The origin of somatostatin fibers in the median eminence and neural lobe of Rana temporaria

Summary A light microscopic immunocytochemical study of the brain of frogs with hypothalamic lesions was performed in order to obtain evidence concerning the origin of somatostatin fibers in the median eminence and neural lobe of the hypophysis. The results indicate that the somatostatin fibers of the neural lobe originate from somatostatin perikarya located in the prechiasmatic part of the hypothalamus and possibly also in the telencephalon. The somatostatin neurons of the pars ventralis tuberis cinerei do not send axons to the neural lobe. The frog median eminence contains axon terminals of somatostatin neurons located in the pars ventralis of the tuber cinereum. Many other somatostatin fibers of the frog median eminence originate from somatostatin neurons located outside the tuber cinereum. Most of these neurons probably lie in the preoptic hypothalamic region.This investigation was supported by a grant from the Belgian Nationaal Fonds voor Geneeskundig Wetenschappelijk Onderzoek     

Experimental identification of a hypothalamic gonadotropic centre

Summary In a large group of adult female and male specimens of Rana temporaria all nervous pathways to the median eminence and hypophysis were extirpated without disturbing the normal blood supply of the median eminence and hypophysis. The total interruption of the nervous pathways persisted in all animals. In all animals gametogenesis and seasonal development of the gonads and of the secondary sexual characteristics were absent. In another large group of animals, the pars ventralis of the tuber cinereum of the hypothalamus was isolated from the brain. The pars ventralis retained its normal connections with the median eminence and hypophysis. The normal blood supply of the whole isolated region (pars ventralis tuberis + median eminence + hypophysis) was preserved. Four months after operation, in the majority of the animals, the isolation was still complete. In all animals, normal gametogenesis and seasonal development of the gonads and of the secondary sexual characteristics had occurred.In accordance with previous experiments, the present experiments showed that the pars ventralis tuberis contains an important gonadotropic centre. The activity of the gonadotropic centre of the pars ventralis tuberis is, at least, mainly exerted by its influence on the gonadotropic function of the pars distalis of the hypophysis. This is shown morphologically by a gradient of nuclear volume and by the mean nuclear volume of the P.A.S. positive cells of the pars distalis. The influence of the gonadotropic centre on the pars distalis is exerted by A.F.-negative nerve fibres to the median eminence. It is highly probable that all these nerve fibres end on the primary capillary network of the hypophysial portal system. The experiments strongly suggested that the axon endings of the gonadotropic centre would influence the gonadotropic activity of the pars distalis of the hypophysis by releasing of gonadotropic releasing factors into the blood capillaries of the median eminence.The gonadotropic centre of the pars ventralis tuberis is necessary for the normal gametogenesis and for the seasonal development of the gonads and of the secondary sexual characteristics. So far as concerned the gametogenesis and the seasonal development of the gonads, the function of the gonadotropic centre is largely autonomous.Exclusion of the function of a part of the pars ventralis tuberis produces quantitative, but no qualitative changes in gametogenesis.     

The gonadotropic centre of the tuber cinereum hypothalami and ovulation

Summary In adult female specimens of Rana temporaria, the pars ventralis of the tuber cinereum of the hypothalamus was isolated from the brain. The isolated pars ventralis retained its normal connections with the median eminence and hypophysis. The normal blood supply of the whole isolated region (pars ventralis tuberis + median eminence + hypophysis) was preserved.The results showed (1) that, in accordance with previous experiments, as far as concerned gametogenesis, seasonal development of the gonads and of the secondary sexual characteristics, the function of the gonadotropic centre of the pars ventralis tuberis is largely independent of nervous connections between the pars ventralis of the tuber cinereum and the brain; (2) that, in animals in which the pars ventralis tuberis was isolated from the brain at the beginning of the seasonal development of the ovaries, the L.H. secretion is disturbed during the ovulation period; (3) that consequently, in such animals, the stimulating activity of the isolated gonadotropic centre on the L.H. secretion of the hypophysis is too weak to cause ovulation; (4) that nervous connections between the isolated region and the preoptic region of the brain are indispensable for ovulation.     

The origin of the aldehyde-fuchsin-negative nerve fibres of the median eminence of the hypophysis: a gonadotropic centre

Summary In a large number of adult female and male specimens of Rana temporaria, the pars ventralis of the tuber cinereum of the hypothalamus was completely isolated from the brain. The pars ventralis retained its normal connections with the hypophysis. The vascular supply of the whole isolated region was left intact. Two and a half months after operation, in some animals, the interruption of the nervous pathways was still complete. In these animals it could be shown that all or nearly all A. F.-negative nerve fibres of the median eminence are axons of nerve cells localized in the area periventricularis of the pars ventralis tuberis. According to our results in Rana, the same A. F.-negative nerve fibres of the higher vertebrates are probably also axons of homologous nuclei of the tuber cinereum.In accordance with previous experiments, it could be concluded that the nerve cells of which the A. F.-negative fibres are axons, form an important centre that, via the pars distalis of the hypophysis, is responsible for the seasonal development of the ovaries, eggs and oviducts in the females and for the seasonal development of the thumb pads in the males.     

Fine structure of the median eminence and the pars nervosa of the bullfrog,Rana catesbeiana

Summary The hypothalamic neurosecretory system of the bullfrog, Rana catesbeiana, was studied with light- and electron microscopy. The median eminence is roughly divided into two portions. The upper portion mostly consists of ependymal cells, glial cells and preoptico-hypophysial nerve tract, whereas in the lower portion, neurosecretory axons, glial cells, processes of glial and ependymal cells, and fine blood vessels of the hypothalamic portal vein are located. A part of the neurosecretory axons of the preoptico-hypophysial tract proceeds to the lower portion of the median eminence. These axons are arranged perpendicularly to the capillaries of the hypothalamic portal vein. The glial cells are densely located in the area of the median eminence where neurosecretory material is abundant. The neurosecretory material in the neurosecretory cells, their axons, the median eminence and the pars nervosa of the bullfrog shows a positive reaction to PAS treatment.The neurohemal area of the median eminence is occupied by many neurosecretory and non-neurosecretory axons, containing neurosecretory granules and/or synaptic vesicles. The axonal portions with the synaptic vesicles which are considered to be the nerve endings abut on the capillaries of the portal system. The size of synaptic vesicles in the axon terminals containing few neurosecretory granules is larger than those in the endings with many neurosecretory granules. Infrequently glial and ependymal processes are interposed between the nerve endings and the capillary wall.In the hilar region of the infundibulum, synapses are frequently observed between the thin fibers with or without neurosecretory granules and dendrites of non-neurosecretory neurons. The probable functions of these synapses are briefly discussed on the basis of our findings. Both in the hilar region of the infundibulum and in the pars nervosa, electron-dense neurosecretory granules of two different sizes were observed. The median eminence contains only one type of granules.The fine structure of the pars nervosa shows similar structures to those of the median eminence. Both in the median eminence and the pars nervosa, the fenestrated endothelium of the capillaries was frequently observed. The thick perivascular connective tissue space containing fibroblasts and collagen fibrils was observed both in the median eminence and the pars nervosa. Vesicles in the cytoplasm of the endothelial cells which appear to take a part in the transendothelial transport were observed.This investigation was supported in part by United States Public Health Service Research Grant, No. A-3678, to Hideshi Kobayashi from the National Institute of Arthritis and Metabolic Diseases and partly by a grant for Fundamental Scientific Research from the Ministry of Education of Japan. The authors wish to express their thanks to Prof. K. Takewaki for his kind encouragement.     

Immuno-cytochemical demonstration,in the external region of the amphibian median eminence,of separate vasotocinergic and mesotocinergic nerve fibres

Summary By means of single and double immuno-enzyme cytochemical staining techniques, it was shown that the external region of the amphibian median eminence contains separate vasotocinergic and mesotocinergic nerve fibres. Moreover, it was demonstrated that the vasotocinergic fibres also contain neurophysin. In animals in which the hypothalamic magnocellular neurosecretory preoptic nuclei had been completely removed, the immuno-reactive vasotocinergic and mesotocinergic fibres of the median eminence had disappeared. From this result, it is concluded that, at least the great majority of the vasotocinergic and mesotocinergic fibres of the external region of the amphibian median eminence are processes of neurosecretory perikarya located in the hypothalamic magnocellular preoptic nuclei. On the other hand, our results do not exclude the possibility that a minority of these neurosecretory fibres originate from small immuno-reactive perikarya which were found in the tuber cinereum. The observation that both kinds of processes accumulate around blood capillaries of the hypophysial portal system strongly suggests that they play a role in the control of the activity of the pars distalis of the hypophysis.This investigation was supported by a grant from the Belgian Nationaal Fonds voor Geneeskundig Wetenschappelijk Onderzoek     

Identification of adenohypophysiotropic neurohormone producing neurosecretory cells in Rana temporaria

Summary Electron microscopic investigation of neural isolated and normal pars ventralis of the tuber cinereum showed the presence, in Rana temporaria, of a tubero-hypophysial neurosecretory system. A striking structural resemblance between this parvicellular, aldehydefuchsin negative, tubero-hypophysial neurosecretory system and the magnocellular, aldehydefuchsin positive, hypothalamo-hypophysial neurosecretory system was observed. Six different neurosecretory cell types are described, characterized by different shape and size of their respective secretory granules. The nature of the secretory product of these cells is briefly discussed.Of the results of this investigation, a preliminary note has been published (Dierickx, 1971).The authors wish to thank Prof. Dr. H. Steyaert and Dr. W. Moerdijk of the Laboratory for Mathematical Statistics, Ghent University, for invaluable advice and assistance with their computers. They thank Dr. G. De Waele for his help with the scanning electron microscopy.     

Electron microscopic and pharmacological studies on the rat median eminence

Summary The rat median eminence contains at least three kinds of granules or vesicles: 1. large electron-dense granules (perhaps carriers of neurohypophysial hormones), 2. small electron-dense granules with or without haloes (perhaps carriers of catecholamines) and 3. synaptic vesicle-like structures (perhaps carriers of acetylcholine). The former two electrondense granules exist in separate axons but they coexist with the latter vesicles in the same axons.The pars nervosa shows basically a similar structure to the median eminence. However, the axons containing the small electron-dense granules are very few. In the pars tuberalis, there are at least two types of cells: the cells of one type contain much cytoplasm with large round nuclei and those of the other type contain a small amount of cytoplasm with polymorphic nuclei. The cells of the former include multivesicular bodies and secretory granules, but those of the latter do not. Some of capillaries of the primary plexus are surrounded by the cells of the pars tuberalis on one side and by neurosecretory axon endings on the other side.The median eminence contains high concentration of acetylcholine or an acetylcholine-like substance and shows neurohypophysial hormone activity.Aided by Grant A-3678 from the United States National Institute of Arthritis and Metabolic Diseases. The authors are indebted to Dr. Welsh, Harvard University, for the kind gift of Mytolon.     

The nerve fibres controlling the gonadotropic activity of the hypophysis of Rana temporaria

Summary In 27 adult female specimens of Rana temporaria, the total interruption of the nervous pathways to the median eminence and the hypophysis was performed, without disturbing the normal blood supply of the hypophysis. In eight cases this complete interruption persisted three months after operation. In these animals there occurred no seasonal development of the eggs, ovaries and oviducts. In the 19 other operated animals a varying degree of regeneration of the interrupted nerve fibres occurred. In these animals a varying number of ripe eggs was present, but in many of them there existed a significant difference between the weight of their ovaries compared with that of the ovaries of the control animals. In all control animals the seasonal development of the ovaries and oviducts was normal. From these and from previous experiments it could be concluded that the A. F. negative nerve fibres of the median eminence, not originating from the magnocellular neurosecretory preoptic nuclei, are responsible for the normal seasonal development of the ovaries and oviducts in Rana temporaria. It must be these nerve fibres that release gonadotropic releasing factors into the blood capillaries of the median eminence. The A. F. positive neurosecretory fibres, the portal circulation of the pars distalis and the pituicytes of the median eminence play only a secondary part during this period of control of the gonadotropic activity of the hypophysis.     

The hypothalamo-hypophysial system of hypophysectomized rats

Summary The median eminence (ME) of hypophysectomized rats was studied by means of light and electron microscopy. Paraldehyde-fuchsin (PAF)-positive material is seen in the external zone (EZ) of the ME 2–5 days after the operation. Its amount gradually increases especially in the caudal part of the ME during the following few days. Some PAF-positive fibers make contact with the subependymally located blood capillaries. In the most caudal region of the recessus infundibuli they penetrate into the third ventricle. PAF-positive material decreases markedly from the ME of rats two months after hypophysectomy and exposure to a 1% salt load. Fibers of types A₁, A₂ and B containing granules of 120–220 nm, 100–150 nm and 80–100 nm in diameter, respectively, are seen in the EZ of the ME in hypophysectomized rats, although almost exclusively A₂- and B-type structures make contact with the primary portal capillaries in intact animals. All types of neurosecretory fibers establish contact with the subependymal nonfenestrated blood capillaries and penetrate the recessus infundibuli. Some neurosecretory terminals of different types make direct contact with the glandular cells of the pars tuberalis or are separated from them by a thin basal lamina. It is assumed that mainly neurosecretory fibers of types A₂ and B are permanently connected with the primary portal capillaries in the EZ of the ME in intact mammals, while the overwhelming majority of fibers of A₁-type shows ingrowth during the course of postoperative reparation. The possible physiological significance of the described changes is discussed.     

Mitotic Figures in the Median Eminence of the Hypothalamus

The median eminence of the hypothalamus is part of the avenue by which neurosecreted hormones from the hypothalamic nuclei reach the pars nervosa (neural lobe) of the pituitary and eventually the bloodstream. Lithium treatment and osmotic stress increases the transport of neurosecretory hormones to the pituitary in the adult rat. Specialized astrocytes termed pituicytes in the pars nervosa of the pituitary participate in the secretory process and also develop considerable mitotic activity. The present work reveals similar mitotic figures in cells within the median eminence following 3 days of lithium treatment. The location and appearance of these mitoses add to the evidence that pituicytes are present in the median eminence. Moreover, mitoses occur within the ependymal (tanycyte) layer of the median eminence. Thus, the present results suggest that the tanycyte layer may contain pituicytes, indicating that the hypothalamus possesses specialized cells for modulating neurosecretion in response to osmotic challenges.     

A gradient of activity in the hypophysis of Rana temporaria

Summary In normal pars distalis of Rana temporaria, a gradient of activity of the P.A.S.-positive cells is always present. In animals, with complete and permanent interruption of all nervous pathways to the median eminence and hypophysis and without disturbance of the normal blood supply of the hypophysis, a subtotal disappearance of the gradient of activity occurred and the total activity of the P.A.S.-cells of the pars distalis was lowered. The activity of the P.A.S.-positive cells regressed in a typical way, suggesting a humoral mechanism.In animals with marked regeneration of the interrupted nerve fibres, a normal gradient of activity was present. From these and previous experiments, it is concluded that the gradient of activity of the P.A.S.-positive cells of the pars distalis of Rana temporaria is largely dependent on the A.F.-negative fibres of the median eminence, not originating from the magnocellular neurosecretory preoptic nuclei. It is suggested that these nerve fibres exert their influence on the activity of the P.A.S.-positive cells via substances released in the blood capillaries of the median eminence.The gradient of activity and its variations are mainly related to the gonadotropic function of the pars distalis.I am indebted to miss M. T. Bakeeff and miss C. Everaert who were so kind to check the karyometric results.     

The structure and vascularization of the pars tuberalis of the hypophysis of Rana temporaria

Summary The pars tuberalis of the hypophysis of Rana temporaria presents the general structural and the cytological characteristics of an endocrine gland. It is composed of elongated cells with long, branching processes ending on the external basement membrane of the pericapillary space. The pars tuberalis cells produce secretory granules which are accumulated in the pericapillary endings of the processes.Corresponding to its separate localization, the pars tuberalis of Rana temporaria has a separate vascularization of which the efferent capillaries anastomose with the capillary plexus of the median eminence. The general direction of the blood flow of the pars tuberalis is towards the capillaries of the median eminence. Also, the secretory products of the pars tuberalis pass into the blood stream of the hypophysial portal system.Several characteristics of the pars tuberalis show that its function must be different from that of the pars distalis of the hypophysis. Moreover, in contrast with the pars distalis, the activity of the pars tuberalis is not regulated by neurohumoral factors.The results show that a role of the pars tuberalis in the regulation of the activity of the pars distalis of the hypophysis is not excluded.     

The function of the hypophysis without preoptic neurosecretory control

Summary In a large number of adult female and male specimens of Rana temporaria, a total extirpation of the magnocellular neurosecretory preoptic nuclei was performed. Several months after operation, no regeneration of the neurosecretory nuclei had occurred. The results showed 1. that all A.F.-positive neurosecretory nerve fibres of the hypothalamo-hypophysial region originate from nerve cell bodies localized in the magnocellular preoptic nuclei; 2. that the magnocellular preoptic nuclei do not play an important part in the inhibitory control of the activity of the pars intermedia of Rana temporaria; 3. that the possible role of the magnocellular preoptic nuclei in the seasonal development of the gonads and of the secondary sexual characteristics of Rana, could be only of secondary importance; 4. that, in accordance with previous experiments, in Rana, ovulation is dependent on preoptic hypothalamic structures. Some results of the present and of other experiments can be interpreted as supporting the hypothesis of a possible dual higher control over the gonadotropic centre (Dierickx, 1965b, 1966, 1967) of the pars ventralis of the tuber cinereum.A part of these studies has been reported previously in abstract form (Dierickx, 1963a, 1963b, 1965a).     

The hypothalamo-hypophysial system in Acipenseridae

Summary The proximal neurosecretory contact region (PCR) of Acipenseridae, a homologue of the tetrapod median eminence, has been studied by light, fluorescence and electron microscopy. It occupies the rostral and chiefly the ventral surfaces of the hypothalamic tuber cinereum. PAF-positive fibres occur in the zone of the preoptico-hypophysial tract but their terminal enlargements are concentrated mainly in the external zone. They make contact with the primary portal capillaries situated in the pia mater. Monoaminergic fibres and terminals with an intense green fluorescence are localized in the same regions. The fibres of some bipolar monoaminergic neurons of the PCR make contact both with the third ventricle and the primary portal capillaries.Three types of granule-containing neurosecretory fibres and terminals have been recognized in the PCR. Fibres of types A₁ (d=120–300 nm) and A₂ (d=100–170 nm) are peptidergic PAF-positive, although some fibres, including some of type A₁; belong possibly to PAF-negative type. Monoaminergic type B fibres have granules 80–100 nm in diameter. Neurosecretory terminals and vascular endfeet of tanycytes make contact with the 70 nm thick outer basement membrane of the primary portal capillaries. Several laminae of thin horizontally oriented tanycyte processes form a boundary between the external zone and the preoptico-hypophysial tract. Few neuroglial cells with pale cytoplasm, numerous lysosomes and lipofuscin granules are seen in this region. It is hypothesized that, as in other vertebrates, both peptide hypophysiotropic neurohormones and monoamines are discharged from the PCR into the portal circulation and affect the activity of the glandular cells of the pars distalis.Dedicated to Prof. Dr. Drs. h.c. W. Bargmann on the occasion of his 70th birthday     

Monoamine oxidase in the hypothalamo-hypophysial region of the teleosts,Anguilla japonica and Oryzias latipes

Summary Distribution of monoamine oxidase (MAO) was histochemically examined in the hypothalamo-hypophysial region of the eel (Anguilla japonica) and the medaka (Oryzias latipes) with a modified Glenner's tryptamine-tetrazolium method. The hypothalamic neurosecretory cells showed very weak MAO activity in their perikarya. MAO-positive fibers were present in close contact with the neurosecretory cells, suggesting that monoaminergic fibers participate in the control of neurosecretory cell activity. The nucleus lateralis tuberis (NLT) contained cells exhibiting strong MAO activity. These cells must be monoaminergic neurons.In the anterior region of the neurohypophysis of both eel and medaka, two bundles of MAO-positive fibers originating from the NLT proceed down along each side of the third ventricle into the pars distalis. This suggests that monoaminergic neurons of the NLT are involved in the release of hormones from the pars distalis. In addition to these tracts, numerous MAO-positive fibers proceed backward from the post-optic area and end around the blood capillaries located between the neurohypophysis and the pars intermedia in both species.I wish to express my gratitude to Prof. H. Kobayashi for his valuable advice during the course of this study. I am indebted to Prof. S. Uchida, Ocean Research Institute, University of Tokyo, for supplying the eels.     

The hypothalamo-hypophysial system in the ground squirrels, Citellus erythrogenys Brandt and Citellus undulatus Pallas. I. Microanatomy and cytomorphology of the gomori-positive neurosecretory system with special reference to its state during hibernation.

Microanatomy of the peptidergic hypothalamo-hypophysial neurosecretory system in the ground squirrels, C. erythrogenys and C. undulatus, is similar to that of other rodents, but differs in details. Bundles of neurosecretory fibres with single neurosecretory cells along their course connect the hypothalamic neurosecretory centers into the one system. The dorso-caudal portions of the NPV reach the level of the columna fornix. The NPO is a special indepedent formation and appears to have some specific functions in the HHNS. The infundibular recess penetrates deeply ihto the hypophysial stem, but does not reach the posterior pituitary. The glandular epithelium of the tuberal portions of the adenohypophysis surrounds the median eminence and a considerable part of the lateral surface of the tuber cinereum. Different functional types of neurosecretory cells reflect certain phases of their secretory cycle. Correlations between their size, structure and the content of NSM are found. A low activity of the HHNS is observed in both C. erythrogenys and C. undulatus during December-January. A higher activity of the HHNS in torpid C. undulatus than in C. erythrogenys is established. This correlates with a difference in deepness of torpor in C. erythrogenys and C. undulatus. An important role of the HHNS in hibernation, i.e. in realization of adaption of species in ontogenesis is suggested.     

Morphology and function of capillary networks in subregions of the rat tuber cinereum

Summary The differentiated cytology, cytochemistry, and functions within subdivisions of the tuber cinereum prompted this morphometric and physiological investigation of capillaries in the median eminence and arcuate nucleus of albino rats. Morphometric studies established that the external zone of the median eminence had 3–5 times the number and surface area of true and sinusoidal capillaries than the internal or subependymal median eminence zones, or either of two subdivisions examined in the arcuate nucleus. Type-I true capillaries, around which Virchow-Robin spaces comprise 1% of arcuate tissue area, were situated proximally to the median eminence border. This finding is consistent with a premise that confluent pericapillary spaces enable infiltration of arcuate neurons by factors from capillary blood from the median eminence or Virchow-Robin spaces. Physiologically, the rate of penetration across the median eminence capillaries by blood-borne [¹⁴C]-aminoisobutyric acid (a neutral amino acid used as a capillary permeability tracer) was 142 times greater than for capillaries in the distal arcuate nucleus within 12 s of tracer administration. A new finding was that the proximal arcuate nucleus had a permeability x surface area product of 69 l g^–1 min^–1, 34 times greater than that in more distal aspects of the tuber where blood-brain barrier properties exist. We also found that the microcirculatory transit time of a plasma space marker, [¹⁴C]sucrose, was considerably longer (1.2 s) in the median eminence and proximal arcuate nucleus than in the distal arcuate or ventromedial nucleus (0.4 s). By virtue of its high capillary permeability and extensive blood-tissue surface area, including the wide Virchow-Robin spaces, the median eminence external zone could be a gateway for flooding other tuberal compartments with blood-borne factors. This effect may be compounded by capillary bed specializations in the proximal arcuate nucleus where Type-I true capillaries, Type-III sinusoids, and pericapillary spaces are confluent with those in the median eminence. The results indicate that the proximal arcuate parenchyma could be exposed to circulating neuroactive substances on a moment-to-moment basis.Dedicated to Dr. Milton W. Brightman of Bethesda, Maryland, USA on the occasion of his 67th birthday and tribute as Craigie scholar at the First Craigie Conference on Brain Capillaries, Toronto, Ontario, June 24, 1990     

Immunocytochemical evidence for α-melanocyte-stimulating hormone in the hypothalamus of the frog Rana esculenta

Summary The distribution of immunoreactive -melanocyte-stimulating hormone (-MSH) within the brain of the frog, Rana esculenta, has been studied on adjacent serial sections using an indirect immunofluorescence technique. Immunoreactive cell bodies are found in the anterior part of the preoptic nucleus and in some ventral subependymal cerebrospinal fluid-contacting elements, and in the nucleus infundibularis ventralis. Numerous -MSH-like immunoreactive fibers are present in the preoptic area, in the pars ventralis of the tuber cinereum, and in the outer layer of the median eminence. This staining pattern is completely eliminated after preabsorbing the antiserum with the corresponding antigen, but blocking tests with -MSH-related peptides do not lead to any change in the immunoreaction. From these results it may be inferred that an -MSH-like system is present in the hypothalamic neurosecretory area of R. esculenta, and is probably related to its hypophysiotropic functions.The results are compared to the distribution of -MSH within the hypothalamus of reptiles and mammals.This work was supported by a grant from the M.P.I. (60%)