SEASONAL CHANGES IN THE ADRENAL STEROID TISSUE OF THE COMMON EIDER SOMATERIA MOLLISSIMA AND ITS RELATION TO ORGANIC METABOLISM IN NORMAL AND OIL-POLLUTED BIRDS

The annual cycle of the interrenal (steroid secreting) tissue of the adrenal gland of the Common Eider is described in terms of its fractional volume and mean nuclear diameter. These measurements indicate a close relationship between interrenal activity and the organic metabolism of the bird throughout the year. High interrenal activity is generally accompanied by utilization of energy reserves, for example during incubation and by oil-polluted birds, and low activity is associated with the replenishment of body tissues. Heavy fat deposition in April, before egg-laying, occurs at a time of increasing interrenal activity which apparently induces hyperphagia which in turn prevents the catabolic effects of high glucocorticoid production. The significance of the annual interrenal cycle in relation to breeding and moulting is discussed for the Eider and compared with a variety of other species.     

A light- and electron-microscopic investigation of gametogenesis in Typosyllis pulchra (Berkeley and Berkeley) (Polychaeta: Syllidae)

Summary The ultrastructure of the interrenal (adrenocortical) cells of trout (Salmo fario L.) was studied after dexamethasone treatment. A procedure for identifying and isolating interrenal tissue fragments from the surrounding head kidney tissue prior to their preparation for electron microscopy is described. The peripheral plasma cortisol concentrations were measured in order to evaluate the steroidogenic activity of this tissue.The interrenal cells of control animals contain numerous mitochondria with tubular cristae, and a well developed and highly organized smooth endoplasmic reticulum (SER). The scarcity, or absence, of lipid droplets contrasts markedly with the abundance of SER.Treatment with dexamethasone results in a decrease steroidogenic activity of the interrenal cells, as indicated by the fall in plasma cortisol concentrations. The interrenal cells are small, but still contain numerous mitochondria. The SER is poorly developed, but masses of densely intermeshed smooth cisternae subsist. Lipid droplets do not accumulate in these cells; this peculiarity is discussed in connection with the virtual absence of liposomes in teleost interrenal cells.     

Functional morphology of the interrenal and chromaffin cells in the teleosts, Rasbora daniconius (Hamilton), Barbus stigma (Cuv. et Val.) and Channa gachua (Hamilton).

The interrenal cells in Rasbora daniconius, Barbus stigma and Channa gachua are mainly found around the postcardinal vein and its major branches in the haemopoietic head-kidney. The chromaffin cells which are identified by the positive chromaffin reaction are found in the walls of the postcardinal vein or dispersed among the interrenal cells. delta5-3beta-HSDH and G-6-PDH activity was observed in the interrenal cells of all three teleosts. The present work indicates that the interrenal cells are capable of steroid biosynthesis and the chromaffin cells contain biologically active catecholamines.     

Gonadotropins and sex hormones modulate interrenal function in soft-shelled turtle

The aim of the current work was to investigate the role of gonadotropins and female sex hormones on interrenal activity in soft-shelled turtles, Lissemys punctata punctata. 1) FSH treatment (3 microg/100 g body wt daily for 10 days) caused interrenal hypertrophy with increased nuclear diameter, raises of acid phosphatase and alkaline phosphatase concentrations, and depletions of cholesterol (except the free fraction) and ascorbic acid levels from the interrenal gland. However, LH treatment (3 microg/100 g body wt daily for 10 days) failed to produce any perceptible change in the interrenal activity. The combined treatments of FSH and LH (3 microg each/100 gm body wt daily for 10 days) produced no further change beyond that of FSH alone. 2) Estrogen treatment with the low dose (25 microg/100 g body wt daily for 10 days) had no effect, but with higher doses (50 microg or 100 microg/100 gm body wt daily for 10 days) is caused interrenal stimulation by inducing the same manifestations to those of FSH. The degree of manifestations was higher with the higher dose (100 microg daily) than that of the moderate dose (50 microg daily). Progesterone treatment with the low dose (25 microg /100 g body wt daily for 10 days) had no significant effect, but with the moderate (50 microg daily) and higher (100 microg daily) doses suppressed interrenal activity by showing the reverse changes to those of estrogen. The degree of manifestations was higher with the higher dose than that of the moderate one. The combined treatments of estrogen and progesterone (100 microg each/100 g body wt daily for 10 days) caused interrenal stimulation but to a lesser extent than that of estrogen alone. The findings are briefly discussed.     

Morphofunctional research on the interrenal gland of the frog Rana temporaria following arginine vasotocin administration

Studies have been made of the effect of injections of hypothalamic nonapeptide neurohormone, arginine vasotocin, on functional condition of the interrenal gland in mature frogs. In unoperated, sham-operated and in animals 10 days after hypophysectomy, single and especially three subsequent injections of arginine vasotocin (5 x 10(-9) M per 1 kg of the body weight) result in evident activation of glandular cells of the interrenal gland which is manifested in the increase of the volume of their nuclei and cytoplasmic area, as well as in the dilatation of the blood vessels. Activation of the interrenal gland in hypophysectomized frogs, which lack endogenous ACTH, indicate the direct para-adenohypophyseal influences of nonapeptide hypothalamic hormones on the activity of glandular cells in the peripheral endocrine glands, in particular, the interrenal gland of the grass frog.     

Interrenal of a female gymnophione amphibian,Ichthyophis beddomei,during the annual reproductive cycle

The interrenal (adrenal) of Ichthyophis beddomei lies on the ventral side of the kidney, distributed in four zones. It is separated from the renal tissue by a thin layer of connective tissue and contains both adrenocortical and chromaffin cells. Adrenocortical tissue constitutes a major portion of the interrenal islets; the chromaffin tissue consists of a few cells located at the peripheries of the interrenal islets. Histochemical studies demonstrate the presence of Δ⁵3β-hydroxysteroid dehydrogenase, 17 β-hydroxysteroid dehydrogenase, glucose-6-phosphate dehydrogenase, succinate dehydrogenase, and sudanophilic lipids in the adrenocortical tissue, suggesting its steroidogenic potential. Annual histometric and histochemical studies show two peaks of interrenal activity: (1) during the breeding phase of the reproductive cycle (January and February) and (2) during the season of heavy monsoon rains (June and July) in the postbreeding phase.     

Social dominance and interrenal cell activity in rainbow trout,Salmo gairdneri (Pisces,Salmonidae)

Synopsis The relationship between standing in a dominance hierarchy and physiological stress was studied in rainbow trout. Individual fish were assigned relative dominance ranks, based on behavioral observations in a large, simulated stream tank. These ranks were compared to histometric measures of interrenal cell activity. Fish, isolated individually in the stream tank had significantly lower levels of interrenal activity than fish from the crowded holding tank. Groups of fish in the stream tank formed stable, linear dominance hierarchies. Interrenal activity correlated inversely with dominance rank, with the exception that top ranking fish had higher activity than expected. Possible cause and effect relationships are discussed.     

Comparative cytomorphology of the avian adrenocortical tissue.

The histology and histochemistry of the interrenal gland of twenty avian species have been described in this report. The avian interrenal tissue, on the basis of nuclear orientation and other cytomorphic features, can be classified into six cytologically distinct types. Cytomorphic organization within the interrenal tissue of these avian species also indicates some regional demarcation into subcapsular and central zones. Zonal demarcation in the interrenal tissue of these birds has also been indicated from chemocytological investigation. Paradoxically not much of correspondence was noticed in the cytologic and cytochemical patterns. The cytological and cytochemical studies fail to indicate any phylogenetic trend within the interrenal tissue of the birds under investigation.     

Interrenal responses to high ambient temperature in soft-shelled turtle, Lissemys punctata punctata

An exposure to ambient temperature of 25 degrees C had no perceptible effect on interrenal function but further increase of temperature to 35 degrees C caused nuclear hypertrophy with increase of nuclear diameter, RNA concentration, acid phosphatase and alkaline phosphatase activities, accompanied by quantitative depletions of cholesterol (free, esterified and total) and ascorbic acid levels in the interrenal gland of the soft-shelled turtle Lissemys p. punctata. Similar manifestations of stimulation, except in the nucleus, were marked after exposure to 38 degrees C, but the degree of response in respect of esterified and free cholesterol levels was higher at 38 degrees C than at 35 degrees C. Moreover, withdrawal of 38 degrees C temperature and subsequently maintaining at 25 degrees C for 15 days showed reverse manifestations to those of 35 degrees C/38 degrees C, leading to a tendency towards normalcy. It is suggested that high a ambient temperature of 35 degrees C significantly stimulates interrenal function of Lissemys turtles, but further increase of 38 degrees C does not cause further overall stimulation, and withdrawal of higher temperature (38 degrees C) shows a tendency towards normalcy. It is also suggested that (a) high ambient temperature causes thermal stress, (b) it is reversible and (c) it acts on interrenal activity presumably via CRF-ACTH-axis in turtles.     

Effect of ACTH and bilateral gonadectomy on the adrenocortical tissue of an air-breathing fish -- Heteropneustes fossilis (Bloch).

The mammalian ACTH (20 I.U.) injected in Heteropneustes fossilis (Bloch) caused hyperplasia of the adrenocortical tissue. The interrenal as well as chrome cells became vacuolated. The interrenal nuclear size increased by 15% as an average than the control fishes. The glycogen granules were found to be absent from the interrenal cells. The effect of bilateral gonadectomy in both sexes has been marked in the gradual decrease of the interrenal nuclear diameter which 12 weeks after the operation, recorded nearly 25% reduction in size. The glycogen content was reduced considerably. Removal of gonads during maturation period bringing inhibition of hyperplasia of the gland at the time of spawning possibly indicates action of gonadal steroids on the physiological alterations of the interrenal cells.     

The effect of injections of arginine vasotocin and mesotocin on the function of the interrenal gland in the frog Rana temporaria 2 months after the removal of the anterior lobe of the hypophysis]

Morphometric and biochemical studies have been made on the activity of interrenal gland in the frog Rana temporaria after removal of the anterior lobe of pituitary and injections of arginine vasotocin and mesotocin. It was demonstrated that the activity of this gland after the operation decreases increasing after arginine vasotocin injections.     

Histophysiologische Untersuchungen der Entwicklung des Interrenalorgans beim Krallenfrosch (Xenopus laevis Daudin)

Zusammenfassung Mit histologischen und histochemischen Methoden lassen sich bei Xenopus-Larven 4 Zonen des interrenalen Gewebes (Nebennieren-Rinden-Gewebes) in caudo-cranialer Ausdehnung unterscheiden, die verschiedenen Phasen der Nebennierenentwicklung entsprechen. Diese Zonen wurden auf Grund der in ihnen am häufigsten auftretenden Zelldifferenzierungsformen als Blastemzone, Zone der Organbildung, Zone der Ausdifferenzierung und Zone der Funktionsphasen bezeichnet. Während der Entwicklung verändert sich ihre Ausdehnung, sie verschieben sich von cranial nach caudal. Daneben kommt es während der Ausdifferenzierung zu einer Verlagerung der Interrenalzellen von dorsal nach ventral. Die ausdifferenzierten Interrenalzellen durchlaufen einen Funktionszyklus. Charakteristische Differenzierungformen sind hierbei: wenig stimulierte, stark stimulierte, überstimulierte Interrenalzellen sowie Zellen in der Restitutionsform und Degenerationsform.Kerngrößenmessungen und Bestimmungen der Steroiddehydrogenase-Aktivität an Normaltieren zeigen eine Abhängigkeit der Nebennierenaktivität von dem Entwicklungsstadium. Maxima der Aktivität liegen im Stadium 59 und am Ende der Metamorphose vor.Untersuchungen des Organes nach Hypophysektomie und/oder ACTH-Behandlung machen den Einfluß der Hypophyse auf die Normalentwicklung der Nebenniere deutlich. Hypophysektomie verhindert zwar nicht die Abgliederung der Nebenniereninseln vom Blastem, die Zellen differenzieren sich jedoch nicht vollständig aus, sondern verharren in einer funktionellen Ruhepause. ACTH-Behandlung beschleunigt die Differenzierung und funktionelle Entwicklung der Interrenalzellen und führt zu einer Vergrößerung der einzelnen Interrenalkomplexe. Sie kompensiert auch die hemmende Wirkung der Hypophysektomie auf die Nebenniere. Außerdem vermag ACTH die durch Hypophysektomie bewirkte Wachstums-und Entwicklungshemmung teilweise auszugleichen.

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Histophysiological investigations of the development of the interrenal organ in the clawed toad (Xenopus laevis Daudin)

Summary Histological and histochemical investigations have given evidence for 4 zones of the interrenal tissue in caudo-cranial direction in Xenopus-larvae. These zones correspond to 4 phases of the interrenal development. Due to the representative differentiation form of the interrenal cells they were characterized as: 1) zone of the blastema, 2) zone of the newly formed interrenal organ, 3) zone of differentiation and 4) zone of the various functional stages. During the development these zones shift from cranial to caudal. Besides there is a displacement during the differentiation of the interrenal cells in dorso-ventral direction. Differentiated interrenal cells undergo a functional cycle: hardly stimulated, clearly stimulated and overstimulated adrenocortical cells as well as cells in the state of restitution and degeneration are to be observed.Calculation of the mean nuclear diameter and the activity of the steroiddehydrogenase in normal animals demonstrate a correlation between the activity of the adrenal cortex and the stage of development. Maxima of this activity are found in stage 59 and at the end of metamorphosis. Experiments with hypophysectomy and/or ACTH-treatment clearly demonstrate the influence of the pituitary on the adrenocortex during normal development. Hypophysectomy does not prevent the separation of interrenal islets from the blastema. These cells, however, do not completely differentiate but stay in an inactive phase of their function. ACTH-treatment accelerates the differentiation and functional development of the interrenal cells and further results in an enlargement of the single adrenocortical islets. This treatment compensates the inhibition of the adrenal cortex after hypophysectomy and partly compensates the retardation of development and growth in hypophysectomized animals.

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Mit dankenswerter Unterstützung durch die Deutsche Forschungsgemeinschaft.     

Seasonal studies on the adrenal homologue of female Puntius sophore (Ham.).

In female P. sophore the adrenal homologue is located inside its head kidneys and comprises interrenal cells and chromaffin cells. The clumps of interrenal cells surround the postcardinal vein and its major branches; they also lie in the haemopoietic tissue. The chromaffin cells mostly lie in the endothelium of the post-cardinal vein, while a few of them also lie interspersed among the interrenal cells. The seasonal study of the adrenal homologue shows that the interrenal cells undergo changes in bulk and composition in different months, while the chromaffin cells do not show any significant variation.     

Parallel early development of zebrafish interrenal glands and pronephros: differential control by wt1 and ff1b

Steroids are synthesized mainly from the adrenal cortex. Adrenal deficiencies are often associated with problems related to its development, which is not fully understood. To better understand adrenocortical development, we studied zebrafish because of the ease of embryo manipulation. The adrenocortical equivalent in zebrafish is called the interrenal, because it is embedded in the kidney. We find that interrenal development parallels that of the embryonic kidney (pronephros). Primordial interrenal cells first appear as bilateral intermediate mesoderm expressing ff1b in a region ventral to the third somite. These cells then migrate toward the axial midline and fuse together. The pronephric primordia are wt1-expressing cells located next to the interrenal. They also migrate to the axial midline and fuse to become glomeruli at later developmental stages. Our gene knockdown experiments indicate that wt1 is required for its initial restricted expression in pronephric primordia, pronephric cell migration and fusion. wt1 also appears to be involved in interrenal development and ff1b expression. Similarly, ff1b is required for interrenal differentiation and activation of the differentiated gene, cyp11a1. Our results show that the zebrafish interrenal and pronephros are situated close together and go through parallel developmental processes but are governed by different signaling events.     

Correlated changes in the structure of the anterior pituitary gland,testes and interrenal tissue during sexual maturation of male lizards

Summary The anterior pituitary gland, testes and interrenal glands of a series of young males of the teiid Cnemidophorus l. lemniscatus (L.) have been studied by light microscopy in order to correlate the changes occurring during sexual maturation. In the testes of the smallest animals, spermatogenesis does not progress beyond primary spermatocytes and there is no differentiated interstitial tissue. In medium-sized animals, spermatids and some interstitial cells appear, and in the largest lizards, spermatogenesis is completely established and Leydig cells abound. Simultaneously with the development of the testes, interrenal glands undergo great hypertrophy and hyperplasia, especially in the peripheral reactive zone. Starting in animals of intermediate size, the anterior hypophysis exhibits a considerable hypertrophy of two rostral cell types: the chromophobic corticotrophs and the acidophilic PAS-positive cells considered to be interstitiotrophs. These cells show large, vesicular nuclei and prominent nucleoli, signs of enhanced cellular activity. The hypertrophy begins in the dorso-rostral region of the gland close to the median eminence, at the site of entry of the portal vessels. This suggests a hypothalamic influence on the function of these pars distalis cells. The scattered basophilic gonadotrophs or folliculotrophs are scarce, small, and do not vary appreciably among the animals studied. The hyperactivity of corticotrophs may account for enlargement of the interrenal glands. Testicular development is apparently related to an increased activity of interstitiotrophs but to a stable level of activity in folliculotrophs.This research forms part of project No. 31.26.S1-0244 supported by the Consejo Nacional de Investigaciones Científicas y Tecnológicas.     

Histophysiologic response of the pituitary-inertrenal system to salicylate and adrenocorticotropin in catfish, Heteropneustes iossilis (Bloch).

The interrenal and pituitary cells showed increase in nuclear diameter eight days after treatment with ACTH, salicylate or salicylate + ACTH. After ACTH treatment, however, the cytoplasm in the interrenal cells was granular. Similarly, the lead-haematoxylin positive (PbH + ve) cells in the rostral pars distalis also showed partial depletion of stainable granules after ACTH treatment. Salicylate or salicylate ACTH caused depletion of cytoplasmic material in both the interrenal as well as PbH + ve cells.     

Endothelium is required for the promotion of interrenal morphogenetic movement during early zebrafish development

The adrenal cortex has a complex vasculature that is essential for growth, tissue maintenance, and access of secreted steroids to the bloodstream. However, the interaction between vasculature and adrenal cortex during early organogenesis remains largely unclear. In this study, we focused on the zebrafish counterpart of adrenal cortex, interrenal tissue, to explore the possible role of endothelium in the development of steroidogenic tissues. The ontogeny of interrenal tissue was found to be tightly associated with the endothelial cells (ECs) that constitute the axial vessels. The early interrenal primordia emerge as two clusters of cells that migrate centrally and converge at the midline, whereas the central convergence was abrogated in the avascular cloche (clo) mutant. Neither loss of blood circulation nor perturbations of vessel assembly could account for the interrenal convergence defect, implying a role of endothelial signaling prior to the formation of axial blood vessels. Moreover, as the absence of trunk endothelium in clo mutant was rescued by the forced expression of SCL, the interrenal fusion defect could be alleviated. We thus conclude that endothelial signaling is involved in the morphogenetic movement of early interrenal tissue.     

The influence of social rank on adenohypophysial cell activity in Salmo irideus

Summary In nine cell types of the adenohypophysis in untreated adult rainbow trout, histologically different activity phases, seasonal changes in activity, and the relation between certain cell types and the interrenal gland, thyroid or gonads were investigated by light and, occasionally, by electron microscopy. Special attention was given to the effect of social rank on the synthetic activity in adenohypophysial cells of trout kept in small groups in which a social hierarchy with one (light) dominant and several (dark) submissives is established.Cell types in the rostral pars distalis were azocarminophil (I) or amphiphil (II). Proximal pars distalis cell types were slightly basophil (IV), orangeophil (V), strongly basophil (VI) or chromophobe (VII). In the pars intermedia, cell types were amphiphil (VIII) or very slightly basophil (IX). Type III was a non-secretory supporting (?) cell.Histologically different activity phases abounded in type IV cells, which mainly occurred in the proximal pars distalis but were also found dispersed in the rostral pars distalis, the pars intermedia and the neurohypophysis.Influences of social rank were pronounced in type IV cells. Phases with a high synthetic activity were exclusively found in submissive animals, phases with a low synthetic activity occurred in dominants. As a positive relation existed between type IV cell activity and the social rank dependent activity of the interrenal gland, it was suggested that type IV cells produce ACTH.In (dominant) male trout treated with DOCG or ACTH, colloid-containing type IV cell phases, reflecting accumulation of the secretory product, were found. This supported the earlier suggestion that ACTH in the trout is produced in the basophil type IV cells and not, as reported in the literature, in cells comparable to type II.The author is greatly indebted to Miss M.C. Wentzel, Miss I. Stulen, Mr. J. Veening and Dr. J.G. van Rhijn for their help with histological techniques, interrenal cell measurements and statistical aspects