Comparative ultrastructural investigations of the pineal organ of the blind cave fish,Anoptichthys jordani,and its ancestor,the eyed river fish,Astyanax mexicanus

A comparative ultrastructural study has been made of the pineal organ in specimens of two closely related populations of the characid fish, Astyanaz mexicanus. The specimens of one population are living in the river, under natural light conditions. The specimens of the other population, originally described as Anoptichthys jordani, are living in a completely dark cave. In specimens of both populations the pineal organ consists of a spindle shaped end-vesicle, connected to the diencephalic roof by a slender stalk. The pineal tissue is compact and consists predominantly of glia-like supporting cells and sensory cells resembling the photoreceptor cells of the lateral vertebrate eye. Phagocytotic microglia-like cells can be found in close contact with the outer segments of the sensory cells. Nerve cells are located in the neighbourhood of neuropil formations, in which synaptic contacts are established between sensory cells and nerve cells. From these nerve cells fibers are emerging, forming the pineal tract that runs down the pineal stalk towards the diencephalon. On the basis of the ultrastructure described by other authors it is concluded that the pineal organ in specimens of the river population of Astyanax mexicanus resembles the pineal organ of other fish species. In specimens of the river population, reared under normal light-dark conditions for 3, 9 or 18 months, conspicuous morphological changes have not been detected in the presumably light-sensitive outer segments of the sensory cells or in other parts of the pineal tissue. In specimens of the cave populations, reared under identical conditions, an age-dependent, gradual regression of the regular outer segment organization of the pineal sensory cells takes place. In other parts of the pineal tissue, only small morphological changes can be observed. In specimens of the cave population, reared in constant darkness, the regression of the pineal outer segment organization begins earlier and is obvious. It is postulated that the gradual age-dependent regression of the regular organization of the outer segments in the pineal organ of cave specimens of Astyanax mexicanus is genetically determined and indicates a regressive evolution of the pineal light sensitivity. The expression of the regressive traits is dependent on the environmental light conditions.     

The parapineal and pineal organs of the elver (glass eel), Anguilla anguilla L.

Summary The parapineal organ of the glass eel (elver) consists of approximately 400 cells and is situated to the left of the connection of the pineal stalk to the third ventricle. A conspicuous nerve tract containing approximately 350 fibers arises from the parapineal organ and runs in spatial relationship to the habenular commissure toward the left habenular nucleus. The dominating cell type of the parapineal organ of the elver is a neuron (sensory neuron) of small diameter provided with atypical cilia (9×2+0, or rarely 8×2+0 types). Well-developed photoreceptor outer segments are lacking, and no interstitial cells of ependymal type have been observed with certainty in the parapineal organ. The axonal processes from the nerve cells form the tract leaving the parapineal organ.The pineal organ proper of the elver consists of photoreceptor cells with well-developed outer segments, interstitial cells of ependymal type, and ganglion cells. Axons from the latter form the pineal tract, which leaves the pineal organ and runs in close contact with the subcommissural organ toward the posterior commissure. The proximal part of the pineal stalk contains only a few photoreceptor cells the outer segments of which are less developed than those of the pineal body and the distal part of the pineal stalk.     

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.     

Immunocytochemical localization of serotonin and photoreceptor-specific proteins (rod-opsin,S-antigen) in the pineal complex of the river lamprey,Lampetra japonica,with special reference to photoneuroendocrine cells

Summary The pineal complex of the river lamprey, Lampetra japonica, was examined by means of immunocytochemistry with antisera against serotonin, the precursor of melatonin, and two photoreceptor proteins, rod-opsin (the apoprotein of the photopigment rhodopsin) and S-antigen. Serotonin-immunoreactive cells were observed in both the pineal and the parapineal organ. The proximal portion of the pineal organ (atrium) comprised numerous serotonin-immunoreactive cells displaying spherical somata. In the distal end-vesicle of the pineal organ, the serotonin-immunoreactive elements resembled photoreceptors in their size and shape. These cells projecting into the pineal lumen and toward the basal lamina were especially conspicuous in the ventral portion of the end-vesicle. In addition, single serotonin-immunoreactive nerve cells were found in this location. Retinal photoreceptors were never seen to contain immunoreactive serotonin; amacrine cells were the only retinal elements exhibiting serotonin immunoreaction. Strong S-antigen immunoreactivity was found in numerous photoreceptors located in the pineal end-vesicle. In contrast, the S-antigen immunoreactivity was weak in the spherical cells of the atrium. Thus, the pattern of S-antigen immunoreactivity was roughly opposite to that of serotonin. Similar findings were obtained in the parapineal organ. The rod-opsin immunoreaction was restricted to the outer segments of photoreceptors in the pineal end-vesicle and parapineal organ. No rodopsin immunoreactive outer segments occurred in the proximal portion of the atrium. Double immunostaining was employed to investigate whether immunoreactive opsin and serotonin are colocalized in one and the same cell. This approach revealed that (i) most of the rodopsin-immunoreactive outer segments in the end-vesicle belonged to serotonin-immunonegative photoreceptors; (ii) nearly all serotonin-immunoreactive cells in the end-vesicle bore short rod-opsin-immunoreactive outer segments protruding into the pineal lumen; and (iii) the spherical serotonin-immunoreactive cells in the pineal stalk lacked rod-opsin immunoreaction and an outer segment. These results support the concept that multiple cell lines of the photoreceptor type exist in the pineal complex at an early evolutionary stage.     

Ultrastructure of the pineal organ of the killifish,Fundulus heteroclitus,with special reference to the secretory function

Summary The pineal organ of the killifish, Fundulus heteroclitus, was investigated by electron microscopy under experimental conditions; its general and characteristic features are discussed with respect to the photosensory and secretory function. The strongly convoluted pineal epithelium is usually composed of photoreceptor, ganglion and supporting cells. In addition to the well-differentiated photosensory apparatus, the photoreceptor cell contains presumably immature dense-cored vesicles (140–220 nm in diameter) associated with a well-developed granular endoplasmic reticulum in the perinuclear region and the basal process. These dense-cored vesicles appear rather prominent in fish subjected to darkness. The ganglion cell shows the typical features of a nerve cell; granular endoplasmic reticulum, polysomes, mitochondria and Golgi apparatus are scattered in the electron-lucent cytoplasm around the spherical or oval nucleus. The dendrites of these cells divide into smaller branches and form many sensory synapses with the photoreceptor basal processes. Lipid droplets appear exclusively in the supporting cell, which also contains well-developed granular endoplasmic reticulum and Golgi apparatus. Cytoplasmic protrusions filled with compact dense-cored vesicles (90–220 nm in diameter) are found in dark-adapted fish. The origin of these cytoplasmic protrusions, however, remains unresolved. Thus, the pineal organ of the killifish contains two types of dense-cored vesicles which appear predominantly in darkness. The ultrastructural results suggest that the pineal organ of fish functions not only as a photoreceptor but also as a secretory organ.We thank Dr. Grace Pickford for the fishes.     

Ultrastructure and serotonin immunocytochemistry of the parietal-pineal complex in the Japanese grass lizard, Takydromus tachydromoides.

The fine structure and immunocytochemical localization of serotonin in the cells of the receptor line were studied in the parietal eye and pineal organ proper of the Japanese grass lizard, Takydromus tachydromoides. Typical photoreceptor cells (PC) were the predominant cell type in the receptor line of the parietal eye, the outer segments of which had regular stacks of numerous disks similar to those of cones. The pineal organ contained relatively few PCs, which showed less well-developed outer segments than those of the parietal eye. In contrast, secretory rudimentary photoreceptor cells (SRPC) accounted for the majority of receptor cells in the pineal organ. These cells were structurally characterized by whorl-like lamellar outer segments and numerous dense-cored vesicles (80-280 nm in diameter). A small number of SRPC were also found in the parietal retina, which were similar to those in the pineal organ. In the parietal-pineal complex, numerous mitochondria located in the PC were larger and rounder than those in the SRPC. In the PC, basal processes prossessed only synaptic ribbons, whereas in the SRPC some of these processes contained synaptic ribbons and others contained dense-cored vesicles, rarely having both. Serotonin-immunoreactive cells were found not only in the pineal organ but also in the parietal eye, which closely resembled the cells of the receptor line in their size and shape. Furthermore, on immunoelectron microscopy for serotonin using the protein A-gold technique, gold particles indicating serotonin-immunoreactive sites were restricted in the core of dense-cored vesicles in the SRPC of the pineal organ. Regional differences in the distributions of the PC, SRPC and serotonin-immunoreactivity were found in the parietal-pineal complex.     

Organ of Bellonci of an Antarctic crustacean,the marine isopod Glyptonotus antarcticus

The paired organ of Bellonci protrudes from the optic lobe of the giant Antarctic isopod, Glyptonotus antarcticus. It is linked to the cortex by a broad peduncle. No connection to the cuticle or “sensory pore organ” was found. A cluster of sensory-like cells forms two outer ciliary segments branching into numerous microvilli with microtubules. The putative sensory somata are irregular in shape and contain a very high density of glycogen granules. The two outer segments sprout from two pits of the soma in different directions, forming a right angle. Glial cells wrap around the sensory cells and also delimit lacunae into which bundles of microvilli project. These lacunae contain electron-dense granules of small size and with species-specific patterns. Lacunae and dense granules show features typical of a degeneration process in the sensory cells. This general morphology corresponds to the unilobular type of organ of Bellonci, known in other isopods; it differs from the plurilobular type with onion bodies found in other Crustacea.     

Acetylcholinesterase-containing nerve cells and their distribution in the pineal organ of the goldfish,Carassius auratus

Summary Histochemically, an intense acetylcholinesterase (AChE) reaction has been observed in the perikarya of the nerve cells and in the neuropil formations of the pineal organ in the goldfish, Carassius auratus. A group of AChE-rich nerve cells has also been observed between the caudal end of the pineal stalk and the habenular ganglion. No component of the complex revealed butyrylcholinesterase (BuChE) activity.Two different types of nerve cells were recognized on the basis of their size, AChE activity and distribution. Type I cells are characterized by large perikarya possessing a moderate AChE activity and by the presence of an extensive AChE-rich neuropil formation in their vicinity; they are restricted to the rostro-lateral regions of the pineal vesicle. Type II cells are situated in the medio-rostral area of the pineal vesicle and along the entire length of the stalk, and are smaller than Type I cells; they show an intense AChE activity in their perikarya.The neuropil formations in the medio-rostral area of the pineal vesicle are almost as large as those in the vicinity of the Type I cells; they exhibit a strong AChE activity. In the rostral half of the vesicle several sensory cells are associated with each nerve cell, while in the caudal portion only a few cells are apposed to each nerve cell. Thus, the ratio of the number of sensory cells to that of AChE-containing nerve cells in the anterior half of the pineal vesicle is high when compared with the remaining area. In the anterior half of the vesicle the outer segments of the sensory cells are more distinct and their inner segments possess a higher AChE activity than those in the posterior region and the stalk. A gradation in the degree of development of neuropil formations along the pineal axis is remarkable; their size and AChE activity gradually diminish in a caudal direction. In view of the structural specialization of the rostral region of the pineal organ, it has been argued that its terminal portion is more photosensitive.This work was supported by a fellowship from the Alexander von Humboldt Foundation, Federal Republic of Germany.     

The pineal complex of the three-spined stickleback,Gasterosteus aculeatus L.

Summary The pineal complex of the three-spined stickleback (Gasterosteus aculeatus L.) was investigated by light and electron microscopy, as well as fluorescence histochemistry for demonstration of catecholamines and indolamines. The pineal complex of the stickleback consists of a pineal organ and a small parapineal organ situated on the left side of the pineal stalk. The pineal organ, including the entire stalk, is comprised mainly of ependymal-type interstitial cells and photoreceptor cells with well-developed outer segments. Both unmyelinated and myelinated nerve fibres are present in the pineal organ. Nerve tracts from the stalk enter the habenular and posterior commissures. A small bundle of nerve fibres connects the parapineal organ and the left habenular body. The presence of indolamines (5-HTP, 5-HT) was demonstrated in cell bodies of both the pineal body and the pineal stalk, and catecholaminergic nerve fibres surround the pineal complex.     

Electron microscopic and experimental studies of the pineal organ in the white-crowned sparrow,Zonotrichia leucophrys gambelii

Summary The structure of the pineal organ of Zonotrichia leucophrys gambelii, as revealed by light- and electron-microscopy, resembles that of Passer domesticus (Oksche and Kirschstein, 1969; Ueck, 1970). The typical cellular element is the pinealocyte with certain basic structural features of the pineal photoreceptors of lower vertebrates (see Oksche, 1971). However, instead of the characteristic, cone-like outer segments, there are, as in other species of birds, only bulbous cilia with ectopic whorls of lamellae. This structure of the outer segment is, in a sense, contrary to the demonstration of synaptoid contacts, numerous unmyelinated, and occasional myelinated nerve fibers by electron microscopy. In Nissl preparations it was possible to demonstrate typical nerve cells. The pinealocytes of Z. l. gambelii are secretory; their Golgi complex forms granulated vesicles (800–1,400 Å in diameter) that belong to the group of granular inclusions characteristic of monoamines. Autonomie nerve fibers course within the connective tissue capsule of the pineal organ. In many pinealocytes of Z. l. gambelii, the granular endoplasmic reticulum contains extensively expanded cisternae that are filled with a flocculent material and closely associated with bundles of filaments. In a number of cases such loop-like structures are selectively stainable with aldehyde fuchsin. It was not possible to demonstrate specific secretory activity in the supporting cells. Extirpation of the pineal organ in Z. l. gambelii had no definitely detectable influence on the photoperiodic control of testicular growth.Aves, Passeriformes, Fringillidae.Supported by grants from the Deutsche Forschungsgemeinschaft to Professor Oksche and by the National Science Foundation (GB 11905) to Professor Farner. A part of this investigation was effected while Professor Kobayashi held a Visiting Professorship at the University of Giessen.     

Histological,histochemical and electron microscopical studies on the nervous apparatus of the pineal organ in the tiger salamander,Ambystoma tigrinum

Summary 150–190 photoreceptor cells form a basic structural component of the pineal organ of Ambystoma tigrinum. Most of the outer and inner segments of these cells project into the lumen horizontally. Only 10 percent of the total number of photoreceptor cells are located within the pineal roof which is composed of a single cell layer. The photoreceptor cells are connected with nerve cells by synapses displaying characteristic ribbons. Different types of synaptic contacts, i.e. simple, tangential, dyad, triad and invaginated, are found. They are embedded in extended neuropil zones. A particular type of synapse indicates the presence of interneurons. The basal processes of some photoreceptor cells leave the pineal organ and make synaptic contacts with nervous elements located within the area of the subcommissural organ. Employing the method of Karnovsky and Roots (1964) for histochemical demonstration of acetylcholinesterase (AChE) approximately 70 neurons (intrapineal neurons) can be discerned in the pineal organ of Ambystoma tigrinum. In analogy to the distribution of photoreceptor cells only few nerve cells are observed in the roof portion of the pineal organ. Evidently, two different types of AChE-positive intrapineal neurons are present. About 40–50 AChE-positive neurons (extrapineal neurons) are scattered in the area of the subcommissural organ. In this area two types of nerve cells can be distinguished: 1) neurons which send pinealofugal (afferent) axons toward the posterior commissure and 2) neurons which emit pinealopetal (efferent) axons into or toward the pineal organ.The nervous pathways connecting the pineal organ with the diencephalomesencephalic border area are represented by a distinct pineal pedicle and several accessory pineal tracts.Granular nerve fibers run within the posterior commissure and establish synaptic contacts in the commissural region adjacent to the pineal organ. Some of these granular elements enter the pineal organ.The morphology of the nervous apparatus of the pineal organ of Ambystoma tigrinum is discussed in context with evidence from physiological experiments.In partial fulfillment of the requirements for the degree of Dr. med., Faculty of Medicine, Justus Liebig University, GiessenThe author is indebted to Professors A. Oksche and M. Ueck for their interest in this study. Thanks are due to Professor Ch. Baumann, Giessen, and Professor H. Langer, Bochum, for stimulating discussions. The technical assistance of Miss R. Liesner is gratefully acknowledgedDedicated to Professor Berta Scharrer on the occasion of her 70th birthday. Supported by grants from the Deutsche Forschungsgemeinschaft to A.O. and M.U.     

Photoreceptor-like outer segments in the pineal organ of the lovebird,Uroloncha domestica (Aves: Passeriformes)

Summary In the pineal organ of the lovebird, Uroloncha domestica, bulbous, cup-shaped and elongated outer segments of photoreceptor-like pinealocytes are demonstrated by scanning electron microscopy. These scarce outer segments, 4–11 m in length, extend into the pineal lumen. The present structural observations speak in favor of photosensitive pinealocytes in the pineal organ of Uroloncha domestica. The relation of the photoreceptor-like pinealocytes to acetylcholinesterase-positive nerve cells and a nervous connection between the pineal and the brain indicate that the pineal organ of this passeriform species may be the site of neuroendocrine and photoreceptive functions.Supported by a fellowship from the Japan Society for the Promotion of Science to M. UeckSupported by a grant from the Ministry of Education of Japan to K. Wake and by a grant of the Deutsche Forschungsgemeinschaft to M. Ueck     

The structure of the postantennal organ in Onychiurus sp. (Insecta: Collembola) and its connection to the central nervous system

Summary The postantennal organ in Onychiurus (group armatus) is a sensory organ comprising one sensory cell, several enveloping cells and cuticular structures.The perikaryon of the sensory cell is located in the central nervous system and distally gives off a dendrite in which one inner and two outer segments are distinguishable. Two ciliary structures connect the outer dendritic segments with the inner segment. The outer segments divide repeatedly, basal to the cuticular structures, into small branches which end distally beneath the cuticular wall. The wall of the cuticular structures is very thin and is pierced by numerous funnel-shaped pores. The pores are filled with electron-dense material which forms a continuous sheath underneath the cuticle. This material encases the small dendritic branches and the processes of the enveloping cells which occupy the lumen of the cuticular structures. There are three types of enveloping cells: one inner, several outer and one basal. Their processes differ in the manner in which they envelop the various regions of the dendrite.At the beginning of moulting outer dendritic branches are not found within the cuticular structures of the organ. They may be assumed to retract inwardly. However, in the later stages, when the cuticle is fully formed, the outer dendritic segments appear to divide. It is assumed that the small dendritic branches reach their targets before ecdysis. The electrondense material which clogs the intermoult cuticular pores is absent until the final stages of the moulting cycle.Supported by a grant from the Deutscher Akademischer Austauschdienst.     

The pineal organ of Raja clavata: Opsin immunoreactivity and ultrastructure

Summary The pineal organ of Raja clavata was studied by light and electron microscopy, including the immunocytochemical antiopsin reaction. The pineal organ of the ray consists of three portions: (i) a large proximal pineal, (ii) a long tube-like connecting stalk, and (iii) a short distal terminal enlargement. This latter end-vesicle lies in the deep connective tissue layers of the braincase. All portions of the pineal are composed of pinealocytes, intrinsic neurons, ependymal/glial cells, and bundles of nerve fibers embedded in thin neuropil formations. The inner segments of the pinealocytes protrude into the lumen in all parts of the organ and usually contain basal bodies and numerous mitochondria. Often, two outer segments were found to arise from the basal bodies of a single inner segment. By means of light-microscopic immunocytochemistry the outer segments showed a strong antiopsin reaction.The axons of the pinealocytes form ribbon-containing synapses on dendritelike profiles, which appear to belong to the intrinsic pineal neurons. There are other axo-dendritic synapses established by presynaptic terminals lacking ribbons and containing granular and synaptic vesicles. Pineal neurons may contain granular vesicles approximately 60–100 nm in diameter; their processes contribute to the bundles of unmyelinated axons.The fine structural organization of the pineal organ and the opsin immunoreactivity of the outer segments of the pinealocytes indicate a photoreceptive capacity of the organ. The double outer segments represent a peculiar multiplication of the photoreceptor structures.This investigation was supported by grants from the Deutsche Forschungsgemeinschaft to A. Oksche (Ok 1/24; 1/25: Mechanismen biologischer Uhren)On leave from the 2nd Department of Anatomy, Semmelweis OTE, Budapest, Hungary     

Morphology of the pineal organ in the salamander Ensatina eschscholtzi

The pineal organ of Ensatina eschscholtzi, a terrestrial and secretive species of salamander of the family Plethodontidae, is a photoreceptive structure lying on the dorsal surface of the diencephalon. The pineal is flattened with a broad lumen and consists of three cell types: photoreceptors, supportive cells, and neurons. Pineal photoreceptors are typical vertebrate photoreceptors and possess outer segment formations which, however, are frequently contorted and disorganized. Sloughing of apical portions of outer segments and vesiculation along the lateral edges of outer segment membrane disks are consistently observed and presumed to represent mechanisms of outer segment membrane recycling. Photoreceptors have basal processes which synapse with neural dendrites. Synapses between photoreceptor basal processes are occasionally observed. All synapses are characterized by synaptic ribbon structures of variable number, size, and configuration. Dense-core vesicles are occasionally observed mingled with clear synaptic vesicles within photoreceptor basal processes. Supportive cells within the pineal function in phagocytosis and recycling of shed outer segment membrane material, and neurons are localized at the lateral margins of the organ. The latter send axons into the ipsilateral side of the dorsal diencephalon. The pineal organ of Ensatina shows marked variation in overall size (cell total), cell type proportions, absolute neuron number, and ratio of photoreceptor number to neuron number for individual pineals. None of these morphological parameters is correlated with body size, sex, or season, and it is assumed that such variability represents significant variation in photosensory capabilities. It is suggested that the pineal organ of Ensatina is a partially degenerate photoreceptive structure.     

Ontogenetic development of the pineal organ,parapineal organ,and retina of the three-spined stickleback,Gasterosteus aculeatus L. (Teleostei)

The ontogenetic developments of the pineal organ, parapineal organ, and retina were studied by the use of light and electron microscopy in embryos and fry of the teleost, Gasterosteus aculeatus, from 60 to 168 h after fertilization. Sixty to 66 h after fertilization, the primordium of the pineal complex is discernible in the diencephalic roofplate; the parapineal anlage is located rostral to the pineal anlage. Photoreceptor cells endowed with outer segments are present in the embryonic pineal organ already after 72 h, whereas outer segments of retinal photoreceptors could not be demonstrated before 144 h (hatching occurs between 120-144 h). Furthermore, neuropil formations with synaptic specializations are present in the rostral part of the pineal organ 108 h after fertilization. At 72 h, the embryonic parapineal parenchyma is already differentiated into parapinealocytes, which give rise to the parapineal tract, and glia-resembling elements. Although parapinealocytes carry cilia (9 X 2 + 0), only a single outer segment of the photoreceptor type could be demonstrated in the parapineal organ of one adult stickleback. Photoreceptors present in the pineal organ of unhatched embryos are hardly involved in visual functions, but may already at this early developmental stage serve as photoneuroendocrine transducers.     

Immunocytochemical localization of Vitamin A in the retina and pineal organ of the frog,Rana esculenta

Summary Vitamin A immunoreactive sites were studied in the retina and pincal organ of the frog,Rana esculenta, by the peroxidase antiperoxidase, avidin-biotinperoxidase and immunogold methods. Indark-adapted material, strong immunoreaction was found in the outer and inner segments of the photoreceptor cells of both retina and pineal organ, as well as in the pigment epithelium, retinal Müller cells and pineal ependymal cells. Inlight-adapted retina, cones and green (blue-sensitive) rods were immunopositive.At the electron microscopic level, immunogold particles were found on the membranes of the photoreceptor outer segments as well as on the membranes of the endoplasmic reticulum and mitochondria. Individual retinal photoreceptor cells exhibited strong immunoreaction in the distal portion of the inner segment, the ciliary connecting piece and the electron-dense material covering the outer segment. In the pigment epithelium, the immunolabeling varied in intensity in the basal and apical cytoplasm and phagocytosed outer segments.The immunocytochemical results indicate that retinoids (retinal, retinol and possibly retinoic acid) are present not only in the photoreceptor cells of the retina but also in those of the pineal organ. The light-dependent differences in the immunoreactivity of vitamin A underlines its essential role in the visual cycle of the photopigments. Our results suggest that the pineal ependyma plays a role comparable to that of the Müller cells and pigment epithelium of the retina with regard to the transport and storage of vitamin A. The presence of a retinoid in nuclei, mitochondria and cytoplasmic membranes suggests an additional role of vitamin A in other metabolic processes.Dedicated to Professor Dr. T.H. Schiebler on the occasion of his 65th birthdaySupported by the Hungarian OTKA grant Nr. 1619 to B.V., and a grant from the Pardee Foundation to G.H.W.     

TEM studies on the lattice organs of cirripede cypris larvae (Crustacea, Thecostraca, Cirripedia)

 Lattice organs consist of five pairs of sensory organs situated on the dorsal carapace in cypris larvae of the Crustacea Cirripedia. The lattice organs in cypris larvae of Trypetesa lampas (Acrothoracica) and Peltogaster paguri (Rhizocephala) represent the two main types found in cirripedes, but only minor differences exist at the TEM level. Each lattice organ is innervated by two bipolar, primary receptor cells. The inner dendritic segment of each receptor cell carries two outer dendritic segments. The outer dendritic segments contain modified cilia with a short ciliary segment (9×2+0 structure). Two sheath cells envelop the dendrite except for the distal ends of the outer dendritic segments. This distal end enters a cavity in the carapace cuticle and reaches a terminal pore situated at the far end of the cavity. The cuticle above the cavity is modified. In both species the epicuticle is partly perforated by numerous small pores and the underlying exocuticle is much thinner and less electron dense than the regular exocuticle. Lattice organs very probably have a chemosensory function and are homologous with the sensory dorsal organ of other crustacean taxa. Accepted: 18 August 1998     

The pineal-paraphyseal complex of sea turtles. I. Light microscopic description

The pineal-paraphyseal complex of sea turtles is an impressively large structure which projects dorsally and anteriorly above the prosencephalon. The complex was examined by light microscopy in several age classes of green sea turtles (Chelonia mydas) and from juvenile loggerhead turtles (Caretta caretta). The paraphysis is extensively fused to the distal portion of the pineal body, suggesting an interrelated function for these two tissues. No duct or canal was observed connecting the pineal lumen to the third ventricle. Two pineal cell types are described which appear to correspond to the neuroglial supportive cells and the secretory rudimentary photoreceptor cells of other amniotic vertebrates. A possible luminal secretion in the form of apical protrusions is produced by the latter cell type. No typical photoreceptive outer segments were observed.     

Immunocytochemical and electron-microscopic investigations of the pineal organ in adult agamid lizards,Uromastix hardwicki

Summary Lacertilian species display a remarkable diversity in the organization of the neural apparatus of their pineal organ (epiphysis cerebri). The occurrence of immunoreactive S-antigen and opsin was investigated in the retina and pineal organ of adult lizards, Uromastix hardwicki. In this species, numerous retinal photoreceptors displayed S-antigen-like immunoreactivity, whereas only very few pinealocytes were labeled. Immunoreactive opsin was found neither in retinal photoreceptors nor in pinealocytes. Electron microscopy showed that all pinealocytes of Uromastix hardwicki resemble modified pineal photoreceptors. A peculiar observation is the existence of a previously undescribed membrane system in the inner segments of these cells. It is evidently derived from the rough endoplasmic reticulum but consists of smooth membranes. The modified pineal photoreceptor cells of Uromastix hardwicki were never seen to establish synaptic contacts with somata or dendrites of intrapineal neurons, which are extremely rare. Vesiclecrowned ribbons are prominent in the basal processes of the receptor cells, facing the basal lamina or establishing receptor-receptor and receptor-interstitial type synaptoid contacts. Dense-core granules (60–250 nm in diameter) speak in favor of a secretory activity of the pinealocytes. Attention is drawn to the existence of receptor-receptor and receptor-interstitial cell contacts indicating intramural cellular relationships that deserve further study.Supported by the Deutsche Forschungsgemeinschaft (Ko 758/31) and the Deutscher Akademischer Austauschdienst (Senior DAAD Research Fellowship to M.A.H.)