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J F F Powell, S L Krueckl, P M Collins and N M Sherwood

Abstract

Three species of fish have become important in the study of reproduction and development. Rockfish are a model for developmental studies of live-bearing perch-like fish, whereas medaka and zebrafish are models for developmental and genetic studies. The forms of GnRH are identified in the brains of each of these fish and in the pituitary of the rockfish to investigate the role of GnRH in reproduction. Here, we report that grass rockfish (Sebastes rastrelliger) have three forms of GnRH in brain extracts as determined by HPLC elution position and RIA. These forms are identified as sea bream GnRH, chicken GnRH-II and salmon GnRH. In contrast, only two forms of GnRH were detected in brain extracts of medaka (Oryzias latipes) and zebrafish (Brachydanio rerio): salmon GnRH and chicken GnRH-II. Rockfish is distinct from medaka and zebrafish in that the most abundant form of GnRH in the rockfish pituitary is sea bream GnRH, whereas this form is absent in the other two fishes. The identification of sea bream GnRH in the rockfish brain and pituitary extracts indicates that the phylogenetic emergence of sea bream GnRH is earlier than the order Perciformes.

Journal of Endocrinology (1996) 150, 17–23

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G. J. Boer and M. Rieutort

Levels of GH in serum were assayed during the development of heterozygous (HET) control and vasopressin-deficient homozygous (HOM) Brattleboro rats. In early postnatal growth no differences in GH concentrations were present between HET and HOM rats for the rapid decline in serum levels of GH in the first week and the constant period up to day 24 of age thereafter. However, higher values were found in 55-day-old HOM rats and lower values at the age of 9 months. It is concluded that the stunted development of the body and brain of HOM rats is not GH-related, and that changes or anomalies in GH secretion appear only after neurogenesis has been completed.

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N Zmora, D Gonzalez-Martinez, JA Munoz-Cueto, T Madigou, E Mananos-Sanchez, SZ Doste, Y Zohar, O Kah and A Elizur

The cDNA sequences encoding three GnRH forms, sea bream GnRH (sbGnRH), salmon GnRH (sGnRH) and chicken GnRH II (cGnRH II), were cloned from the brain of European sea bass, Dicentrarchus labrax. Comparison of their deduced amino acid sequences to the same forms in the gilthead sea bream, Sparus aurata, and striped bass, Morone saxatilis, revealed high homology of the prepro-cGnRH II (94% and 98% respectively), and prepro-sGnRH (92% to both species). The sbGnRH exhibited dissimilar identities, with high homology to the striped bass (93%), and lower homology (59%) to the gilthead sea bream. Two transcript types were identified for the GnRH-associated peptide (GAP)-sGnRH as well as for the GAP-cGnRH II, which suggests a possible alternative splicing followed by the addition of an early stop codon. In order to obtain antibodies specific for the three GnRH precursors, recombinant GAP proteins were produced. The differential expression of the three GnRHs previously reported in the brain by means of in situ hybridization, using riboprobes corresponding to the GAP-coding regions, was fully confirmed by immunocytochemistry using antibodies raised against the recombinant GAP proteins, indicating that the transcripts are translated into functional proteins. Moreover, this approach allowed us to follow, for the first time, the specific projections of the different cell groups: sGAP fibers are distributed mainly in the forebrain with few projections reaching the pituitary, sbGAP fibers are mainly present in the preoptic area, mediobasal hypothalamus and predominantly project to the pars distalis of the pituitary, whereas cGnRH II fibers have a widespread distribution primarily in the posterior brain, and do not project to the pituitary. These new tools will be extremely useful to study further the development, regulation and functional significance of three independent GnRH systems in the brain of vertebrate species.

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S C van Buul-Offers, K de Haan, M G Reijnen-Gresnigt, D Meinsma, M Jansen, S L Oei, E J Bonte, J S Sussenbach and J L Van den Brande

Abstract

In order to determine the effects of IGF-II overexpression on growth of mice, transgenic mice were produced carrying one of three different H-2Kb human IGF-II minigenes in which different non-coding exons (exon 5, truncated exon 5 or exon 6) preceded the coding exons 7, 8 and 9. These were spaced by truncated introns and for proper polyadenylation an SV40 polyadenylation signal was incorporated. The highest levels of IGF-II minigene mRNA expression were found in lines containing the truncated exon 5 construct (II5′). Those containing exon 6 (II6) had less expression and 5 constructs (II5) gave only moderate levels of mRNA expression. In general mRNA expression was highest in thymus and spleen, low in liver and kidney and absent in the brain. In addition, one 115' line showed expression in the brain. Serum IGF-II levels at 8 weeks of age were increased 7- to 8-fold in homozygous transgenic lines with construct II5′ without brain expression and 2- to 3-fold in the one that showed expression in the brain; serum IGF-I levels were unchanged. Serum IGFs in the lines containing the constructs 115 and 116 were not different from those of the controls. In all cases body length and weight as well as the weight of several organs such as brain, liver, kidneys, heart and spleen when expressed as a function of age did not differ from controls. Only the thymus showed a significant increase in weight in the transgenics II5′.

Inbreeding of 2 lines containing construct 115' with pituitary deficient Snell dwarf mice did not influence body length or weight despite increased serum IGF-II levels. Again the thymus showed a marked increase in growth. The biological activity of the IGF-II peptide was further demonstrated by increased serum IGF-binding protein-3 in the transgenic dwarf mice, as shown by Western ligand blotting.

In summary, overexpression of IGF-II in transgenic normal and dwarf mice does not affect overall body growth, but causes increased growth of the thymus. This suggests a role for IGF-II in thymic development by paracrine/autocrine action.

Journal of Endocrinology (1995) 144, 491–502

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P. SÖDERSTEN

Lordosis behaviour was induced in immature 20-day-old male rats by sequential treatment with oestradiol benzoate (OB) and progesterone, but prepubertal male rats were behaviourally less sensitive to the OB and progesterone treatment than were female rats. Thus, the sex difference in the lordosis response was present early during development. Castration at various times after birth showed that the capacity of immature rats to show lordosis is normally inhibited by an action of testicular secretions exerted during the first 10 days of life. Treatment of day 0 castrated rats with OB, either as a single injection given on the day of birth or as daily injections given on the first 10 days after birth, was much more effective in inhibiting the display of lordosis behaviour at 30 and 37 days of age than was treatment with testosterone benzoate (TB). Treatment with dihydrotestosterone benzoate neonatally had no inhibitory effect. Treatment of intact male rats or day 0 castrated OB-or TB-treated rats with the anti-oestrogen ethamoxytriphetol (MER-25) during the first 10 days of life antagonized the inhibitory effect of the testes and of the OB or TB treatment on the development of the lordosis response. It is suggested that during normal development oestradiol formed in the brain from testosterone in the circulation acts during the first 10 days of life to inhibit the capacity of male rats to show lordosis when adult.

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B. T. DONOVAN

SUMMARY

The effect of hypophysectomy or of division of the pituitary stalk on the growth and function of the corpora lutea in the ovaries of the pseudopregnant ferret was studied, and compared with the changes seen in the ovaries and uterus of normal animals. Hypophysectomy caused a fall in the weight of the ovaries and uterus and regression of the corpora lutea. Isolation of the pituitary gland from the brain was compatible with full development of the corpora lutea and did not interfere with the growth of the uterus during the first 4 weeks of pseudopregnancy. Later on, the ovarian and uterine weights fell below those of control animals. Blank operations, and stalk section with subsequent regeneration of the portal vessels, did not disturb luteal function. It is concluded that the pituitary gland of the pseudopregnant ferret secretes a luteotrophic hormone and that an additional factor, possibly oestrogen, may be required for optimal luteal activity.

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RYOKO KAKIHANA, STEPHEN BLUM and SEYMOUR KESSLER

SUMMARY

The development of the pituitary-adrenocortical stress response was studied in CBA/J × DBA/2J hybrid mice. On the basis of the plasma corticosterone response 15 min after a subcutaneous injection of histamine dihydrochloride (50 mg/kg), the first three neonatal weeks could be divided into stress-nonresponsive (3–211 days) and stress-responsive 16–21 days) periods. During the former period, corticosterone levels in the brains of the non-stressed control mice were 63% higher than those of comparable mice during the latter period. Histamine stress significantly increased corticosterone concentrations in the brain during both these periods, but the increase was much greater (88%) during the stress-responsive period than during the stress-nonresponsive period (29%).

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A. J. ZOLOVICK

Agents known to alter synthesis, storage, release, reuptake or catabolism of the monoamines (Lippmann, 1968; Rubinstein & Sawyer, 1970), and excess amounts of 5-hydroxytryptamine (5-HT), can both interfere with ovulation and ovarian development (Brown, 1967; Vaughan, Benson & Norris, 1970). Conversely, manipulation of sex steroid levels alters hypothalamic catecholamine content (Lichtensteiger, Korpela, Langemann & Keller, 1969), and cyclic variations in hypothalamic catecholamines (Lichtensteiger, 1969) and monoamine oxidase activity have been observed during the oestrous cycle (Zolovick, Pearse, Boehlke & Eleftheriou, 1966).

Whether one monoaminergic system alone regulates gonadotrophin secretion or whether secretion is dependent upon a more complex integrative mechanism, as suggested by Lippmann (1968), remains to be determined. In most investigations, one or a combination of several pharmacological agents have been employed to alter brain levels of the various monoamines. However, these agents do not permit the selective manipulation of only one monoaminergic system without affecting others. Recently, intracisternal

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R J A Helliwell and L M Williams

Abstract

The pineal hormone, melatonin, is important in the timing of seasonal reproduction in the sheep. Melatonin of maternal origin readily crosses the placenta; its function in the fetal sheep is, however, unclear. To gain an insight into the role of melatonin in ovine development we have identified specific melatonin receptors throughout gestation using 2-[125I]iodomelatonin and quantitative in vitro autoradiography. Specific binding was found at the earliest time studied at 30 days of gestation, over the developing thyroid (term=145 days). At 31 days of gestation specific labelling was found over the thyroid and pituitary glands, the spinal nerves, nasal cavity and developing bronchi. This binding was diminished by over 50% in the presence of 10−4 m GTPγS (an analogue of guanosine triphosphate) indicating that the 2-[125I]iodomelatonin binding at this early stage of gestation represents a receptor coupled to a regulatory G-protein. By 40 days of gestation specific binding was found over the nasal epithelium, cochlear epithelium, regions of the brain, especially the hind brain and the vestibulocochlear and glossopharyngeal nerves, and both the pars distalis and pars tuberalis of the pituitary. As gestation proceeded, labelling over the pars distalis appeared to become more scattered in nature while that on the pars tuberalis remained consistent. Saturation studies of both the neuronal and pituitary binding sites at 121 days of gestation and in the newborn lamb revealed a single class of high-affinity binding sites with K d values in the picomolar range. Also at 121 days of gestation, binding over the fetal pars tuberalis was diminished in a dose-dependent manner by GTPγS, again confirming that specific binding is indicative of a receptor coupled to a regulatory G-protein. These data demonstrate a potential for sensitivity to melatonin from early in gestation, as well as the developmentally specific expression of the melatonin receptor in certain tissues, and suggest a wider role for melatonin in ovine fetal development than previously considered.

Journal of Endocrinology (1994) 142, 475–484

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G Schreiber

In larger mammals, thyroid hormone-binding plasma proteins are albumin, transthyretin (TTR) and thyroxine (T4)-binding globulin. They differ characteristically in affinities and release rates for T4 and triiodothyronine (T3). Together, they form a 'buffering' system counteracting thyroid hormone permeation from aqueous to lipid phases. Evolution led to important differences in the expression pattern of these three proteins in tissues. In adult liver, TTR is only made in eutherians and herbivorous marsupials. During development, it is also made in tadpole and fish liver. More intense TTR synthesis than in liver is found in the choroid plexus of reptilians, birds and mammals, but none in the choroid plexus of amphibians and fish, i.e. species without a neocortex. All brain-made TTR is secreted into the cerebrospinal fluid, where it becomes the major thyroid hormone-binding protein. During ontogeny, the maximum TTR synthesis in the choroid plexus precedes that of the growth rate of the brain and occurs during the period of maximum neuroblast replication. TTR is only one component in a network of factors determining thyroid hormone distribution. This explains why, under laboratory conditions, TTR-knockout mice show no major abnormalities. The ratio of TTR affinity for T4 over affinity for T3 is higher in eutherians than in reptiles and birds. This favors T4 transport from blood to brain providing more substrate for conversion of the biologically less active T4 into the biologically more active T3 by the tissue-specific brain deiodinases. The change in affinity of TTR during evolution involves a shortening and an increase in the hydrophilicity of the N-terminal regions of the TTR subunits. The molecular mechanism for this change is a stepwise shift of the splice site at the intron 1/exon 2 border of the TTR gene. The shift probably results from a sequence of single base mutations. Thus, TTR evolution provides an example for a molecular mechanism of positive Darwinian evolution. The amino acid sequences of fish and amphibian TTRs are very similar to those in mammals, suggesting that substantial TTR evolution occurred before the vertebrate stage. Open reading frames for TTR-like sequences already exist in Caenorhabditis elegans, yeast and Escherichia coli genomes.