The significance of small pulses of gonadotrophin-releasing hormone

1987 ◽  
Vol 113 (3) ◽  
pp. 413-418 ◽  
Author(s):  
I. J. Clarke ◽  
J. T. Cummins
Keyword(s):  
Pulse Amplitude ◽  
Releasing Hormone ◽  
Gonadotrophin Releasing Hormone ◽  
Series Of Experiments

ABSTRACT A series of experiments was conducted to ascertain the significance of 'small' pulses of gonadotrophin-releasing hormone (GnRH). In the first experiment, ovariectomized hypothalamo-pituitary disconnected (HPD) ewes were given 250 ng pulses of GnRH every 2 h for 1 week, 25 ng pulses every 2 h for 24 h, 25 ng pulses hourly for 24 h and then alternating hourly pulses of 25 and 250 ng. During the 25 ng pulses, LH was not detectable in plasma and FSH concentrations declined after 2 days. Following the 25 ng pulses, the resumption of 250 ng pulses led to exaggerated LH responses (mean ± s.e.m. pulse amplitude 18·7 ± 1·7 vs 10·2 ± 1·2 μg/l in the first week). In a second experiment, ovariectomized–HPD ewes were maintained on 250 ng GnRH pulses every 2 h for 1 week and were then given three 25 ng pulses mid-way between the 250 ng pulses. Samples of blood were taken over three 250 ng pulses without 25 ng insertions and over three pulses with insertions. The insertion of 25 ng GnRH pulses did not cause LH pulses in their own right and did not alter the LH responses to the 250 ng pulses. In a third experiment, 50 ng GnRH pulses were inserted between the 250 ng GnRH pulses, as in experiment 2; these 50 ng pulses caused small LH pulses and led to a reduction in the response of the LH pulse amplitude to the 250 ng pulses. The 'small' LH pulses which occurred in response to 50 ng GnRH compensated for the reduced responses to the 250 ng pulses. Hence, the integrated area under the LH curve and between successive 250 ng pulses remained the same, irrespective of the 50 ng insertions. From these data we conclude that 'small' GnRH pulses alone can sustain ongoing LH synthesis without release, leading to an accumulation of releasable LH, and that the insertion of 'small' GnRH pulses may modify the pattern of pituitary responsiveness to 'large' GnRH pulses. J. Endocr. (1987) 113, 413–418

Influence of the characteristics of pulses of gonadotrophin releasing hormone on the dynamics of luteinizing hormone release from perifused sheep pituitary cells

1983 ◽  
Vol 98 (3) ◽  
pp. 411-421 ◽  
Author(s):  
R. P. McIntosh ◽  
J. E. A. McIntosh
Keyword(s):  
Luteinizing Hormone ◽  
Anterior Pituitary ◽  
Pulse Width ◽  
Pulse Amplitude ◽  
Threshold Value ◽  
Pituitary Cells ◽  
Hormone Release ◽  
Releasing Hormone ◽  
Lh Release ◽  
Gonadotrophin Releasing Hormone

The effects were studied of varying the frequency, width and amplitude of pulses of gonadotrophin releasing hormone (GnRH) on the release of LH from anterior pituitary cells. Dispersed sheep cells supported in Sephadex were perifused with medium for 10 h and stimulated with different constant pulse patterns of GnRH. The timing of release of LH was measured by radioimmunoassay of the effluent fractions. Pulses of GnRH ranging in duration from 2 min every 8 min to 16 min every 128 min, and in concentration from 1·7 pmol/l to 250 nmol/l were applied to the cells, as well as continuous stimulation. Comparisons of differences between LH release patterns among samples of the same preparation of cells were used to demonstrate the effects of different GnRH stimulatory regimes. It was concluded that (1) the frequency of GnRH stimulation was important to the nature of LH release (periods shorter than about 16 min between pulses reduced LH output and caused faster desensitization of response), (2) the pulse width of GnRH input was important (the rising edge of the pulse produced greater LH output per unit of GnRH input than did continued application of GnRH within a pulse and wider pulses combined with shorter periods reduced LH output) and (3) over a threshold value of 5–10 nmol GnRH/1 pulse amplitude had little further influence on LH output or rate of desensitization in dispersed cells. These findings reinforce the hypothesis that the rising edge of the GnRH pulse is the major stimulant to LH release.

Secretion rates and short-term patterns of gonadotrophin-releasing hormone, FSH and LH in the normal stallion in the breeding season

1988 ◽  
Vol 117 (2) ◽  
pp. 197-206 ◽  
Author(s):  
C. H. G. Irvine ◽  
S. L. Alexander
Keyword(s):  
Breeding Season ◽  
Venous Blood ◽  
Pulse Amplitude ◽  
Interpulse Interval ◽  
Statistical Testing ◽  
Pulse Interval ◽  
Apparent Difference ◽  
Releasing Hormone ◽  
Gonadotrophin Releasing Hormone ◽  
Lh Secretion

ABSTRACT Pituitary venous blood was collected by a painless non-surgical cannulation method from five ambulatory stallions at 5-min intervals for 5–6 h during the breeding season. In four adult stallions, statistical analysis showed that pulses of gonadotrophin-releasing hormone (GnRH) and LH were coincident (P <0·01), as were pulses of FSH and LH (P <0·05). Furthermore, the patterns of changes in concentration of FSH and LH were highly correlated in each of the four stallions. However, seemingly ineffective pulses of GnRH were also observed, with 28% of GnRH pulses failing to induce a significant gonadotrophin pulse. In the four adult stallions the amplitude of pituitary venous gonadotrophin pulses varied markedly but no correlation with GnRH pulse amplitude was observed. Peak secretion of FSH, but not LH, during pulses was correlated with the length of the interpulse interval. Consequently, the ratio of FSH to LH during peaks was least (P <0·02) when the interpulse interval was 30 min or less. Thus, differential FSH and LH secretion was achieved within a constant steroid milieu. Two stallions had regular contact with oestrous mares, and in these horses the secretion of GnRH and gonadotrophins occurred almost continuously with rapid, rhythmic pulses superimposed upon a tonic background. Mean (± s.d.) interval between GnRH pulses was 31·4 ± 9·8 min and 27·7 ± 10·1 min. This secretory pattern was not observed in the two stallions which had infrequent contact with oestrous mares, although the small numbers precluded statistical testing of this apparent difference. No GnRH pulses were observed in one of these stallions, while in the other mean (± s.d.) GnRH pulse interval was 45·0 ± 48·7 min, the large variance being partly due to rapid pulses during a period in which the stallion teased mares. The fifth stallion was pubertal, and GnRH and LH secretion occurred in 15 and 0% of samples respectively, while low levels of FSH secretion were observed in 37% of samples and jugular testosterone levels were immeasurably low. We conclude that there is a statistically significant synchrony between pulses of GnRH, LH and FSH in the pituitary venous blood of stallions. Furthermore, decreasing intervals between gonadotrophin pulses result in a significant reduction in secretion of FSH but not LH. J. Endocr. (1988) 117, 197–206

Neonatal immunisation against a novel gonadotrophin-releasing hormone construct delays the onset of gonadal growth and puberty in bull calves

2012 ◽  
Vol 24 (7) ◽  
pp. 973 ◽  
Author(s):  
J. H. Hernandez-Medrano ◽  
R. W. Williams ◽  
A. R. Peters ◽  
D. Hannant ◽  
B. K. Campbell ◽  
...  
Keyword(s):  
Pulse Amplitude ◽  
Reproductive Organ ◽  
Saline Control ◽  
Testicular Development ◽  
Releasing Hormone ◽  
Bull Calves ◽  
Antibody Titres ◽  
Endocrine Status ◽  
Gonadotrophin Releasing Hormone ◽  
Pulse Characteristics

The aim of the present study was to investigate the effect of the neonatal immunisation of bull calves against a novel gonadotrophin-releasing hormone (GnRH) construct, comprised of GnRH coupled to the glycoprotein D subunit of the bovine herpes virus-1 (GnRH–BHV1 gD), on endocrine status, reproductive organ development and carcass quality. Eighteen bull calves received either GnRH construct (n = 9) or saline (control; n = 9) at 2, 6 and 13.5 weeks of age. Blood samples were taken to determine antibody titres against GnRH, FSH and testosterone (T) concentrations and LH pulse characteristics, with testicular circumference monitored monthly. Immunisation reduced LH pulse amplitude (P < 0.05) and T concentrations (P < 0.05), particularly at the peak in anti-GnRH titres after the second booster at 16 weeks of age (P < 0.001), but not when titres fell. Despite antibody titres decreasing after 16 weeks, immunisation reduced testicular size between 16 to 57 weeks of age (P < 0.05), provoking an 8-week delay in puberty onset, defined as testicular circumference ≥14 cm. In conclusion, neonatal immunisation induced a significant immune response against GnRH, provoking a temporary endocrine disturbance that had a long-term effect on testicular development, delaying the onset of puberty. These results support the hypothesis that a developmental window exists during testicular development, such that disturbance of the endocrine drive to the gonads during this period results in a longer-term impairment of gonadal function.

Effects of modifying gonadotrophin-releasing hormone input before and after the oestrogen-induced LH surge in ovariectomized ewes with hypothalamo-pituitary disconnection

1990 ◽  
Vol 127 (2) ◽  
pp. 223-233 ◽  
Author(s):  
D. J. Phillips ◽  
J. T. Cummins ◽  
I. J. Clarke
Keyword(s):  
Continuous Infusion ◽  
Positive Feedback ◽  
Pulse Amplitude ◽  
Secretory Component ◽  
Similar Response ◽  
Lh Surge ◽  
Releasing Hormone ◽  
Before And After ◽  
Gonadotrophin Releasing Hormone ◽  
Sufficient Magnitude

ABSTRACT The patterns of gonadotrophin-releasing hormone (GnRH) input to the pituitary gland that affect the expression of a positive-feedback event by oestrogen on LH secretion were investigated in ovariectomized ewes with hypothalamo-pituitary disconnection (HPD). In experiment 1, ovariectomized HPD ewes were given hourly i.v. pulses of 250 ng GnRH and an i.m. injection of 50 μg oestradiol benzoate (OB). The ewes were given a bolus pulse of 2·25 μg GnRH 16 h after injection of OB, followed by half-hourly pulses of 250 ng GnRH for 14 h (treatment A). The LH surge response was significantly (P <0·05) greater in these ewes compared with that in ewes given a continuous infusion of GnRH (250 ng/h) after the OB injection, followed by a continuous infusion of 500 ng GnRH/h after the bolus pulse of GnRH (treatment B). When no GnRH was administered after the OB injection, except for the bolus pulse of GnRH (treatment C), the surge response was significantly (P <0·05) reduced compared with that in treatment A, and was reduced compared with treatment B. These data suggest that GnRH pulses are important in the generation of the OB-induced LH surge, but that a baseline secretory component can prime the pituitary to some extent. experiment 2, a doubling of the continuous infusion dose of GnRH used in treatment B to 500 ng/h before the bolus pulse of GnRH and to 1 μg/h afterwards (treatment D) gave a similar response compared with treatment A, suggesting that if the baseline input of GnRH is of sufficient magnitude, it can overcome the lack of pulsatile input. In experiment 3, halving the GnRH pulse amplitude used in treatment A from 250 to 125 ng (treatment E) did not reduce the LH surge response, implying that when the GnRH input is in a pulsatile mode, the amplitude of GnRH pulses is less important than the pulsatile nature per se. In experiment 4, removal of GnRH input after the bolus pulse of GnRH (treatment F) significantly (P <0·05) reduced the surge response compared with when pulses were maintained (treatment A), indicating that GnRH input is still required once the LH surge has been initiated. Collectively, these experiments show that several forms of GnRH delivery, both pulsatile and baseline, can result in the full expression of a positive-feedback response in ovariectomized ewes treated with oestrogen. Journal of Endocrinology (1990) 127, 223–233

The oestrogen-induced surge of LH requires a 'signal' pattern of gonadotrophin-releasing hormone input to the pituitary gland in the ewe

1989 ◽  
Vol 122 (1) ◽  
pp. 127-134 ◽  
Author(s):  
I. J. Clarke ◽  
J. T. Cummins ◽  
M. Jenkin ◽  
D. J. Phillips
Keyword(s):  
Pituitary Gland ◽  
Pulse Amplitude ◽  
Pulse Frequency ◽  
Lh Surge ◽  
Releasing Hormone ◽  
Gnrh Secretion ◽  
Signal Pattern ◽  
Oestradiol Benzoate ◽  
Gonadotrophin Releasing Hormone ◽  
Baseline Values

ABSTRACT Two experiments were conducted with ovariectomized and hypothalamo-pituitary disconnected (HPD) ewes to ascertain the pattern of inputs, to the pituitary gland, of gonadotrophin-releasing hormone (GnRH) necessary for the full expression of an oestrogen-induced LH surge. The standard GnRH replacement to these sheep was to give pulses of 250 ng (i.v.) every 2 h; at the onset of experimentation, pulses were given hourly. In experiment 1, groups of sheep (n = 7) were given an i.m. injection of 50 μg oestradiol benzoate, and after 10 h the GnRH pulse frequency or pulse amplitude was doubled. Monitoring of plasma LH concentrations showed that a doubling of pulse frequency produced a marked increase in baseline values, whereas a doubling of amplitude had little effect on the LH response. In a second experiment, ovariectomized HPD sheep that had received hourly pulses of GnRH for 16 h after an i.m. injection of oil or 50 μg oestradiol benzoate were given either a 'bolus' (2·25 μg GnRH) or a 'volley' (500 ng GnRH pulses 10 min apart for 30 min, plus a 500 ng pulse 15 min later). Both groups then received GnRH pulses (250 ng) every 30 min for the next 13 h. Oestrogen enhanced the LH responses to the GnRH treatments, and the amount of LH released was similar in ovariectomized HPD ewes given oestrogen plus bolus or volley GnRH treatments and ovariectomized hypothalamopituitary intact ewes given oestrogen. These results suggest that the oestrogen-induced LH surge is initiated by a 'signal' pattern of GnRH secretion from the hypothalamus. Journal of Endocrinology (1989) 122, 127–134

Effects of luteolysis during late pregnancy on pituitary responsiveness to gonadotrophin-releasing hormone in the rat

1991 ◽  
Vol 128 (3) ◽  
pp. 411-418
Author(s):  
T. R. Koiter ◽  
G. C. J. van der Schaaf-Verdonk ◽  
G. A. Schuiling
Keyword(s):  
Plasma Concentrations ◽  
Control Level ◽  
Late Pregnancy ◽  
Ovariectomized Rats ◽  
Normal Delivery ◽  
Releasing Hormone ◽  
Oestradiol Treatment ◽  
Progesterone Treatment ◽  
Gonadotrophin Releasing Hormone ◽  
Series Of Experiments

ABSTRACT We investigated whether the increase in the gonadotrophin response to gonadotrophin-releasing hormone (GnRH) during the last days of pregnancy and the occurrence of parturition on day 22 of pregnancy in rats are due to the increase in the plasma concentrations of oestradiol-17β after luteolysis, which occurs around day 20. In a first series of experiments we studied the effects of s.c. implantation of two capsules containing oestradiol on basal and GnRH-stimulated secretion of LH and FSH before and after luteolysis. Before luteolysis, ovariectomy increased basal LH and FSH; oestradiol treatment prevented this increase partly (FSH) or completely (LH). Ovariectomy also lowered the LH response to the infusion of GnRH (100 ng/h). Oestradiol treatment, on the other hand, increased the LH and FSH responses of both intact and ovariectomized rats above the level in intact non-treated control rats. After luteolysis, ovariectomy increased basal FSH only. Treatment with oestradiol did not prevent the increase in basal FSH and ovariectomy diminished the LH response to GnRH infusion. Oestradiol treatment maintained the LH response in ovariectomized rats at the control level and increased the FSH responses of both intact and ovariectomized rats to a higher level than in control rats. Furthermore, the LH and FSH responses of the oestradiol-treated groups of intact and ovariectomized rats were higher after luteolysis than before. In a second series of experiments two capsules containing progesterone were s.c. implanted before or after luteolysis. Progesterone treatment suppressed the plasma concentration of oestradiol and the gonadotrophin responses to infusion of GnRH on the expected day of parturition in both groups of rats. Parturition was delayed only in the rats in which progesterone treatment had started before luteolysis. It was concluded that throughout pregnancy ovarian factors suppress basal FSH and that the increase in responsiveness to GnRH after luteolysis is due partly to an increase in oestradiol production and partly to an ovarian factor which augments the action of oestradiol. Furthermore, normal delivery does not require high plasma concentrations of oestradiol during the last day of pregnancy. Journal of Endocrinology (1991) 128, 411–418

Effects of castration on luteinizing hormone secretion and response to gonadotrophin-releasing hormone in sheep infected with Trypanosoma congolense

1996 ◽  
Vol 134 (1) ◽  
pp. 115-122 ◽  
Author(s):  
BM Mutayoba ◽  
PD Eckersall ◽  
IA Jeffcoate ◽  
MJA Harvey ◽  
V Cestnik ◽  
...  
Keyword(s):  
Luteinizing Hormone ◽  
Hormone Secretion ◽  
Pulse Amplitude ◽  
Trypanosoma Congolense ◽  
Endocrine Function ◽  
Plasma Testosterone ◽  
Ph Effects ◽  
Luteinizing Hormone Secretion ◽  
Releasing Hormone ◽  
Gonadotrophin Releasing Hormone

Mutayoba BM, Eckersall PD, Jeffcoate IA, Harvey MJA, Cestnik V. Holmes, PH. Effects of castration on luteinizing hormone secretion and response to gonadotrophin-releasing hormone in sheep infected with Trypanosoma congolense, Eur J Endocrinol 1996:134:115–22. ISSN 0804–4643 The effects of trypanosomiasis on the endocrine function of the hypothalamo-pituitary-gonadal axis were investigated before and after castration of Scottish Blackface rams infected with Trypanosoma congolense and uninfected controls. Blood samples were collected at 15-min intervals for 6 h before and at 10,20,40, 60, 80, 100 and 120 min after injection of synthetic gonadotrophin-releasing hormone (GnRH, 20 μg iv) 2 days before infection and 26 and 54 days after infection, with castration being performed 28 days after infection. Mean luteinizing hormone (LH) pulse amplitude was higher (3.3 ± 0.2 vs 2.6 ± 0.3 ng/ml) and mean plasma testosterone concentration was lower (4.1 ± 0.6 vs 7.6 ±1.2 nmol/l) in infected vs control rams 26 days after infection (p < 0.05). Mean plasma LH concentration and pulse amplitude increased in both groups after castration but both were significantly lower in infected compared to control rams (6.6 ±1.5 and 13.0 ± 2.2 ng/ml, p < 0.01; 7.7 ±0.9 and 11.6 ± 0.9 ng/ml, p < 0.001). respectively. However, LH responses to exogenous GnRH were similar in infected and control rams at each stage of the experiment, suggesting that the smaller increase in plasma LH after castration in infected rams was not caused by reduced responsiveness of the pituitary to GnRH but by alterations in GnRH secretion by the hypothalamus or its transport to the adenohypophysis. These results also demonstrate that impairment of testosterone secretion within 4 weeks of T. congolense infection in sheep may be due to testicular rather than pituitary effects. IA Jeffcoate, Department of Veterinary Physiology. University of Glasgow Veterinary School, Bearsden, Glasgow G61 1QH. UK

Biphasic response in the secretion of gonadotrophin-releasing hormone in ovariectomized ewes injected with oestradiol

1989 ◽  
Vol 123 (3) ◽  
pp. 375-382 ◽  
Author(s):  
A. Caraty ◽  
A. Locatelli ◽  
G. B. Martin
Keyword(s):  
Negative Feedback ◽  
Positive Feedback ◽  
Sampling Period ◽  
Pulse Amplitude ◽  
Portal Blood ◽  
Blood Collection ◽  
Biphasic Response ◽  
Releasing Hormone ◽  
Gonadotrophin Releasing Hormone ◽  
Lh Secretion

ABSTRACT In ovariectomized ewes, an injection of oestrogen initially inhibits the tonic secretion of LH, and then induces a large release of LH similar to the preovulatory surge in intact ewes. The pattern of hypothalamic secretion of gonadotrophin-releasing hormone (GnRH) into the pituitary portal blood during this biphasic response to oestrogen was investigated in conscious, unrestrained, ovariectomized adult Ile-de-France ewes during the breeding season. The ewes were ovariectomized and implanted with cannulae for portal blood collection on the same day. Seven days later, portal and peripheral blood samples were collected simultaneously every 5 min for 25 h. The ewes were injected with oestradiol-17β (25 μg i.v. and 25 μg i.m.) 6·25 h after the start of sampling. GnRH and LH were measured by radioimmunoassay in portal and jugular plasma samples respectively. A clear pulsatile pattern of LH secretion was observed before the oestradiol injection in all ewes, followed by the typical biphasic decrease (negative feedback) and increase (positive feedback) in mean concentrations. The sampling period was divided, for analysis, into pretreatment, negative feedback and positive feedback phases. Before injection with oestradiol, the GnRH pulses were clearly defined in portal blood and always synchronized with LH pulses in the peripheral circulation. The frequency was 5·9 ± 0·6 pulses/6 h (mean ± s.e.m.), and the amplitude was 31·6±7·6 pmol/l. During negative feedback, both the frequency (4·2 ± 0·5 pulses/6 h, P<0·01) and amplitude (15·2 ± 4·6 pmol/l, P<0·05) of the GnRH pulses decreased. During positive feedback, there was a large surge in the concentration of GnRH, due primarily to an increase in pulse frequency (11·0±1·3 pulses/6 h, P<0·01). A change in pulse amplitude was not detected, but there was a large increase in the basal level of GnRH (P<0·05). As a consequence of the changes in frequency and amplitude of the pulses, the mean levels of GnRH before injection with oestradiol (5·3 ± 1·0 pmol/l) differed (P<0·05) from those during negative (3·8±0·5 pmol/l) and positive (18·9±4·7 pmol/l) feedback phases. These results show that the biphasic pattern of LH secretion induced by oestrogen injection in short-term ovariectomized ewes is caused by parallel changes in the secretion of GnRH as well as changes in pituitary responsiveness to GnRH. An abrupt increase in the frequency of GnRH pulses appears to be a key component of the positive feedback mechanism which elicits the oestradiol-induced surges of both GnRH and LH. Journal of Endocrinology (1989) 123, 375–382

Neurophysiological control of the secretion of gonadotrophin-releasing hormone and luteinizing hormone in the sheep--a review

1991 ◽  
Vol 3 (2) ◽  
pp. 137 ◽  
Author(s):  
JC Thiery ◽  
GB Martin
Keyword(s):  
Luteinizing Hormone ◽  
Steroid Receptors ◽  
Pulse Amplitude ◽  
Pulse Frequency ◽  
Neural Pathways ◽  
Releasing Hormone ◽  
Diagonal Band ◽  
Gonadotrophin Secretion ◽  
Gnrh Neurons ◽  
Gonadotrophin Releasing Hormone

The anterior pituitary gland secretes pulses of luteinizing hormone (LH) in response to pulses of gonadotrophin-releasing hormone (GnRH) released into the hypophysial portal blood by the hypothalamus. The pulsatile nature of the secretions is very important because the frequency of the pulses is directly related to the activity of the GnRH neurons. We can therefore take advantage of this phenomenon to develop mechanistic interpretations of responses to experimental treatments designed to unravel the neural pathways that influence what is, arguably, the most important individual signal controlling the activity of the reproductive system. We might also resolve the disagreements in the literature covering the neuropharmacology of gonadotrophin secretion. In this review, we describe work towards this end in the sheep. Most (95%) of the 2500 GnRH cell bodies in the sheep brain are located in a region covering the anterior hypothalamus, the medial preoptic area, the diagonal band of Broca, and the septum. The axons of up to 50% of these cells terminate in the organum vasculosum of the lamina terminalis. The remainder terminate in the median eminence and form the final common pathway for the many factors that affect gonadotrophin secretion. Among the factors known to affect the frequency of the pulses (or the activity of the GnRH neurons) are nutrition, pheromones, photoperiod and gonadal steroids (negative and positive feedback). Factors that affect GnRH pulse amplitude are more difficult to determine because variations in pituitary responsiveness prevent the use of LH patterns as a 'bioassay'. Techniques developed recently have allowed the direct measurement of GnRH pulse amplitude and revealed inhibitory effects of oestradiol, but we do not know whether this effect is due to a reduction in the amount of GnRH released by each neurone or a reduction in the number of neurones releasing a pulse. It is unlikely that the factors that alter pulse frequency do so by directly affecting the GnRH cells. For example, it is obvious that other cells, with specific receptors for pheromonal or nutritional stimuli, formulate a signal that is transferred to the GnRH cells via interneurones. Similarly, it is likely that a hypothalamic clock intervenes between photoperiodic inputs and GnRH output. Opioidergic neurons have been proposed as a link in this system, but the complexity of their action makes it unlikely that they directly affect the GnRH neurons. The responses to steroids are simple and rapid, but steroid receptors have not been found in GnRH cells, so at least one other set of interneurones is involved.(ABSTRACT TRUNCATED AT 400 WORDS)

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