A possible neuroendocrine system in polynoid polychaetes

Within the supraesophageal ganglion of polynoids is a vertical fiber tract which has the appearance of a “Y” in transverse sections of the brain, and contains the axons of many neurosecretory cells. The granule-filled terminals of these neurosecretory fibers are found at the base of the tract where they are in contact with the inner surface of the sheath covering the ventral surface of the brain. This sheath separates these neurosecretory endings from an underlying pericapsular epithelium which is thicker in this region. Beneath this pericapsular epithelium is a coelomic sinus. The dorsal blood vessel is located within this sinus and is “innervated” by a pair of fiber bundles that pass out of the brain at the base of the vertical fiber tract. The outer surface of the vessel is covered by epithelioid cells which contact these fiber bundles and the thickened pericapsular epithelium, and sometimes contain granular cytoplasmic inclusions. The lumen of the vessel is continuous with the lumina of a pair of cellular, thickwalled structures of unknown function which are attached to the ventro-lateral margins of the brain. The relationship between neurosecretory endings, enlarged pericapsular cells, coelomic sinus and blood vessel provides morphological evidence for the hypothesis that these structures are elements of a neuroendocrine system, similar in some respects to the brain-infracerebral gland complex of nereid and nephtyid polychaetes.     

The infracerebral gland--a possible neuroendocrine complex in Nereis

The infracerebral gland of Nereis consists of an epithelium covering the ventral surface of the posterior region of the brain. The thickness of the epithelium varies greatly in different species, and it appears especially well developed in Nereis limnicola. Cells of the most numerous type are in direct contact with the base of the brain. Their apical surfaces bound a coelomic sinus, below which is a blood plexus. Other cells are fuchsinophilic and contain many inclusions resembling elementary neurosecretory granules. A third type is rare and resembles glial elements. A number of nerve tracts run from the neuropil to the base of the brain in the region of the gland. Where they impinge upon the capsule they form swellings containing elementary granules and small vesicles. Some axons do not end on the capsule but pass through the capsule and then ramify among the cells of the gland. The swollen endings of other fibers, probably nervous in character, are packed with mitochondria and are scattered over the inner surface of the capsule in the region of the gland. The features described are suggestive of a neuroendocrine complex, and the relation between the brain and the infracerebral gland is in need of experimental analysis in view of the important endocrine functions presently ascribed to the brain in nereids.     

Neuroendocrine control of oocyte development in Poecilocera picta (Insecta: Orthoptera)

The neuroendocrine control of oocyte development in Poecilocera picta Fab. has been described. The secretory activity of the A type of neurosecretory cells has been correlated with ovarian development. In P. picta during the first four days after emergence the neurosecretory material is seen passing down the axons but the cells are largely devoid of neurosecretory material. When the oocytes are developed the A type cells are with stored neurosecretory material.
In P. picta the mature males do not appear to accelerate the process of maturation in females. The females which are reared without males or with castrated males also mature at the same time as the females which are reared with males. The corpus allatum also enlarges and decreases at the same period. The number of resorptive bodies is much more in the females which are reared with castrated males or without males. There appears to be some correlation in the secretion of the neurosecretory material, copulation, and the appearance of resorptive bodies. In P. picta the role of the mature male is only in copulation which very likely allows the cell to synthesize and secrete and release a large amount of neurosecretory material whose discharge in the haemolymph enables a successful development of the oocyte. Corpus allatum appears to be controlled by a precursor from the brain.
Cautery of the cerebral neurosecretory cells, allatectomy and sectioning of the nerves have been done to find out the role of neurosecretory material.     

Presumptive neurosecretion in Leucochloridiomorpha constantiae (Trematoda) and its possible role in governing maturation

A review of literature dealing with neurosecretion in the Platyhelminthes is presented and presumptive neurosecretory cells associated with the cerebral ganglia and longitudinal nerve cords ofLeucochloridiomorpha constantiae metacercariae and adults are described. Both the mean number of aldehyde-fuchsin stained presumptive neurosecretory cells and the intensity of staining are less in adults than in metacercariae. This suggests that maturation in this trematode may be directly or indirectly associated with a reduction in neurosecretion as in nereid annelids. Results of cytochemical studies indicate the neurosecretory product contains a glycoprotein.     

A SURVEY OF NEUROENDOCRINE PHENOMENA IN NON-ARTHROPOD INVERTEBRATES

• 1 Neurosecretory phenomena are apparently ubiquitous among Metazoa.
• 2 In Hydra, neurosecretory products are probably involved in the control of growth and reproduction.
• 3 Secretory elements in the central nervous system of turbellarians probably promote fission, regeneration and reproduction.
• 4 The cerebral ganglia of nemertines are the source of a hormone which exercises an inhibitory influence on maturation of the gonads and the development of somatic sexual characteristics. A principle secreted by the ganglia and/or the associated cerebral organs controls weight regulation. A sex hormone controls sexual differentiation.
• 5 Secretory neurons may influence the production of exsheathing fluid and ecdysis in nematodes.
• 6 In nereid polychaetes, a single hormone which originates from the cerebral neuroendocrine system apparently promotes segment proliferation and inhibits maturation. However, in certain other polychaetes, contrasting endocrine mechanisms seem to operate.
• 7 In lumbricid oligochaetes, a hormone secreted by the cerebral ganglion (and possibly by other nervous centres also) promotes gonadal activity and the differentiation of somatic sexual characters. In some species the ganglion exerts an inhibitory influence on the associated processes of ‘diapause’ and posterior regeneration. The cerebral and suboesophageal ganglia are implicated in the control of osmoregulation. Cerebral neurosecretory cells in limicolous oligochaetes may influence regenerative growth and osmoregulation.
• 8 The cerebral ganglia of leeches secrete a gonadotrophic hormone.
• 9 In gastropod molluscs, hormones secreted by the cerebral ganglia, glandular ‘dorsal bodies’ and/or optic tentacles are responsible for the control of the development of the reproductive tract, and the activity of the gonads. Contrasting mechanisms are thought to operate in the different gastropod groups and there is conflicting evidence particularly with respect to the endocrine functions of the optic tentacles and the gonads. The ‘bag cells’ of the abdominal ganglion of ApZysiu secrete a hormone which induces egg-laying. Cells in the pleural and parietal ganglia probably control osmoregulation in Lymnaea.
• 10 The optic glands of cephalopods secrete a gonadotrophic hormone, but sex hormones are apparently absent. The functional significance of the neurovenous tissues (presumptive neurosecretory complexes of unusual character) is obscure in most cases.
• 11 The radial nerves of starfish are the source of a hormone which induces the production of I-methyl adenine by the follicle cells of the gonad. This second principle stimulates oocyte maturation and the shedding of male and female gametes.
• 12 The relevance of such information to the development of certain biological concepts and to various aspects of comparative physiology is briefly discussed.

     

The Cerebral Neurosecretory System,Secretory End-Foot System and Infracerebral Gland—a Probable Neuroendocrine Complex in Nephtys (Annelida; Polychaeta)

Abstract The brain of Nephtys contains four neurosecretory cell types with distinctive cytoplasmic inclusions, a cells are located uniquely in a single pair of ganglionic nuclei and b cells are represented by a single pair of cells, whereas c cells and d cells have a scattered distribution. Their axons form two types of secretory release structure. First, possible axon collaterals synapse upon slender “dentritic twigs” in the core of the brain. Secondly, two tracts descend to the brain floor to form a “neurosecretory neuropile” (or storage and release complex) in contact with the inner surface of the brain capsule. Other neurosecretory fibres penetrate through the capsule, branch extensively, and terminate in contact with its ventral surface in close association with the “infracerebral gland”. The gland is derived from the pericapsular epithelium and exhibits signs of specialization for glandular function. In contrast to certain other polychaetes, it does not contain secretory neuron perikarya. The secretory end-foot system is poorly developed. Its terminals are located adjacent to the neurosecretory neuropile, which they encircle. The cell bodies are probably represented by four e cells which, like the terminals, contain many mitochondria.     

Studies on the neurosecretory system and retrocerebral endocrine glands of Nezara viridula Linn. (Heteroptera: Pentatomidae)

The neurosecretory system and retrocerebral endocrine glands of Nezara viridula Linn. have been described on the basis of in situ preparations and histological sections employing the paraldehyde fuchsin (PF) and performic acid-victoria blue (PAVB) techniques. In the brain of N. viridula, there are two medial groups–each consisting of five neurosecretory cells which belong to A-type. The lateral neurosecretory cells are absent. The axons of the two groups of medial neurosecretory cells (MNC) compose the two bundles of neurosecretory pathways (NSP) that decussate in the anterodorsal part of the protocerebrum. The two pathways, after the cross-over, run deep into the protocerebrum and deutocerebrum and emerge as NCC-I from the tritocerebrum. The nervi corporis cardiaci-I (NCC-I) of each side which are heavily loaded with NSM terminate in the aorta wall. Thus, the neurosecretory material (NSM), elaborated in the medial neurosecretory cells of the brain, is stored in the aortic wall and nervi corporis cardiaci-I (NCC-I). The NCC-II are very short nerves that originate from the tritocerebrum and terminate in the corpora cardiaca (CC) of their side. Below the aorta, but dorsal to the oesophagus, lie two oval or spherical corpora cardiaca. A corpus allatum (CA) lies posterior to the corpora cardiaca (CC). The corpora cardiaca do not contain NSM; only the intrinsic secretion of their cells has been occasionally observed which stains orange or green with PF staining method. The corpus allatum sometimes exhibits PF positive granules of cerebral origin. A new connection between the corpus allatum and aorta has been recorded. The suboesophageal ganglion contains two neurosecretory cells of A-type which, in structure and staining behaviour, are similar to the medial neurosecretory cells of the brain. The course and termination of axons of suboesophageal ganglion neurosecretory cells, and the storage organ for the secretion of these cells have been reported. It is suggested that the aortic wall and NCC-I axons function as neurohaemal organ for cerebral and suboesophageal secretions.     

Uptake of tritium-labelled biogenic amines by the prostomium of the polychaete Nereis virens (Sars) (Annelida)

The uptake of tritium-labelled 5-HT, noradrenaline, 5-hydroxytrytophan, DOPA and dopamine by the cerebral ganglion and prostomial nervous system of the polychaete Nereis virens has been examined using radioautography at the level of the light microscope. Pronounced uptake of (3)H-5HT occurred in the antennal, palpal, tegumentary and nuchal nerves as well as in ganglionic nuclei 13, 14, 15, 16, 17, 20, 24 and 25, the mid-brain neuropile, the neurosecretory neuropil and the infracerebral organ; (3)H-NA uptake was observed in small cells in the prostomial epidermis, and the infracerebral organ; (3)H-dopamine only in one of two common types of epidermal mucus cells. Prostomial muscles labelled generally with (3)H-NA and at specific sites with (3)H-5HT. These observations support the concept of an efferent serotonergic system originating in several cerebral ganglionic nuclei and serving prostomial muscle and epidermis. Evidence for an afferent adrenergic system is less substantial. The role of dopamine remains obscure.     

Brain neurosecretory cell and ecdysiotropin activity of the non-diapausing,pre-diapausing and diapausing southwestern corn borer, Diatraea grandiosella Dyar

The location and number of brain neurosecretory cells were studied in the larval southwestern corn borer. One posterior, two median and two lateral groups of paraldehyde-fuchsin positive cells were found in each cerebral hemisphere.Implantation of brain parts containing different groups of neurosecretory cells revealed that the median neurosecretory cells contained higher ecdysiotropic activity than the other cell groups. In vitro culture of ecdysial gland with brain or brain-parts extract showed also that the median neurosecretory cells contained much higher ecdysiotropic activity than other neurosecretory cells. To estimate the ecdysiotropic activity of pre-diapausing 6th instar larvae, their brain or brain extract was incubated in culture medium containing an ecdysial gland from a day-4 last-instar non-diapausing larva. Data showed that the ecdysiotropic activity in the pre-diapausing larvae was far lower than in non-diapausing and diapausing larvae.     

The fine structure of the infracerebral complex of Perinereis cultrifera grube (annelida: Polychaeta): C1 and C2 cells

The infracerebral complex of Perinereis cultrifera is located along the posterior, ventral portion of the brain, between the brain capsule and the coelomic sinus. The C₁ cells are characterized by the presence of an entanglement of cell processes along the basal border, numerous filament bundles throug Golgi complexes with small vesicles nearby in the apical region. The C₂ cells, stellate in form, resemble protein-synthesizing neurosecretory cells because of the electrondense granules found in the cell body and its processes. This cell i cells, thereby eliminating direct contact with the coelomic sinus or the blood vessels although processes extend dorsally to the brain capsule. No significant ultrastructural change appears in either cell type during a 24-hr cycle or during the juvenile or reproductive phase of the animal's life.     

Electron microscopical and histochemical observations on the relation between medio-dorsal bodies and neurosecretory cells in the basommatophoran snails Lymnaea stagnalis,Ancylus fluviatilis,Australorbis glabratus and Planorbarius corneus

Summary In Basommatophora medio-dorsal bodies (MDB) are closely attached to the cerebral ganglia, in which, just underneath the bodies, groups of Gomori-positive neurosecretory cells (MDC) occur. It has been suggested that the MDB-cerebral ganglion complex should be regarded as a neuro-endocrine association.In the present study the morphological relation between MDB and the ganglion is histochemically and ultrastructurally investigated in Lymnaea stagnalis, Ancylus fluviatilis, Australorbis glabratus and Planorbarius corneus.Histochemical tests showed the paraldehyde-fuchsin positive material of fibers in the MDB to be different from the neurosecretory material (NSM) in the MDC. At the ultrastructural level no penetration of nerve cell processes through the perineurium, separating the MDB from the ganglion, into the medulla of the MDB was observed. However, excepting for Lymnaea, the perineurium at these places shows particular differentiations. In the medulla of the MDB granule laden profiles (granule ø 700–900 Å) occur. They appeared to be processes of MDB cells.From these results it is concluded that the medulla of the MDB should not be regarded as a neurosecretory neuropile. Apparently, the MDB-cerebral ganglion complex is no neuroendocrine association. Probably the MDB is an endocrine organ. The small electron dense granules of the profiles in the medulla were also found in the MDB cell bodies. They are thought to represent a secretion product. The close morphological relation between MDB and cerebral ganglion may be connected with the origin of the MDB cells from perineural elements.     

Further observations on the fine structure and development of the infracerebral complex (“infracerebral gland”) of Nereis limnicola (Annelida,Polychaeta)

Cell and Tissue Research - The infracerebral complex consists of: (a) two types of ependymoid infracerebral cells located on the ventral surface of the brain, adjacent to a coelomic sinus and blood...     

Endocrine functions of brain in adult and developing mammals

The main prerequisite for organism’s viability is the maintenance of the internal environment despite changes in the external environment, which is provided by the neuroendocrine control system. The key unit in this system is hypothalamus exerting endocrine effects on certain peripheral organs and anterior pituitary. Physiologically active substances of neuronal origin enter blood vessels in the neurohemal parts of hypothalamus where no blood-brain barrier exists. In other parts of the adult brain, the arrival of physiologically active substances is blocked by the blood-brain barrier. According to the generally accepted concept, the neuroendocrine system formation in ontogeny starts with the maturation of peripheral endocrine glands, which initially function autonomously and then are controlled by the anterior pituitary. The brain is engaged in neuroendocrine control after its maturation completes, which results in a closed control system typical of adult mammals. Since neurons start to secrete physiologically active substances soon after their formation and long before interneuronal connections are formed, these cells are thought to have an effect on brain development as inducers. Considering that there is no blood-brain barrier during this period, we proposed the hypothesis that the developing brain functions as a multipotent endocrine organ. This means that tens of physiologically active substances arrive from the brain to the systemic circulation and have an endocrine effect on the whole body development. Dopamine, serotonin, and gonadotropin-releasing hormone were selected as marker physiologically active substances of cerebral origin to test this hypothesis. In adult animals, they act as neurotransmitters or neuromodulators transmitting information from neuron to neuron as well as neurohormones arriving from the hypothalamus with portal blood to the anterior pituitary. Perinatal rats—before the blood-brain barrier is formed—proved to have equally high concentration of dopamine, serotonin, and gonadotropin-releasing hormone in the systemic circulation as in the adult portal system. After the brain-blood barrier is formed, the blood concentration of dopamine and gonadotropin-releasing hormone drops to zero, which indirectly confirms their cerebral origin. Moreover, the decrease in the blood concentration of dopamine, serotonin, and gonadotropin-releasing hormone before the brain-blood barrier formation after the microsurgical disruption of neurons that synthesize them or inhibition of dopamine and serotonin synthesis in the brain directly confirm their cerebral origin. Before the blood-brain barrier formation, dopamine, serotonin, gonadotropin-releasing hormone, and likely many other physiologically active substances of cerebral origin can have endocrine effects on peripheral target organs—anterior pituitary, gonads, kidney, heart, blood vessels, and the proper brain. Although the period of brain functioning as an endocrine organ is not long, it is crucial for the body development since physiologically active substances exert irreversible effects on the targets as morphogenetic factors during this period. Thus, the developing brain from the neuron formation to the establishment of the blood-brain barrier functions as a multipotent endocrine organ participating in endocrine control of the whole body development.     

An ultrastructural in vitro study on the regulation of neurosecretory activity in the freshwater snail Lymnaea stagnalis (L.) with particular reference to Caudo-dorsal cells

The neurosecretory Caudo-Dorsal Cells (CDC) in the cerebral ganglia of the freshwater pulmonate snail Lymnaea stagnalis produce an ovulation stimulating hormone. Previously it has been shown that neuronal and non-neuronal inputs are involved in the regulation of their activity. The degree of autonomy of these cells has been investigated by studying with morphometric methods the ultrastructure of CDC maintained in vitro. CDC of isolated cerebral ganglia which were cultured for 7 days show a considerable rate of synthesis, transport and release of neurohormone. Apparently these processes can proceed in the absence of neuronal and hormonal inputs from outside the cerebral ganglia. Completely isolated CDC, however, do not show neurosecretory activity in vitro; active Golgi zones, indicating the formation of neurosecretory elementary granules, are absent from such cells. Isolation does not seem to affect general cell functions such as protein synthesis and respiration. It is suggested that a neuronal input, originating within the cerebral ganglia, is necessary for the stimulation of CDC neurosecretory activity. Techniques are described for the isolation and culture of neurosecretory cells of L. stagnalis.     

The Neurosecretory Neuron in Neuroendocrine Regulatory Mechanisms

The widespread occurrence of neurosecretory neurons in the animalkingdom suggests a functional significance that is basic andspecial. The explanation of the need for this unusual cell typelies in the fact that it forms a link between the nervous andthe endocrine systems whose functional interdependence formsthe basis for the effectiveness of regulatory mechanisms inthe animal world. These two integrative systems function indifferent ways. The neurosecretory cell, with its dual characteristics,and this cell alone, seems capable of receiving messages in"neural" language, and of transmitting this information in modified"endocrine" language to glandular cells. The neurosecretoryneuron occupies a central position in neuroendocrine interactions,not only because it is geared for communication with the endocrineapparatus, but because it serves as a singular channel ("finalcommon path," E. Scharrer, 1965) through which a multitude ofafferent stimuli, after being processed, are channeled to avariety of endocrine way stations and thus exert control overtheir effector organs.     

Neuroendocrine Regulation of the Development of Seasonal Forms of the Asian Comma Butterfly, Polygonia c-aureum L

In the butterfly, Polygonia c-aureum , development of seasonal forms controlled by the photoperiod and temperature was shown to involve a neuroendocrine system of the brain-corpus cardiacum-corpus allatum complex.
For analysis of the neuroendocrine system concerned, the innervation of the complex was investigated first by cobalt chloride perfusion staining and then by severance of axons, ablation of the candidate cells, injection of a homogenate of these cells and transplantation of corpora cardiaca using pupae programmed to be either summer-form or autumn-form adults.
The results suggested that medial nerve cells produce what is called material producing the summer form.
The seasonal forms of the Asian comma butterfly, Polygonia c-aureum L., summer and autumn forms (Fig. la, b), are determined by the photoperiod and the temperature during the larval period (1–3). Previous studies have given the following results on the physiological mechanism involved in the effect of environmental factors in inducing these seasonal forms. First, the mechanism involves neurosecretory cells located somewhere in the brain (2). Second, the nervous connections between the brain and the corpus cardiacum (NCC I+II (4)) and between the right and left brain lobes are indispensable for the effect (2, 5–7).
The present study consisted of two series of experiments. One was designed to demonstrate morphologically the axonal connection of the corpus cardiacum with the corpus allatum in this butterfly, like that shown in several other insects (8–13). The other series was designed to locate the neurosecretory cells producing material related to the seasonal form and to see if this material is also present in the corpus cardiacum.     

Specification and development of the pars intercerebralis and pars lateralis, neuroendocrine command centers in the Drosophila brain

The central neuroendocrine system in the Drosophila brain includes two centers, the pars intercerebralis (PI) and pars lateralis (PL). The PI and PL contain neurosecretory cells (NSCs) which project their axons to the ring gland, a complex of peripheral endocrine glands flanking the aorta. We present here a developmental and genetic study of the PI and PL. The PI and PL are derived from adjacent neurectodermal placodes in the dorso-medial head. The placodes invaginate during late embryogenesis and become attached to the brain primordium. The PI placode and its derivatives express the homeobox gene Dchx1 and can be followed until the late pupal stage. NSCs labeled by the expression of Drosophila insulin-like peptide (Dilp), FMRF, and myomodulin form part of the Dchx1 expressing PI domain. NSCs of the PL can be followed throughout development by their expression of the adhesion molecule FasII. Decapentaplegic (Dpp), secreted along the dorsal midline of the early embryo, inhibits the formation of the PI and PL placodes; loss of the signal results in an unpaired, enlarged placodeal ectoderm. The other early activated signaling pathway, EGFR, is positively required for the maintenance of the PI placode. Of the dorso-medially expressed head gap genes, only tailless (tll) is required for the specification of the PI. Absence of the corpora cardiaca, the endocrine gland innervated by neurosecretory cells of the PI and PL, does not affect the formation of the PI/PL, indicating that inductive stimuli from their target tissue are not essential for early PI/PL development.