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R. SRIDARAN
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C. E. McCORMACK
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SUMMARY

Continuous monitoring of wheel-running activity and determination of the time of ovulation in rats by serial laparotomies revealed that ovulation followed the onset of running at prooestrus by approximately 9 h (range 7–11 h). This temporal relationship held in rats in which the period of the circadian rhythm had been modified (entrained) by daily exposure to 14 h photoperiods, and in rats in dim continuous light whose rhythms were non-entrained (freerunning). Knowledge of this temporal relationship between the two rhythms made it possible to give bright light signals at known points in the circadian cycle of the rat and to observe the effects on the timing of running and ovulation in subsequent cycles. Giving daily light signals near the onset of running (i.e. at subjective dusk) delayed, whereas giving signals near the end of running (i.e. at subjective dawn) advanced, the time of running and ovulation in subsequent cycles. These results indicate that in rats exposed to the usual laboratory photoperiod the delaying effect of dusk light and the advancing effect of dawn light balance one another; thus the preovulatory surge of LH occurs at a relatively consistent time at prooestrus.

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A. M. McNicol
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I. D. Penman
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A. E. Duffy
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ABSTRACT

Using a metaphase arrest technique, mitotic activity was quantified in the adrenal cortex over a 24-h period in 14-day-old male Sprague–Dawley rats before functional rhythmicity of the hypothalamic pituitary-adrenal (HPA) axis is established, and after its onset, in 6- to 7-week-old rats. At all times, proliferative activity was greater in the younger animals, as previously reported. A significant circadian rhythm was identified in both groups, but the timing of the peak differed, lying between 17.00 and 21.00 h at 14 days and 11.00 and 15.00 h at 6–7 weeks. These results raise the possibility that functional rhythmicity of the HPA axis may alter an inherent proliferative rhythm.

Journal of Endocrinology (1989) 120, 307–310

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G. CHAZAL
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M. FAUDON
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F. GOGAN
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M. HERY
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C. KORDON
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E. LAPLANTE
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Implantation of a solid source of oestradiol into ovariectomized rats produced constant plasma concentrations of the hormone over a long period of time. Under these conditions, LH is released in a circadian pattern with a very marked peak in the afternoon. This circadian rhythm is synchronized to the light–darkness cycle, since it follows exactly a shift in the nycthemeral cycle. The first peak appeared on day 3 after placement of the oestrogen implant; its amplitude was constant from days 3 to 9 after implantation, and decreased gradually during prolonged implantation. The afternoon peak was not correlated with changes in the pituitary sensitivity to exogenous LH releasing hormone (LH-RH), since the LH response to increasing doses of the peptide could be superimposed in the morning and in the afternoon. However, the decreased amplitude of the rhythm observed after more than 9 days of implantation seemed to depend upon a progressive desensitization of the pituitary gland to LH-RH. Pituitary LH content also decreased as a function of implantation time. It is concluded that, under conditions of constant plasma oestradiol concentrations and of constant pituitary sensitivity to LH-RH, a daily activation of the neural trigger releasing pituitary gonadotrophins occurs.

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IRVING ZUCKER
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CATHERINE P. CRAMER
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E. L. BITTMAN
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SUMMARY

Locomotor activity of male hamsters was recorded during long-term exposure to constant light (LL), constant darkness (DD) and during entrainment (modification of a circadian rhythm) to a 14 h light: 10 h darkness photoperiod (14L: 10D). In LL the period of the activity cycle was substantially longer in hypophysectomized than in control animals. This difference persisted during tests in DD. Although hypophysectomy reduced the duration of the active phase in some hamsters, overall the difference between the groups was not significant. The phase angle of onset of activity in 14L: 10D was not affected by hypophysectomy. Hypophysectomized female hamsters tested in DD had activity rhythms whose periods were longer than those of control animals; they were also significantly less active than corresponding controls during the first 4 h of the subjective night but the duration of the active phase did not differ significantly between the groups. These results suggest that hormones of the pituitary-gonadal axis modulate the period of circadian oscillation.

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W. DUPONT
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P. BOURGEOIS
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A. REINBERG
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R. VAILLANT
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For a period of 21 months between May 1974 and September 1976, circadian variations in the plasma concentration of corticosterone were studied by competitive protein-binding techniques in mature male and female edible frogs living in their natural environment. Blood samples were taken from 8 to 12 frogs six times daily and conventional and cosinor methods were used for statistical analysis.

Circadian rhythms were not detected during February and March (time of hibernation). Circannual rhythms were detected in three parameters of the circadian rhythm. The mean concentration of corticosterone over a 24 h period (24 h mean) reached a peak on 1 May (between 15 April and 15 May; 95% limits of confidence); the annual mean value of the 24 h means was 1·97 ± 0·25 (s.e.m.) μg/100 ml, with an amplitude of 0·66 μg/100 ml (0·53–0·79 μg/100 ml; 95% limits of confidence). Circadian variations in the concentration of corticosterone were largest in May (peak of reproductive activity). The time at which the peak concentration of corticosterone occurred showed circannual variations; peak values were detected around 24.00 h in May, 19.00 h in July and 08.00 h in November.

Both circadian and circannual variations have therefore been demonstrated in an endocrine function of an amphibian in its natural habitat.

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A. H. Cincotta
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A. H. Meier
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ABSTRACT

Lipogenesis and blood glucose concentrations were determined at 4-hourly intervals in control and insulin-treated golden hamsters maintained on 14-h daily photoperiods (08·00–22·00 h). Lipogenesis was studied by measuring the incorporation of label into liver and fat pad lipids in animals killed 30 min after i.p. [3H]acetate injection and 2 h after insulin or saline (control) injections. Circadian rhythms of lipogenesis and plasma glucose concentration were present in both control and insulin-treated hamsters. In control animals most lipogenic activity occurred during the dark period and early during the daily photoperiod (14 h light: 10 h darkness). There were dramatic differences in the lipogenic (fat pad) and hypoglycaemic responses to insulin which varied as a function of the time of day at which insulin was injected. Insulin stimulated fivefold increases in lipid deposition (fat pad incorporation) when injected late during the dark period but had little or no effect 4–8 h after the onset of light. Daily injections for 8 days also produced variable cumulative effects on body fat stores as a function of the time of day. Insulin injected late during the dark period stimulated a 40% increase in abdominal fat weight over controls, whereas insulin injected at 4 and 12 h after the onset of light had no effect on abdominal fat weight. Insulin decreased plasma glucose concentrations markedly at 8 and 20–24 h after the onset of light but had no apparent hypoglycaemic activity (120 min after its injection) at 4 h after the onset of light. These response rhythms coupled with rhythms of insulin secretion provide a basis for temporal synergisms which could produce a spectrum of physiological conditions as a function of the phase relations between the rhythms.

J. Endocr. (1984) 103, 141–146

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N. D. Martensz
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S. V. Vellucci
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L. M. Fuller
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B. J. Everitt
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E. B. Keverne
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J. Herbert
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ABSTRACT

Circadian rhythms in cortisol and testosterone in both blood and cerebrospinal fluid (CSF) were studied in four groups of male and female talapoin monkeys. Samples were taken 4 h apart under two conditions: whilst the sexes were kept separate (isosexual) and again after 24 h of interaction (heterosexual). There were similar rhythms in cortisol in males and females during the isosexual condition, though in blood (but not in CSF) mean levels were higher in females. Heterosexual interaction increased cortisol levels in both sexes (though more so in males), and also altered the shape of the rhythm, acrophase being delayed by 4 h in males and by 2 h in females. The amplitude of the rhythm was not altered. Cortisol levels were positively correlated in both males and females with the amount of aggression received from other males, but not from females nor with the animals' social rank.

Circadian rhythms in serum testosterone in males were also altered by heterosexual interaction. Access to females delayed acrophase by 2 h, but had no effect on mean levels (unlike the effect on cortisol). As for cortisol, the amplitude of the testosterone rhythm remained unchanged. Serum testosterone was negatively correlated with aggression from males, but not from females nor with sexual interaction. This was associated with a pronounced decrease in the levels of testosterone during the night, not observed in males receiving no aggression from others. There was a non-significant trend towards a positive correlation between social rank and serum testosterone.

These results show that social behaviour in groupliving primates has major effects on the parameters of the circadian pattern of secretion of both cortisol and testosterone. Aggression received from males seems to be a potent factor associated with the daily rhythms in both hormones, though there may be rank-related effects in the case of testosterone.

J. Endocr. (1987) 115, 107–120

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F.C. Logue
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W.D. Fraser
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D.St.J. O'Reilly
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G.H. Beastall
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ABSTRACT

A pronounced circadian rhythm has been demonstrated for intact parathyroid hormone (1-84) in the serum of normal male adults. The broad nocturnal rise of parathyroid hormone (1-84) secretion appears to be of physiological significance, for it is accompanied by a significant rise in nephrogenous cyclic adenosine monophosphate. The rate of return of parathyroid hormone (1-84) to baseline concentrations varies between individuals, an observation which has implications for the optimal time of sampling for the investigation of possible mild hyperparathyroidism.

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C. E. MCCORMACK
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RAJAGOPALA SRIDARAN
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SUMMARY

In order to determine whether the timing of ovulation in rats was controlled by an endogenous circadian rhythm, the hour of ovulation was determined by observing tubal ova during laparotomy in adult rats exposed to full animal room illumination (150 lux) during daily photoperiods of 14 h (full LD), continuous 150 lux illumination (full LL), daily dim (0·2 lux) photoperiods of 14 h (dim LD), continuous 0·2 lux illumination (dim LL) or continuous darkness (DD). Rats in all groups except those exposed to full LL continued normal cyclic ovulation. By the second oestrous cycle, most rats in the full LL group failed to ovulate, even though they showed characteristic cyclic changes in the vaginal smear pattern. The hour at which ovulation occurred was similar in rats exposed to full LD, dim LD or DD but was delayed in rats exposed to full LL or dim LL; the longer the period of exposure, the greater was the delay. For a given length of exposure, ovulation was delayed more in full LL than in dim LL. The full LL used in this study produced persistent vaginal oestrus within 40 days, whereas the dim LL did not. The delayed ovulation in rats exposed to dim LL was associated with a delayed preovulatory surge of LH. These results are consistent with the hypothesis that the timing of the preovulatory surge of LH and ovulation are controlled by an endogenous circadian rhythm, which in most rats has a periodicity in continuous light of slightly longer than 24 h.

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M. B. ter HAAR
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P. C. B. MacKINNON
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M. G. BULMER
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SUMMARY

The incorporation of [35S]methionine into protein in various regions of the brain and in the anterior pituitary, and serum luteinizing hormone (LH) and follicle-stimulating hormone (FSH) levels were measured at 6 h intervals throughout a 24 h period in the following groups of Wistar rats: (1) normal adult males and females; (2) adult genetic males or females which had been respectively 'feminized' or 'masculinized' by androgen deprivation or administration in neonatal life. Similar measurements were made at 12 h intervals in adult male rats which had been castrated at 7 or 15 days of age.

Serum LH levels showed a circadian rhythmicity in normal adult animals of both sexes, with peak levels in the male occurring 6 h earlier than those in the female. There was no statistically significant circadian rhythm in FSH levels in any group of animals. In all groups of castrated animals LH and FSH levels were raised but no circadian rhythms were observed.

Incorporation of [35S]methionine into protein in all cerebral areas showed circadian rhythms, the peak values of which, in the adult males, were almost 8 h (120°) out of phase with those of the adult females. In the 'feminized' genetic males or 'masculinized' genetic females the rhythmic phase was reversed to that of the opposite genetic sex. Animals castrated at 7 days or 15 days of age did not appear to show a rhythm.

A 12 h rhythm of incorporation was apparent in the anterior pituitary of the normal adult male and the adult 'masculinized' female; no significant rhythm was seen in the normal adult female and the 'feminized' male.

It is suggested that a 'female type' rhythm of incorporation in the brain may be associated with the maintenance of oestrous cycles.

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