RELATION BETWEEN OVARIAN MATURATION AND ACTIVITY OF THE CORPORA ALLATA IN SEASONAL FORMS OF THE BUTTERFLY, POLYGONIA C-AUREUM L.

In summer form adults of Polygonia c-aureum, the ovary develops soon after emergence and eggs mature a few days later. However, in autumn form adults, having been kept in laboratory conditions, only several mature eggs are found about four weeks after emergence. The allatectomy during the larval period prevents egg maturation in adults of both forms. When the corpora allata from the summer form of both sexes are transplanted into allatectomized adults of the autumn or summer form, the ovary develops as in the case of summer form adults. But the corpora allata of both sexes of the autumn form are not effective in inducing the ovarian maturation. Environmental factors, especially day-length and temperature during the larval period, may act through the neuroendocrine system to induce the activity of the corpora allata in the adult life.     

Système endocrine et physiologie de la diapause imaginale chez le criquet égyptien, Anacridium aegyptium

The Egyptian locust, Anacridium aegyptium, has four protocerebral neurosecretory centres: the A to B neurosecretory cells of the pars intercerebralis (the A cells are rich in fuchsinophil material and the B cells are devoid of fuchsinophil neurosecretion), the voluminous C neurosecretory cells poor in neurosecretion, and the median sub-ocellar neurosecretory cells.From September to the beginning of January, imaginal diapause is characterized by an accumulation of the median neurosecretion in the pars intercerebralis-corpora cardiaca system, by small corpora allata, and, in the female, by a stop in oöcyte development although the male's sexual activity is still not altered. Allatectomy suppresses neither the male's sexual behaviour nor its fecundity. From January, the increase of the photoperiod causes a release of the median neurosecretion in both sexes, an increase of the volume of the corpora allata, and breaks ovarian diapause.In autumn, the implantation of the male's or female's corpora allata of Anacridium does not stimulate ovarian growth of diapausing females. On the contrary, the implantation of corpora allata or of pars intercerebralis or of corpora cardiaca of Locusta migratoria migratorioides (locust without diapause) causes ovarian development of the diapausing females of Anacridium. Thus, in the two sexes of the Egyptian locust, the corpora allata are inactive during the female ovarian diapause. The imaginal diapause of Anacridium affects both sexes (stocking of median neurosecretion, arrest of the corpora allata). If diapause does not seem to affect the male's development, it is because its sexual activity is free from the pars intercerebralis and corpora allata.The corpora allata of Anacridium show a sexual dimorphism in the active adult: they are smaller in the male and have more mitosis in the female. An explanation of this dimorphism is advanced.     

Rupture de la diapause ovarienne d'Anacridium aegyptium par stimulation électrique des cellules neurosécrétrices médianes de la pars intercerebralis

Oögenesis and the physiological activity of the corpora allata were studied in adult females of the Egyptian locust (Anacridium aegyptium), in ovarian diapause, after electrical stimulation in vivo of the pars intercerebralis. This stimulation provokes (1) a decrease in the quantity of fuchsinophilic material present in the median neurosecretory cell bodies and in the internal cardiac tract, (2) an increase in the physiological activity of the corpora allata (measured by its chromatropic effect on larvae of Locusta), and (3) rupture of the ovarian diapause (advance of maturation of the oöcytes and oviposition by 5 months, and initiation of the ovarian cycle).In the control animals, the same electrical stimulations of various regions of the central nervous system (tritocerebrum, first ganglion of the abdominal cord) have no effect on these phenomena.In allatectomized females, electrical stimulations of the pars intercerebralis are followed by a slight growth of oöcytes, without a deposit of yellow vitellus. The diapause is not broken. Section of the allatocardiac nerves or rupture of the allatocardiac and allato-suboesophageal nervous connexions do not change the physiological state of the corpora allata. In the case of females in which the corpora allata have been disconnected, electrical stimulations of the pars intercerebralis succeed in activating the corpora allata and breaking the ovarian diapause. The aggregate of these results confirms that in locusts the control of the brain over the physiological activity of the corpora allata is above all neuroendocrine.     

Autogeny and endocrine involvement in the reproduction of tropical flesh flies (Diptera: Sarcophagidae)

Autogeny is highly developed among the Sarcophagidae in tropical Africa. Unlike their temperate region relatives, the tropical species examined can mature the first batch of eggs without an adult protein meal, and they do so without the long period of delay usually seen in autogenous species. Protein deprivation, however, does reduce the number of eggs matured by about one-third. The tropical and temperate flies appear to share a common endocrine mechanism controlling egg maturation. Removal of the medial neurosecretory cells of the pars intercerebralis prevents egg maturation. Reimplantation of clusters of MNC restores the ability to mature eggs. Extirpation of the corpora allata reduces the percentage of females that mature eggs to about 70%, and the females that do mature eggs produce fewer eggs. The significant difference between the tropical and temperate species centers on the trigger that sets the neuroendocrine system into action. For the tropical species, the brain is activated immediately and does not require the trigger of a protein meal.     

Experimental evidence for a neuroendocrine control of ecdysone biosynthesis in adult females of Locusta migratoria

The considerable increase in ecdysteroid concentration which occurs in normal Locusta ovaries at the end of each cycle of oöcyte maturation is prevented if the median neurosecretory cells of the pars intercerebralis are cauterized, or if the corpora cardiaca are excised 24 hr before the onset of ecdysone synthesis in normal females. Implantation of additional brain-corpora cardiaca complexes into young vitellogenic females advances the time of ecdysone synthesis by some 12 hr. Oöcyte growth itself is not affected in these different types of experiments.It is inferred from the data of the present study that ecdysone synthesis in the follicle cell epithelium of maturing Locusta ovaries is stimulated by a neurohormone produced in the median neurosecretory cells of the pars intercerebralis and secreted into the blood via the corpora cardiaca.     

Untersuchungen über den einfluss der königinnensubstanz auf die entwicklung der endokrinen drüsen bei der arbeiterin der honigbiene (Apis mellifica)

Zusammenfassung Die Corpora allata der Bienearbeiterinnen machen während des Adultlebens zwei Wachstumsphasen durch. Im Laufe der ersten Phase wird das Wachstum durch Königinnensubstanz gehemmt; die zweite dagegen tritt nur bei Anwesenheit einer Königin auf. Das Wachstum der Corpora allata weiselloser Bienen ist mit der Eireifung korreliert. Ein Hormon der Corpora allata dürfte für die Eireifung notwending sein (Abb. 4).Ausser dem Hormon der Corpora allata muss für das Oocytenwachstum ein weiterer Faktor vorhanden sein. Wahrscheinlich handelt es sich um ein Sekret der neurosekretorischen Zellen der Pars intercerebralis des Gehirns. Die Königin hemmt direkt oder indirekt das Kernwachstum in den neurosekretorischen Zellen der Arbeiterin. Es ist fraglich, ob es sich auch hier um eine Wirklung der Königinnensubstanz handelt.

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Summary The corpora allata of adult worker bees show growth periods. During the first period growth is inhibited by queen substance while the second growth period only occurs when a queen is present. A correlation between the volume of the corpora allata and oocyte suggests that oocyte maturation is dependent on the presence of a hormone of the corpora allata.Besides of the hormone of the corpora allata at least one other factor is necessary for oocyte maturation. It is probably contained in the neurosecretory material produced in the pars intercerebralis of the brain. The growth of the nuclei of the neurosecretory cells of the brain is inhibited by the presence of a queen, but it is doubtful whether this is due to the action of queen substance.

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Résumé Les corpora allata des ouvrières adultes d'abeilles ont deux phases de croissance. Lors de la première phase, la croissance est inhibée par la phéromone de la reine, tandis que la deuxième phase ne se produit qu'en présence d'une reine. Une corrélation entre le volume des corpora allata et la croissance des oocytes suggère que l'oogenèse dépend d'une hormone des corpora allata.Or, un autre facteur au moins est encore nécessaire pour la maturation des oocytes. Il est probablement sécrété par les cellules de la pars intercerebralis du cerveau. La croissance des noyaux des cellules neurosécrétrices du cerveau est inhibée par la reine, mais le mode d'action de la phéromone dans cette inhibition reste inconnu.

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Durchgeführt mit Hilfe eines Forschungskredites des Schweiz. Nationalfonds an Herrn Prof. Dr.M. Lüscher.     

Intervention des cellules neurosecretrices medianes dans la castration parasitaire d'Anacridium aegyptium (Orthoptère)

The median neurosecretory cells (M-NSC) of Anacridium aegyptium were studied by light and electron microscopy during the ovarian diapause (autumn and winter) and the ovarian activity (spring) in normal females and in females infected by a tachinid fly Metacemyia calloti. In spring, the M-NSC contain comparatively much stainable material in normal females than in parasitized ones. This difference results from the ovarian maturity of normal females and the ovarian immaturity of parasitized females.The activity of the M-NSC was assessed in spring using the evolution of the radioactivity of the pars intercerebralis (PI) and corpora cardiaca (CC) after injection of ³⁵S-cysteine. The activity of the M-NSC is impaired in the parasitized females without ovarian development.The ovarian diapause was broken by massive implantations of corpora allata (CA) or electrical stimulations of the PI. Implantations of CA induce in parasitized females the same state of ovarian development than during natural activity in spring. Electrical stimulations of the PI. which increase the activity of the M-NSC. enhance the ovarian development of the parasitized females. A hypoactivity of the M-NSC is involved in the ‘castration’ of the Egyptian grasshopper.     

Neuroendocrine regulation of corpus allatum activity in Manduca sexta: The endocrine basis for starvation-induced supernumerary larval moult

When newly-ecdysed 5th instar larvae of Manduca sexta were starved for 3 days and thereafter fed on standard diet the majority (90%) of the surviving larvae moulted into 6th instars. Allatectomy prior to starvation abolished the supernumerary moult, while denervation of the corpora allata (CA) had no effect.Cautery of medial neurosecretory cells, but not of the lateral cells, prevented supernumerary moulting and pupation ensued. Transplantation of brains from young 5th instar donors into larvae, whose medial neurosecretory cells were cauterized prior to starvation, restored the extra larval moult. Neither CA nor corpora cardiaca (CC) could be substituted for the medial neurosecretory cells.For induction of the supernumerary moult the medial neurosecretory cells are required only until day 1 after refeeding whereas the CA are required until day 3 after refeeding. Allatectomy on day 3 after refeeding resulted in the production of black 6th instar larvae.We conclude that starvation-induced supernumerary moulting is due to activation of the CA by allatotropin produced by medial neurosecretory cells in the brain. The anteromedial cells (group II) appear to be the source of allatotropin.     

Daily protein synthesis in the pars intercerebralis and corpora allata ofApis mellifica L. with and without training to a foraging schedule

Summary The rate of protein synthesis by the neurosecretory cells in the pars intercerebralis and in the corpora allata was determined by autoradiography. Bees that forage throughout the day display a daily rhythm in that rate; the synthesis curve could be shifted considerably in each of two different training experiments.The total amount of protein synthesized per day is distinctly greater for bees that collect all day than for those with activity limited by the training schedule.The corpora allata also exhibit a daily rhythm in protein synthesis, and this too can be altered by training. The relationship to collecting activity are less clear than in the case of the pars intercerebralis, but are similar in direction of shift.We thank Mrs. L. v. Szczepanski for valuable assistance in preparing the autoradiographs; funds and equipment for their preparation were supplied by the Gesellschaft für Strahlen- und Umweltforschung m.b.H. Neuherberg/München (Institute of Biology: Prof. O. Hug).     

ENDOCRINOLOGICAL STUDIES ON SPERMATOGENESIS IN THE SILKWORM, BOMBYX MORI L.

When embryonic testes 1 or 2 days before hatching were transplanted into 5 kinds of hosts (5th instar larvae, “fresh” pupae just after pupation, isolated pupal abdomens and 3-day-old pupae with and without their original corpora allata), the testes transplanted into 5th instar larvae grew most conspicuously and spermiogenesis began in many cysts. A few spermatidal cysts were observed in the testes transplanted into fresh pupae. No signs of maturation were observed in the testes transplanted into other hosts. It is concluded that prothoracic gland hormone might be responsible for the precocious maturation of young testes. On the other hand, the initiation of spermiogenesis was delayed, in comparing with controls, in the 3rd instar larval testes as follows: the testes transplanted into 3-day-old pupae, the testes transplanted into isolated pupal abdomens together with adult corpora allata and the testis into which corpus allatum of a 3rd instar larva was inserted. It is concluded that corpora allata hormone exerts an inhibitory effect on spermatogenesis.     

Controle neuroendocrine du fonctionnement du corpus allatum chez Rhodnius prolixus

Cauterization of the dorsal portion of the protocerebrum effected immediately after cessation of feeding in adult females blocks vitellogenesis. The growth of the oöcytes can be re-established by the implantation of corpora allata or by the application of juvenile hormone analogue, and it is suggested that the protocerebrum has a corpus allatum stimulating influence. The corpus allatum deprived of all its cerebral nervous connexions by cutting the aorta just behind the corpora cardiaca induces abundant vitellogenesis. Females with denervated corpora allata or pars intercerebralis cauterized 24 hr after a meal show a phase of vitellogenesis more active than that in normal female adults. This suggests the existence of a cerebral inhibitory centre: its activity through a nervous or endocrine path is discussed.     

Developmental Changes in the Bombyxin-and Insulin-like Immunoreactive Neurosecretory System in the Wax Moth, Galleria mellonella

Four medial neurosecretory cells (MNC) and 4 lateral neurosecretory cells (LNC) in each brain hemisphere, and one pair of cells in each thoracic ganglion (TG) of Galleria larva react with antibodies against bombyxin and insulin. Material secreted from the MNC and LNC is released mainly in the corpora allata, and that from the TG through the ventral median nerves. Intrinsic secretory cells of the corpora cardiaca (CC) also contain bombyxin-like, but not insulin-like material. The immunoreactivities all disappear during molts and reappear with resumption of feeding. In the MNC and TG they reappear for less than a day, but in cells of the CC immunoreactivity reappears for the whole feeding period. Before pupation, the LNC become temporarily immunopositive towards the end of feeding period, and the MNC and TG during the wandering period, i.e. at the time of prothoracic gland stimulation. Immunoreactivity disappears during the pupal molt. In pupae it is present in the 4 pairs of MNC and 1–2 pairs of LNC 12–48 hr after ecdysis, and in cells of the CC from 12 hr after ecdysis until the end of the pupal instar. In adult, immunoreactivity is restricted to 2 pairs of the LNC and to CC cells.     

NADPH diaphorase and nitric oxide synthase in the corpus cardiacum-corpus allatum of the cockroach Diploptera punctata

Juvenile hormone synthesis by corpora allata is regulated partly by allatostatin containing nerves from the brain that innervate the corpora cardiaca and the corpora allata. To investigate whether NO also participates in the regulation of juvenile hormone synthesis, antibody against NO synthase and the histochemical test for NADPH diaphorase activity, a marker for NO synthase, were applied to the corpora cardiaca-corpora allata of Diploptera punctata. Strong NADPH diaphorase activity occurred in corpus allatum cells but not in nerve fibers in the corpora allata or corpora cardiaca. In contrast, NO immunoreactivity occurred in nerves in the corpora cardiaca but not within the corpora allata. NO and allatostatin were not colocalized. NO synthase and NADPH diaphorase activity were localized in similar areas of the subesophageal ganglion and cells in the pars intercerebralis of the brain. Positive correlation of the quantity of NADPH diaphorase activity with juvenile hormone synthesis during the gonadotrophic cycle and lack of such correlation in subesophageal ganglia suggest that NADPH diaphorase activity reflects the necessity of NADPH in the pathway of juvenile hormone synthesis. These data suggest that NO is unlikely to play a significant role in the regulation of the corpora allata.     

The influence of some chemosterilants and caffeine on the neuroendocrine system of adult male Ephestia kühniella

Injection of caffeine into adult male Ephestia kühniella induced a reduction in the amount of neurosecretory material in the resorcin fuchsin stainable (A-) cells of the pars intercerebralis. Both hemel (hexamethyl melamine) and tepa (tris (1-aziridinyl) phosphine oxide) treatments resulted in an increase in stainable neurosecretory products in the cytoplasm of the A-cells. The increase after hemel treatment was largely a result of increased synthesis of neurosecretory material as indicated by increased nuclear volume and nuclear-cytoplasmic ratio. No changes were induced by hemel in the corpora cardiaca and allata of the adult male E.kühniella. This disruption in neuroendocrine function is believed to be involved in the observed mating aberrations (permanent copulation and sterile matings) by adult males after hemel treatment.     

Evidence for a crustacean hyperglycemic hormone-like molecule in the nervous system of the stick insect,Carausius morosus

Summary Histological sections of the brain, suboesophageal ganglion, and the corpora cardiaca/corpora allata complex were examined for the presence of crustacean hyperglycemic hormone-like immunoreactive material. With the use of an antiserum directed against the hyperglycemic hormone of Carcinus maenas, immunofluorescence was found in the median portion of the pars intercerebralis, and the corpora cardiaca. Extracts of corpora cardiaca were examined by radioimmunoassay for competitive binding to the antiserum; one pair of corpora cardiaca contains at least 7 pg crustacean hyperglycemic hormone-like material.     

Interaction of the neuroendocrine system and ovaries in sheep blowflies, Lucilia sericata, sterilized with tepa

ABSTRACT. Female Lucilia sericata Mg. fed a reproductive diet of meat were sterilized by injection of 10μg of tepa. Light microscopy revealed that the medial neurosecretory cells of the brain of sterilized flies were highly active compared with the same cells in untreated flies of similar age and nutritional status. The corpora allata of sterilized flies were also greatly enlarged. The evidence suggests that developing oocytes secrete a substance that inhibits the corpora allata hormone, thereby suppressing brain neurosecretory cell activity and preventing oogenesis in the subsequent oocytes.     

Neurosecretory system of the earwig Labedura riparia, and the rôle of the aorta as a neurohaemal organ

The neurosecretory system of Labedura riparia has been described from sections and whole mounts using a variety of techniques. The pars intercerebralis contains two clusters of medial neurosecretory cells (MNC), each cluster consisting of 8 to 10 A-cells and occasional B-cells. The lateral sides of the brain have a few B-cells. The axons of the median neurosecretory cells terminate in the cephalic aorta (AO), whereas the axons of the lateral neurosecretory cells (LNC) terminate in the corpora cardiaca (CC). It appears that the neurosecretory material (NSM) elaborated in the MNC is stored in the cephalic aorta and that elaborated in the LNC is stored in the corpora cardiaca, which are two oval or elongate bodies composed of large chromophobe and small chromophil cells. Posteriorly there is the oval or elongate corpus allatum (CA) attached to the CC by thick nerves. The CA consists of one cell type only. Both CC and CA contain no A-cell neurosecretory material. It has been suggested that the neurosecretory system of L. riparia is composed of two complexes. One is formed by the medial neurosecretory cells for which the aorta functions as a neurohaemal organ, and the other is formed by lateral neurosecretory cells-lateral neurosecretory pathways-nervi corporis cardiaci-II in which the corpora cardiaca function as a neurohaemal organ.     

Precocene sensitivity of corpora allata in adult female Locusta migratoria after electrocoagulation of the pars intercerebralis neurosecretory cells

Destruction, by electrocoagulation, of the median neurosecretory cells of the pars intercerebralis of 2-day old adult female Locusta migratoria completely suppressed normal juvenile hormone-biosynthetic activity of the corpora allata in most animals. For example, 6 days after electrocoagulation the rates of spontaneous juvenile hormone biosynthesis, measured by radiochemical assay of freshly isolated glands, showed a median value of less than $\text{1}\text{100}$ that of the corresponding sham operated controls, which were then in mid-vitellogenesis. Injection of graded doses (200–1600 μg) of precocene I at this time, followed by assays five days later, resulted in a similar decline of both corpus allatum volume and precocene epoxidase activity (measured by radiochemical assays of precocene I dihydrodiol formation in vitro) in pars intercerebralis-coagulated and sham-operated animals, when expressed as a percentage of their own zero precocene controls. Electrocoagulation of the pars intercerebralis largely prevented the normal increase in both corpus allatum volume and its epoxidase activity, so that by age 13 days these parameters were about 2.5-fold lower in coagulated vs control (sham operated) animals. In fact, electrocoagulation had no marked effect on the value of epoxidase activity per unit corpus allatum volume. It is concluded that the corpora allata from this species and stage are sensitive to precocene irrespective of whether they are active in biosynthesis of juvenile hormone.     

Neuroendocrine control of vitellogenesis in the dragonfly, Orthetrum chrysis (Selys) (Odonata: Libellulidae).

1. Histological correlative studies on the cerebral neurosecretory cells, the corpora allata and the oocyte development strongly suggest the involvement of cerebral NSM mostly produced by the A cells of pars intercerebralis and the CA hormone in the vitellogenesis. 2. Biochemical studies reveal the incorporation of haemolymph proteins and lipids in the yolk during vitellogenesis. 3. Exogenous administration of the extract of the brain-CC and FME demonstrates, to some extent, the stimulation of protein synthesis during vitellogenesis in the haemolymph by the cerebral NSM and the CA hormone, while the lipid metabolism lies, mostly, under the control of juvenile hormone.     

Allatostatin-immunoreactive neurons projecting to the corpora allata of adult Diploptera punctata

Summary A monoclonal antibody against allatostatin I was used to demonstrate the allatostatin-immunoreactive pathways between the brain and the corpus cardiacum-corpus allatum complex in the adult cockroach Diploptera punctata. The antibody was two to three orders of magnitude more sensitive to allatostatin I than to the other four known members of the allatostatin family. Whole and sectioned brains in which immunoreactivity was localized with horseradish peroxidase-H₂O₂-diaminobenzidine reaction showed strongly immunoreactive cells in the pars lateralis of the brain with axons leading to and arborizing in the corpus cardiacum and the corpus allatum. Although many neurosecretory cells of the pars intercerebralis project to the corpora allata only, four strongly immunoreactive cells were evident here (two pairs on either side), and these did not project to the corpus cardiacum and corpus allatum but rather terminated within the protocerebrum in areas in which lateral cells also formed arborizations. Immunoreactivity was found in many other cells in the brain, especially in the tritocerebrum.