distinguish the three areas of the olfactory epithelium, allowing measurement of sensory epithelium volume, and serves as an excellent marker for the changes in the sensory epithelium that occur during metamorphosis.     

The sensory innervation of the pineal organ in the lizard,Lacerta viridis,with remarks on its position in the trend of pineal phylogenetic structural and functional evolution

Summary The sensory innervation of the pineal organ of adult Lacerta viridis has been investigated. Some specimens of Lacerta muralis lillfordi were also used. In the pineal epithelium, a small number of nerve cell pericarya of a sensory type are present. They lie either solitary or in small clusters close to the basement membrane. The axons originating from the nerve cell bodies, i. e. the pineal sensory nerve fibers, first course in the intraepithelial nerve fiber layer which is only locally present and contains a restricted number of unmyelinated fibers. In Lacerta viridis, the pineal fibers generally leave the epithelium at the proximal part of the organ proper. They then form small bundles which run along the outer surface of the basement membrane in the leptomeningeal connective tissue covering. At the proximal end of the pineal stalk the single bundles assemble constituting the pineal nerve. In Lacerta muralis the fibers leave the pineal epithelium at the proximal end of the stalk running farther down within the epithelium. Many fibers become myelinated after leaving the pineal epithelium. The pineal nerve runs ventralward in the midplane just caudal to the habenular commissure to which no fibers are given off. Continuing their ventralward course between the habenular commissure and the rostral end of the posterior commissure which is traversed by some of them, the pineal fibers reach the dorsal border of the subcommissural organ. Small separate aberrant pineal bundles traverse the posterior commissure at various more caudal levels. Having reached the dorsal border of the subcommissural organ, part of the pineal fibers continue their ventralward course directly running along the lateral sides of this organ to reach the periventricular nerve fiber layer lateral and ventral to it. A restricted number of fibers first turns in a caudal direction running between the base of the posterior commissure and the base of the subcommissural organ before turning ventralward to reach the periventricular layer. Most probably, pineal fibers do neither join the posterior commissural system nor innervate the subcommissural organ. Once having reached the periventricular layer, some pineal fibers curve in a rostral direction while others, before doing so, send a collateral in a caudal direction. Both, the main fibers and the collaterals, contribute to the formation of the periventricular layer. The sites of termination of the pineal fibers could not be ascertained.From the presence of intraepithelial sensory nerve cell bodies and from literature data on the ultrastructure of pineal neurosensory cells it is concluded that the adult pineal organ of Lacerta has a, although rudimentary, (photo)sensory function. The demonstration by our guest-worker Dr. W. B. Quay, of the intraepithelial presence of a tryptamine compound, probably serotonin, points, moreover, to a secretory function of this organ.In adult Lacerta a well-developed parietal nerve connects the parietal eye with the left lateral habenular nucleus. It traverses the habenular commissure.In gratitude and with admiration this paper is dedicated to Prof. Berta Scharrer and to the memory of Prof. Ernst Scharrer.