Mechanism of delayed ovulation in a vespertilionid bat, scotophilus heathi: role of gonadotropin, insulin, and insulin-like growth factor-1

The aim of this study was to evaluate the effects of luteinizing hormone (LH), follicle-stimulating hormone (FSH), insulin, and insulin-like growth factor-1 (IGF-1) on ovarian androstenedione synthesis to understand the mechanism responsible for delayed ovulation in Scotophilus heathi. We found that LH stimulated a dose-dependent increase in androstenedione synthesis by the ovary in vitro. This study also showed a clear seasonal variation in the ability of the ovary to produce androstenedione in vitro in response to LH and FSH stimulation. In response to LH and FSH, maximum quantities of androstenedione were produced during recrudescence in November. The same doses of gonadotropins during the preovulatory period in February stimulated comparatively low androstenedione secretion by the ovary. On the basis of these data, we suggest that in S. heathi, ovarian responsiveness to LH and FSH peaks during recrudescence. This study also showed a seasonal variation in the effects of insulin and IGF-1 on ovarian androstenedione production in vitro. Peak ovarian responsiveness to insulin and IGF-1 was observed during quiescence in September. It is hypothesized that increased insulin/IGF-1 sensitivity during September may be responsible for increased responsiveness to LH. Increased LH release, if coincident with the period of enhanced ovarian responsiveness to LH, may result in the excessive androstenedione production responsible for delayed ovulation in S. heathi.     

Seasonal adiposity and androstenedione production as a possible mechanism for asynchronous reproductive activity between males and females of vespertilionid bat, Scotophilus heathi

Relationship between changes in the body weight and circulating androstenedione concentration was examined in order to find out their involvement in asynchronous gametic cycle of male and female S. heathi. Body weight of both male and female bats increased from September due to accumulation of white adipose tissue and reached to a peak in November. The body weight declined gradually due to consumption of adipose tissue during December and January reaching basal level in February. A significant correlation between body weight and serum androstenedione concentration was observed. Correlated with maximum body weight, peak androstenedione level was found during November in both male and female bats. Peak androstenedione level during November may be responsible for stimulating spermatogenesis in the male, but may suppress follicular growth and ovulation in the female, thus resulting in asynchronous gametic cycle of male and female S. heathi.     

Seasonal adiposity and delayed ovulation in a vespertilionid bat,Scotophilus heathi: role of tumor necrosis factor-alpha

The aim of this article was to evaluate the physiological significance of tumor necrosis factor-alpha (TNF-alpha) in seasonal accumulation of adipose tissue, hyperinsulinemia, and anovulation in Scotophilus heathi. The result showed seasonal variations in the circulating TNF-alpha level. A higher level of circulating TNF-alpha was observed during quiescence and recrudescence, whereas a lower level of TNF-alpha was observed during winter dormancy and the preovulatory period. An increased circulating TNF-alpha level coincided closely with accumulation of adipose tissue and hyperinsulinemia. Immunocytochemical localization of TNF-alpha in the ovary showed immunoreactivity mainly in the oocytes and theca-interstitial cells. The oocytes of small and medium-sized follicles showed strong TNF-alpha immunostaining, whereas weak immunoreactivity was observed in the large antral follicles. The atretic follicles showed mild TNF-alpha immunostaining. TNF-alpha immunoreactivity in the ovary was slightly higher during the quiescence and preovulatory periods compared with the periods of recrudescence and winter dormancy. TNF-alpha alone significantly increased androstenedione and estradiol production by the ovary in vitro but did not augment the luteinizing hormone (LH)-induced androstenedione production. However, TNF-alpha did augment LH-induced estradiol production. The results of this study suggest the involvement of TNF-alpha in the interaction among adipose tissue accumulation, insulin resistance, and ovarian activity in S. heathi.     

Seasonal adiposity, correlative changes in metabolic factors and unique reproductive activity in a vespertilionid bat, Scotophilus heathi

The aim of this study was to determine the effect of changes in body mass, fat reserves and feeding activity on circulating levels of lipids, glucose, protein and metabolic hormones in a vespertilionid bat, Scotophilus heathi. Furthermore, the relationship between changes in metabolic factors and hormones with the unique reproductive features of female S. heathi was also examined. The results of this study showed annual variation in body mass, fat reserve and feeding activity, which correlated significantly with circulating levels of lipids, protein and metabolic hormones. Increased corticosterone level during September-October in S. heathi promotes increased feeding activity, which in turn induces hyperinsulinemia in S. heathi during November. Hyperinsulinemia together with low body temperature in November facilitates fat accumulation in bat. Coinciding with the period of fat accumulation raises serum leptin level, which has been demonstrated to suppress ovarian activities thus causing delayed ovulation in S. heathi. Circulating levels of lipids were high during winter dormancy, which may provide energy to stored sperms. The study thus suggests that the unique reproductive features of female vespertilionid bat are strongly linked to fat deposition. J. Exp. Zool. 309A:94-110, 2008. (c) 2008 Wiley-Liss, Inc.     

Hormonal induction of ovulation stimulates atresia of antral follicles in a vespertilionid bat, Scotophilus heathi

Scotophilus heathi is a seasonally monoestrous subtropical vespertilionid bat found at Varanasi, India. Although the antral follicles remain present in the ovaries of S. heathi from November till March, ovulation is delayed in this species until early March. In order to understand the mechanism of ovulation suppression during this period of delayed ovulation, the effects of human chorionic gonadotropin (hCG), pregnant mare's serum gonadotropin (PMSG), follicle stimulating hormone (FSH) and gonadotropin releasing hormone agonist (GnRH agonist) on ovarian morphology and steroid concentration were investigated. Hormonal treatments were given as a single i.p. dose 24 h after capture. The bats were sacrificed 48 h after the injection. Treatment with hCG, PMSG, FSH and GnRH agonist failed to induce ovulation in S. heathi, although these hormones produced a high degree of ovarian stimulation. The administration of hCG and PMSG induced ovarian enlargement, intense hyperemia, marked changes in the interstitial cells (ICs), development of several antral follicles and a varying degree of abnormalities in the oocytes of most of the antral follicles. In the bats treated with hCG, PMSG and GnRH agonist, androstenedione concentration increased significantly to extraordinarily high levels, whereas estradiol concentration decreased. Administration of FSH caused regression of ICs and pyknosis of granulosa cells in the majority of antral follicles. FSH did not enhance androstenedione concentration. The results of the present study suggest that the failure of hormonal treatments to induce ovulation during the period of delayed ovulation might be due to a seasonal desensitization of ovarian follicles in S. heathi. The hormonal treatment instead stimulated the ICs to produce a high level of androstenedione resulting in atretic changes of the antral follicles.     

Immunolocalization of cytochrome P450 side chain cleavage, 17-alpha-hydroxylase and aromatase in the ovary of vespertilionid bat (Scotophilus heathi) during different phases of ovulatory delay.

Immunocytochemical localization of steroidogenic enzymes, cytochrome P450 side chain cleavage, 17-alpha-hydroxylase and aromatase, was performed in the ovaries of Scotophilus heathi during reproductive cycle, with reference to the period of delayed ovulation. Moderate immunoreactivity of side chain cleavage enzyme and 17-alpha-hydroxylase was observed mainly in thecal cells and interstitial cells of the ovarian stroma during quiescence. Thecal cells positive for 17-alpha-hydroxylase were found even around the primary follicles. The peak immunoreactivity for all the three enzymes was observed during recrudescence. It coincided with high circulating steroid levels during this period. In the stroma, immunoreactivity for side chain cleavage and 17-alpha-hydroxylase was so extensive that it almost occupied the entire interfollicular area of the ovary. Aromatase immunoreactivity declined, but side chain cleavage enzyme and 17-alpha-hydroxylase remained extensive during the period of delayed ovulation. This suggests a high androgen and low estrogen synthesis during the period of delayed ovulation. There was a marked decline in 17-alpha-hydroxylase and an increase in aromatase immunoreactivity during the preovulatory period, suggesting a decrease in androgen and increase in estrogen synthesis. The results suggest thecal cells and interstitial cells of the stroma as the major site of steroidogenesis in the ovary of S. heathi. Over production of androgen is attributed to the extensive development of 17-alpha-hydroxylase positive interstitial cells in the ovarian stroma, and this may be responsible for delayed ovulation in Scotophilus heathi.     

Changes in serum leptin, insulin, androstenedione and luteinizing hormone during ovarian cycle in the bat, Taphozous longimanus

Detailed reproductive pattern and associated endocrine characteristics have been documented in only a few species of order Chiroptera. The aim of the present study was to examine the changes in body weight, serum insulin, leptin, androstenedione and luteinizing hormone (LH) concentrations during annual ovarian cycle in the sheath-tailed bat, Taphozous longimanus. Bats were sampled over three years. Leptin, a satiety hormone produced primarily by adipose tissue, provides information to feeding center of the brain about nutritional status, fat mass, appetite and energy expenditure. The circulating concentration of leptin begins to increase from October and attains a peak in December. The peak serum leptin concentration coincides with body weight in November before winter dormancy in December. The serum leptin levels dissociate from body weight during December. The other peaks of serum leptin levels coincide with late stages of the two successive pregnancies. The serum insulin concentration begins to increase from September and attains a peak during December. The insulin concentration remains low from January to August. The circulating androstenedione concentration begins to increase in October, reaching a peak in December. This increase in androstenedione concentration correlated with the period of heavy accumulation of abdominal fat and increase in body weight. There was a sharp decline in androstenedione concentration and body weight in January. The serum LH shows peaks, in November, coinciding with the peaked body weight, the other peaks in January and May, coinciding with ovulation for the two successive pregnancies. The high leptin and insulin levels might be responsible for the maintenance of reproductive response and gonadal function during adverse environmental condition in the winter, while high androstenedione, and associated body weight along with LH might be responsible for maintaining basal gonadal function. We conclude that high leptin, androstenedione and insulin serve, as signal for the reproductive functions in that sufficient body fat are available to meet the caloric demands and maintain normal function during adverse winter conditions.     

Adiposity and Androstenedione Production in Relation to Delayed Ovulation in the Indian bat,Scotophilus heathi

Observations on body weight, circulating androstenedione concentrations and morphology of ovarian stroma were made in Scotophilus heathi during the period of delayed ovulation to make a comparison with polycystic ovarian syndrome in women. Body weight of bats increased from a level of 31.00 ± 0.30 g in August and reached a peak of 45.00 ± 0.46 g in November. This increase in body weight was due to accumulation of adipose tissue. The body weight declined gradually from December onwards and finally reached a basal level in March. The circulating androstenedione concentration showed a gradual increase from 36.80 ± 15.54 ng/ml in August and reached a peak level of 220.50 ± 50.10 ng/ml in November. Androstenedione concentration reached the lowest level in the March, just before ovulation. Morphological study showed extensive distribution of luteinized stromal cells or interstitial cells (ICs). Morphometric study showed that during the period of ovulatory delay, more than 75% area of the ovary was occupied by the ICs. Hyperandrogenism, anovulation, obesity (fat deposition) and stromal hyperthecosis present during delayed ovulation in S. heathi may serve as an experimental model for some aspects of the polycystic ovarian condition in women.     

Seasonal changes in circulating steroid concentration and their correlation with the ovarian activity in the female Indian sheath-tailed bat, Taphozous longimanus

The relationship between ovarian activity and circulating steroid concentration was studied in the female sheath-tailed bat, Taphozous longimanus. T. longimanus breeds twice in rapid succession during the year at Varanasi, India. Ovarian recrudescence was observed during September, and antral follicles were first observed during the month of October. Circulating androstenedione concentration showed an increase beginning in October, reaching a peak in December. This increase in androstenedione concentration correlated with the period of heavy accumulation of adipose tissue and increase in body mass. Antral follicles grow slowly during the period of high circulating androstenedione concentration from October to December. There was a sharp decline in androstenedione concentration during January. Simultaneously with the decline in androstenedione concentration, a sharp increase in size of the antral follicle and circulating estradiol concentration was noticed. Soon thereafter, one of the follicles ruptures, followed by fertilization and the commencement of the first pregnancy. During the second pregnancy, antral follicles first appeared during late pregnancy in March in the contralateral ovary that lacked the corpus luteum. One follicle developed rapidly and quickly attained a preovulatory stage in April. This is reflected in a sharp increase in estradiol concentration during this period. Ovulation was observed immediately following the first pregnancy in May. During this period, androstenedione concentration remained low. The results of the present study suggest that high androstenedione concentration during October to December (winter dormancy) may be responsible for slow follicular development and delays ovulation in T. longimanus. It is further hypothesized that the geographical variation in reproductive pattern of T. longimanus could be due to variation in the duration of fat storage and associated changes in the androstenedione concentration.     

Differential modulation of ACTH-stimulated cortisol and androstenedione secretion by insulin

Results of previous clinical studies suggested counter regulatory actions between insulin and DHEA(S). The present studies were performed using primary monolayer cultures of bovine fasciculata-reticularis cells to test the hypothesis that insulin directly affects adrenal androgen secretion. Although having no independent effect, insulin exhibited complex time- and concentration-specific actions on ACTH-stimulated secretion of both C21 (cortisol) and C19 (androstenedione) corticosteroids. In the presence of low concentrations (0.05-0.1 nM) of ACTH, cortisol secretion during a 2 h incubation was about 2-fold greater in the presence than in the absence of insulin (0.01-100 ng/ml). In the presence of a maximal concentration (10 nM) of ACTH, on the other hand, cortisol secretion was not affected by insulin at concentrations less than or equal to 0.1 ng/ml, but was decreased at higher insulin concentrations. ACTH-stimulated androstenedione secretion was not significantly affected by insulin during a short-term (2 h) incubation. During a prolonged (24 h) incubation, insulin produced a concentration-dependent inhibition of ACTH-stimulated cortisol secretion. At an insulin concentration of 100 ng/ml, ACTH (10 nM)-stimulated cortisol secretion declined to a level only 30% of that produced by ACTH alone. In contrast, insulin exhibited biphasic effects on the secretion of androstenedione by cells maintained in the presence of ACTH for 24 h; an effect that was most dramatic in the presence of a maximal concentration of ACTH. At an insulin concentration of 0.1 ng/ml, androstenedione secretion by cells maintained in the presence of 10 nM ACTH was increased approximately 2.5-fold. At higher concentrations of insulin, ACTH-stimulated androstenedione secretion was inhibited to an extent comparable to that in cortisol secretion. The effects of insulin on ACTH-stimulated cortisol and androstenedione secretion could not be accounted for by changes in steroid degradation or a loss in 11 beta-hydroxylase activity. These results indicate that insulin interacts with ACTH to modulate the secretion of both C21 and C19 corticosteroids and that physiological concentrations (less than or equal to 1 ng/ml) of insulin may have a long-term effect to enhance selectively adrenal androgen secretion. These data are consistent with a servo mechanism between insulin and DHEA(S) in vivo and indicate that the correlations observed clinically result, at least in part, from a direct action of insulin to modulate the rate of adrenal androgen production.     

24-hour changes in ACTH, corticosterone, growth hormone, and leptin levels in young male rats subjected to calorie restriction

Calorie restriction of young male rats increases plasma prolactin, decreases luteinizing hormone (LH) and testosterone, and disrupts their 24 h secretory pattern. To study whether this could be the consequence of stress, we examined the 24 h variations of plasma adrenocorticotropic hormone (ACTH) corticosterone, growth hormone (GH), leptin, and adrenal corticosterone. Rats were submitted to a calorie restriction equivalent to a 66% of usual intake for 4 weeks, starting on day 35 of life. Controls were kept in individual cages and allowed to eat a normal calorie regimen. Significantly lower ACTH levels were detected in calorie-restricted rats. Plasma corticosterone levels during the light phase of the daily cycle were significantly higher in calorie-restricted rats. Time-of-day variation in plasma ACTH and corticosterone levels attained significance in calorie-restricted rats only, with a maximum toward the end of the resting phase. The daily pattern of adrenal gland corticosterone mirrored that of circulating corticosterone; however, calorie restriction reduced its levels. Plasma ACTH and corticosterone correlated significantly in controls only. Calorie restriction decreased plasma GH and leptin, and it distorted 24 h rhythmicity. In a second study, plasma ACTH and corticosterone levels were measured in group-caged rats, isolated control rats, and calorie-restricted rats during the light phase of the daily cycle. Plasma ACTH of calorie-restricted rats was lower, and plasma corticosterone was higher, compared with isolated or group-caged controls. The changes in the secretory pattern of hormones hereby reported may be part of the neuroendocrine and metabolic mechanisms evolved to maximize survival during periods of food shortage.     

Pattern of ovarian protein synthesis and secretion during the reproductive cycle of Scotophilus heathi: synthesis of an albumin-like protein.

Ovarian proteins synthesized de novo and secreted in vitro were investigated during different stages of the reproductive cycle of Scotophilus heathi. We found increased ovarian protein synthesis and secretion during the recrudescence and the preovulatory periods, coinciding with two peaks of follicular development and steroidogenesis. The ovaries synthesized protein at a low rate during quiescence and the late phase of delayed ovulation. In vitro analysis using 35 S-methionine incorporation showed increased synthesis of 66 kDa protein during delayed ovulation, but secretion of this protein declined markedly during the preovulatory period. N-terminal sequencing of the 66 kDa protein showed that it shares 70% homology with human serum albumin. Immunocytochemistry indicated that this protein is found primarily in the granulosa cells of the follicles. Whether the increase in albumin production by the ovaries during delayed ovulation is associated with a hyperinsulinemic and hyperandrogenic condition requires further investigation.     

Photoperiodically-induced changes in hypothalamic-pituitary-adrenal axis sensitivity in captive house sparrows (Passer domesticus)

We used captive house sparrows (Passer domesticus) to identify regulatory mechanisms underlying seasonal (mimicked by changes in photoperiod) and diel differences in corticosterone output. Corticosterone responses were measured during three simulated seasons: short-day and long-day photoperiods and while birds underwent a pre-basic molt. Under all three conditions we tested for adrenal sensitivity by injecting exogenous ACTH, for pituitary sensitivity by injecting corticotropin-releasing factor (CRF) and arginine vasotocin (AVT), and for diel changes by repeating the injections during the day and at night. The daytime adrenal sensitivities were greatest on long days, lower on short days, and lowest during molt. These data suggest that reductions in either adrenal sensitivity to ACTH and/or capacity to secrete corticosterone could explain lowered endogenous corticosterone titers during molt. Furthermore, adrenal sensitivity to ACTH and pituitary sensitivity to AVT appeared to be greatest at night. This suggests that both the adrenal's sensitivity to the ACTH signal and the pituitary's capacity to secrete ACTH might provide a mechanism allowing for diel changes in corticosterone titers. This differs substantially from what is known about diel regulation in rodents. Taken together, these data provide further evidence that there are complex regulatory mechanisms controlling diel and seasonal changes in corticosterone titers in birds.     

Effect of aging on 24-hour pattern of stress hormones and leptin in rats

This work analyzes the 24-hour changes of hypothalamic-pituitary-adrenal (HPA) axis activity and leptin release in aged rats. Three- and 22-month-old male Wistar rats were killed at 6 time intervals during a 24-hour cycle (n=8-10 rats/group). Aging augmented plasma ACTH while it decreased plasma and adrenal gland corticosterone levels. Plasma and adrenal corticosterone levels attained high levels during all the scotophase, concomitantly with the maxima in ACTH levels, whereas in aged rats only a brief plasma corticosterone peak at the early scotophase and no time of day variations of adrenal corticosterone were observed. Aging augmented circulating leptin, with a significant interaction "agextime" in the factorial ANOVA, i.e. only in young rats time of day changes were significant, with the lowest values of leptin at the middle of the light period and higher values at night. When plasma leptin was expressed on body weight basis, the age-related differences became not significant but the daily pattern of plasma leptin found in young rats persisted. Plasma and adrenal corticosterone levels correlated significantly with plasma ACTH only in young rats. Likewise, plasma leptin correlated with plasma corticosterone only in young rats. These changes can be attributed to a disrupting effect of aging on the homeostatic mechanisms modulating HPA activity and leptin release.     

Selenium deficiency impairs corticosterone and leptin responses to adrenocorticotropin in the rat

Selenium deficiency causes oxidative stress and impairs steroidogenesis in vitro. Leptin is closely related to the hypothalamo-pituitary-adrenal (HPA) axis. Leptin inhibits the HPA axis at the central level while corticosteroids have been shown to stimulate leptin secretion in most studies. We hypothesized that oxidative stress impairs adrenal steroidogenesis and decreases leptin production in vivo. The goal of this study was to investigate in rats the effects of selenium deficiency and oxidative stress on adrenal function and on leptin concentrations. Weanling rats were fed a selenium-deficient (Se-) or selenium-sufficient (Se+) diet for 4-10 weeks. Selenium deficiency caused a marked decrease in liver (> or = 99%) and adrenal (> or = 81%) glutathione peroxidase (GPx) activities. Selenium deficiency did not affect basal and short-term adrenocorticotropin (ACTH) stimulated corticosterone or leptin concentrations. In contrast, after long-term ACTH stimulation, selenium deficiency caused a doubling in adrenal isoprostane content and blunted the increase in corticosterone and leptin concentrations observed in Se+ animals. Plasma leptin concentrations were 50% lower in Se- compared to Se+ animals following long-term ACTH. Our results suggest that oxidative stress causes a decrease in circulating corticosterone in response to ACTH, and, as a consequence, a decrease in plasma leptin concentrations.     

Photoperiod regulates corticosterone rhythms by altered adrenal sensitivity via melatonin-independent mechanisms in Fischer 344 rats and C57BL/6J mice

Most species living in temperate zones adapt their physiology and behavior to seasonal changes in the environment by using the photoperiod as a primary cue. The mechanisms underlying photoperiodic regulation of stress-related functions are not well understood. In this study, we analyzed the effects of photoperiod on the hypothalamic-pituitary-adrenal axis in photoperiod-sensitive Fischer 344 rats. We first examined how photoperiod affects diurnal variations in plasma concentrations of adrenocorticotropic hormone (ACTH) and corticosterone. ACTH levels did not exhibit diurnal variations under long- and short-day conditions. On the other hand, corticosterone levels exhibited a clear rhythm under short-day condition with a peak during dark phase. This peak was not observed under long-day condition in which a significant rhythm was not detected. To analyze the mechanisms responsible for the photoperiodic regulation of corticosterone rhythms, ACTH was intraperitoneally injected at the onset of the light or dark phase in dexamethasone-treated rats maintained under long- and short-day conditions. ACTH induced higher corticosterone levels in rats examined at dark onset under short-day condition than those maintained under long-day condition. Next, we asked whether melatonin signals are involved in photoperiodic regulation of corticosterone rhythms, and rats were intraperitoneally injected with melatonin at late afternoon under long-day condition for 3 weeks. However, melatonin injections did not affect the corticosterone rhythms. In addition, photoperiodic changes in the amplitude of corticosterone rhythms were also observed in melatonin-deficient C57BL/6J mice, in which expression profiles of several clock genes and steroidgenesis genes in adrenal gland were modified by the photoperiod. Our data suggest that photoperiod regulates corticosterone rhythms by altered adrenal sensitivity through melatonin-independent mechanisms that may involve the adrenal clock.     

Strategic approach of hyperandrogenism in women

Based on the results of the laboratory screening of 35 hirsute patients, a strategic approach of hyperandrogenism is proposed. The determination of plasma concentration of total testosterone (T), androstenedione (A) and dehydroepiandrosterone sulfate (DHEAS) appeared to be the basic investigation before to further explore the adrenal (ACTH test) and/or ovarian (gonadotropin measurements) androgen secretion. Together with the clinical findings, plasma T, A and DHEAS levels generally bring evidence of androgen-secreting tumors or polycystic ovaries, or suggest the possibility of adrenal hyperplasia. In the other cases, only the determination of plasma unbound T level is useful for demonstrating the hyperandrogenism. Finally in patients with 'so-called' idiopathic hirsutism, while evidence of excessive androgen production can be obtained by the laboratory screening, the origin of this hyperandrogenism remains uncertain in most cases.     

Histochemical localization of enzymes and lipids in the ovary of a vespertilionid bat, Scotophilus heathi, during the reproductive cycle.

The present study describes seasonal changes in delta5 3beta hydroxysteroid dehydrogenase (3beta-HSD), glusose-6 phosphates dehydrogenase (G-6-PD), and lipids in the ovary of a vespertilionid bat, Scotophilus heathi. Total lipids and 3beta-HSD activity are restricted to thecal and interstitial cells of the ovary. The total lipids, 3beta-HSD, and G-6-PD significantly increase during recrudescence, and remain high during winter dormancy and breeding as compared to the other reproductive phases. High incidence of lipids and enzyme activity in interstitial cells during the breeding period and at the time of ovulation clearly suggests that these cells are actively involved in steroidogenesis. A decline in enzymes and lipid activity during winter dormancy, which correlates with the declining levels of steroidogenesis, might be the factors responsible for prolonged survival of the Graafian follicle in the ovary of S. heathi.     

Annual variations of adrenal gland hormones in the lizard <Emphasis Type="Italic">Podarcis sicula</Emphasis>

The adrenal gland regulates metabolism and maintains normal electrolyte balance. Adrenal hormones are equivalent in all vertebrates; the chromaffin tissue produces adrenaline and noradrenaline and the steroidogenic tissue produces most of the steroid hormones present in mammals. Podarcis sicula belongs to the Squamata family of lizards and it is the most abundant lizard species in southern Italy. This species shows a reproductive annual cycle and the presence of seasonal variations in the activity of the hypothalamus-hypophyseal-thyroid axis. To investigate the existence of an annual cycle of lizard adrenal gland, we have measured plasma concentrations of corticosterone, ACTH, noradrenaline and adrenaline. We have shown that corticosterone rapidly increased from January to March to reach a peak value that persisted until July, then, it slowly decreased until December. ACTH levels increased from January to May and slowly decreased from July to December. Noradrenaline levels were higher in March and then decreased until December. On the contrary, adrenaline levels increased from March to July and slowly decreased until December. Our results demonstrate the existence of an annual cycle of the lizard adrenal gland activity. This is very interesting because its activity is important to rightly regulate the reproductive status of the Podarcis sicula lizard.     

Dynamics of food availability, body condition and physiological stress response in breeding Black-legged Kittiwakes

1. The seasonal dynamics of body condition (BC), circulating corticosterone levels (baseline, BL) and the adrenocortical response to acute stress (SR) were examined in long-lived Black-legged Kittiwakes, Rissa tridactyla , breeding at Duck (food-poor colony) and Gull (food-rich colony) Islands in lower Cook Inlet, Alaska. It was tested whether the dynamics of corticosterone levels reflect a seasonal change in bird physiological condition due to reproduction and/or variation in foraging conditions.
2. BC declined seasonally, and the decline was more pronounced in birds at the food-poor colony. BL and SR levels of corticosterone rose steadily through the reproductive season, and BL levels were significantly higher in birds on Duck Island compared with those on Gull Island. During the egg-laying and chick-rearing stages, birds had lower SR on Duck Island than on Gull Island.
3. The results suggest that, in addition to a seasonal change in bird physiology during reproduction, local ecological factors such as food availability affect circulating levels of corticosterone and adrenal response to acute stress.