Seasonal changes of serum concentrations of estradiol and progesterone in Bolivian squirrel monkeys (Saimiri sciureus)

Serial measurements of estradiol (E2) and progesterone (P) were used to describe the ovarian cycle of the Bolivian squirrel monkey. Group-caged, sexually mature female squirrel monkeys, housed with males, were sampled daily between 0900 and 1100 hr. Sampling was carried out during the breeding and nonbreeding seasons, for periods of 19–20 days from September 1981 to May 1982. Seasonal differences in serum concentrations of E2 and P were found with low levels of E2 and P and an absence of preovulatory surges of E2 during the nonbreeding season. This pattern was also observed in some animals during the breeding season. An abrupt increase in serum P concentrations in December appeared to signal the onset of cycling. Cycling animals had well-defined peaks of E2 (450–9,500 pg/ml) followed by increasing levels of P, which were >200 ng/ml in some animals. After the breeding season, E2 and P levels returned to their initially low levels. Levels of both steroids in cycling animals were higher than those reported for other primates and for previous measurements made in squirrel monkeys. Cycle length based on time interval between consecutive E2 peaks varied from 6–12 days.     

Suppression of ovarian activity during the breeding season in suckling Japanese monkey (Macaca fuscata)

The menstrual cycle was examined in Japanese monkeys, short-day breeders, with or without nursing babies during the breeding season by determining the change in the plasma concentration of progesterone. Blood samples were obtained weekly from suckling and nonsuckling mothers for 6 months from September to February. Plasma concentrations of progesterone in suckling mothers did not show any apparent fluctuation and remained low throughout the study, while irregular menstruation was observed. In nonsuckling mothers, cyclic changes in plasma progesterone concentration accompanied by regular menstruation occurred from October. Annual birth rates were lower in monkeys housed in individual cages (17.6%) than in free-ranging monkeys in Miyajima colony, near Hiroshima in Japan (37.5%). These results indicate that the suckling stimulus for females with young born in spring is potent enough to suppress ovarian activity during the autumn breeding season.     

Age-related hormonal differences in cycling squirrel monkeys (Saimiri boliviensis boliviensis)

This study was planned to evaluate the poor reproductive performance of aged squirrel monkeys in a self-sustaining breeding colony. Two groups of monkeys aged 6–8 years (Group 1), and >12 years (Group 2) were evaluated. During the midbreeding season, daily blood samples were obtained for a period of 35 days from 10 animals in each group. Blood samples were assayed for serum estradiol (E), progesterone (P), and bioassayable luteinizing hormone (bio-LH) concentrations and the data were analyzed by ANOVA (analysis of variance). Bio-LH surges were less frequent in the aged breeders (Group 2). Total P output during periovulatory period (day ?;3 to day + 3) and the total P output during the luteal phases of the estrus cycles were significantly lower in aged squirrel monkeys (Group 2, P < 0.003). Although E peaks occurred less frequently in Group 2, they had higher concentrations as compared to Group 1 (P < 0.003). Cumulative periovulatory and luteal phase E levels did not differ between the two groups. The reduction in P output in aged squirrel monkeys may interfere with proper implantation, thus contributing to the lower pregnancy rate observed in this age group. The decreased frequency of bio-LH surges indicated less frequent ovulations in the aged animals which may further impair reproduction with aging. © 1994 Wiley-Liss, Inc.     

Mating-related estradiol fluctuations during the estrous cycle of the Bolivian squirrel monkey (Saimiri boliviensis boliviensis)

The female squirrel monkey, Saimiri boliviensis, a New World monkey, has 10-day estrous cycles during the annual breeding season. Measurements of serum estradiol (E) concentrations in females housed with males in breeding pens revealed markedly higher levels than previously reported. Additionally, females in breeding pens appeared to have two distinct patterns of serum E peaks relative to the LH surge. Serum estrogen peaks averaging 5-fold greater than levels on the preceding day were observed on the same day as the LH surge, whereas other females had only a small E rise on the day of the LH surge followed by a 6-fold E rise on the next day. The serum progesterone (P) levels in all animals were depressed for 1-2 days before the LH surge but frequently started to rise on the day of the LH surge. The effect of the presence of a breeding male was examined by studying females housed in a group pen without exposure to a breeding male. In contrast to breeding-pen patterns, relatively small E rises were found in the 10 cycles observed. To further elucidate estrus-related E rises, a limited male-access paradigm was used to isolate mating-related hormone fluctuations. Pre-mating E levels had no marked rises; however, 4 h after mating, whether on the day of the LH surge or the next day, large E rises were found. These studies indicate that the LH surge in cycling squirrel monkeys is consistently preceded by a marked P nadir and associated with relatively small E rises.(ABSTRACT TRUNCATED AT 250 WORDS)     

Breeding and rearing squirrel monkeys (Saimiri sciureus) in captivity.

A breeding colony of squirrel monkeys (Saimiri sciureus) was established to provide animals for behavioral research concerned with early development. The origin of the initial breeders was Peru, Colombia, and Bolivia. During the past 10 years, the colony has grown to 125 adult females, 20 adult males, and 120 immature animals of various ages. The annual conception rate for the last 5 years averaged 68%. This resulted in 84% viable births of which 82% survived past 6 months of age. The majority of the births (65%) occurred during June-August, and 87% during May-September. The most efficient and successful breeding strategy was to form mixed-sexed groups of 10--15 females and 2--3 males before the mating season began and to maintain the integrity of these groups with minimal interference. Progeny were reared apart from their natural mother without difficulty, and their growth and development were found to be similar to those of mother-reared monkeys.     

Effect of age and breeding season on the developmental capacity of oocytes from unstimulated and follicle-stimulating hormone-stimulated rhesus monkeys

Effects of age and season on the developmental capacity of oocytes from unstimulated and FSH-stimulated rhesus monkeys were examined. Immature cumulus-oocyte complexes were matured in vitro in modified CMRL-1066 medium containing 20% bovine calf serum and subjected to in vitro fertilization followed by embryo culture. After fertilization, ova from unstimulated prepubertal monkeys displayed lower development to morula (4%) than those from unstimulated adult females (18% in breeding season and 22% in nonbreeding season). No developmental difference was found between ova from unstimulated adult monkeys in breeding and nonbreeding seasons. However, ova from FSH-primed prepubertal monkeys displayed greater development to blastocyst stage (54%) than those from adult monkeys in the breeding season (16%) and nonbreeding season (0%); and ova from FSH-primed adult females in the breeding season had significantly (P < 0.05) greater developmental competence than those obtained in the nonbreeding season (> or = morula stage, 54% vs. 3%; blastocyst stage, 16% vs. 0%). These data indicate that 1) rhesus monkey oocytes acquire developmental competence in a donor age-dependent manner, and 2) animal age and breeding season modulate the effect of FSH on oocyte developmental competence in the rhesus monkey.     

Age, social rank and lactational status influence ovulatory patterns in seasonally breeding rhesus monkeys

Rhesus monkeys housed outdoors exhibit a distinct breeding season limited to the fall and winter months. Four groups of female rhesus monkeys, multiparous nonlactating (MNL; n = 8), multiparous lactating (ML; n = 6), primiparous lactating (PL; n = 3) and nulliparous first-time ovulators (N; n = 6) were studied to investigate the influence of age, parity, and social dominance rank on the parameters of the breeding season. MNL exhibited the longest season (146 days), and PL the shortest (70 days), with N (106 days) and ML (89 days) intermediate. PL females also had a significantly reduced percentage of normal ovulations compared to other groups. Neither body weight nor estimates of body fat were related to either the timing of the ovulatory season or the quality of ovulations within the season. Parity and social dominance rank were significantly related to the percentage of normal ovulations (r = 0.85), with low-ranking, primiparous females exhibiting the fewest normal ovulations. These data indicate that the presence of a suckling infant acts synergistically with environmental factors to determine the parameters of the breeding season. Furthermore, postpubertal females may be more responsive to those factors that terminate the breeding season, and some factor independent of body weight but associated with low social dominance rank and/or primiparity renders females less capable of normal luteal function during the breeding season.     

Sex steroid hormone binding globulin levels and free 17 beta-estradiol and testosterone in cynomolgus monkeys during different reproductive states

The effect of differences in sex hormone binding globulin (SHBG) levels associated with reproductive status on free 17 beta-estradiol (E2) and testosterone (T) was characterized in cynomolgus monkeys. Cycling female cynomolgus monkeys had higher SHBG levels than males, pregnant and lactating females (P less than 0.05). Ovariectomized females were not different than control females, suggesting no hypoestrogenic effect. The percentage of free T was elevated in pregnant animals (P less than 0.05) compared to normal males and females, but the percentage of free E2 was similar between these groups. Although a gender difference in the percentages of free E2 and T was not detected, there was a gender difference in the free T and E2 concentrations due to endogenous secretion. Increased free E2 concentrations during pregnancy were the result of endogenous secretion rather than the decreased binding capacity of SHBG; the increased percentage of free T during pregnancy significantly increased free T concentrations. These data suggest that the gender difference in SHBG levels in cynomolgus monkeys is due to androgenic influences and that estrogens have minimal influence. Furthermore, the decrease in SHBG levels during pregnancy and lactation may not be entirely dependent on these androgenic influences.     

Effect of estradiol-treated females on all-female groups of rhesus monkeys during the transition between the nonbreeding and breeding seasons

Behavior, sex steroid levels and sex skin color were monitored in 2 out-door-housed all-female groups of rhesus monkeys during the nonbreeding season and into the breeding period. Each group contained gonadally intact and ovariectomized (OVX) females. In one group, 2 OVX females were implanted with estradiol benzoate pellets. Female-to-female sexual behavior, sex skin redness and endocrine indices of ovulation (in intact females) all increased as the breeding season approached. The two groups did not differ with regard to these measures. This lack of difference suggests that the presence of males may be required to mediate socially facilitated out-of-season breeding by females. No strong relationships were detected between steroid levels of intact females and sex skin color or female-to-female sexual behavior in all-female groups.     

The estrous cycle of captive woodchucks (Marmota monax)

Changes in vaginal cytology were assessed and correlated with temporal changes in circulating concentrations of progesterone (P) and estradiol-17 beta (E2) during the breeding season (February to March) in a seasonally breeding rodent, the woodchuck (Marmota monax). Ten individually caged adult females, maintained under laboratory conditions for 3-11 mo, were studied. Vaginal smears were taken each morning for 2 consecutive months beginning 1 February 1990. Seven of 10 females exhibited readily identifiable estrus, characterized by a clear predominance (83%) of cornified cells. The earliest estrous smear was recorded on 3 February and the latest on 12 March. These animals were monoestrous and remained in a prolonged estrous period during their brief breeding season. The average duration of estrus was 18.1 +/- 2.1 days, ranging from 12-27 days. Levels of P and E2 were determined in serum samples taken before, during, and after estrus from 7 females who exhibited estrus. No changes in the circulating levels of P were apparent during the estrous cycle. However, there was a consistent pattern of estradiol secretion characterized by elevated levels of E2 before and during estrus, followed by a significant (p less than 0.05) decline in E2 levels one week after the end of estrus. Elevated levels of E2 preceded and coincided with maximal degree of vaginal cornification. Thus, the termination, but not the onset, of estrus in woodchucks reflected closely the temporal pattern of changes in serum E2 levels during the breeding season.     

Menstrual cycle characteristics of seasonally breeding rhesus monkeys

Rhesus monkeys in seminatural environments exhibit a distinct seasonal mating cycle with conceptions restricted to the fall and winter months. In the present study, the characteristics of menstrual cycles were examined during a 1-year period in twelve rhesus monkeys in whom pregnancy was prevented. Menses occurred throughout the year, but ovulations were observed only in the fall and winter months. Menses in the spring and summer months occurred irregularly and were associated exclusively with anovulatory cycles. The total number of ovulations exhibited by these females during the breeding season ranged from two to six and was positively related to body weight, mean luteal phase progesterone (P) levels of normal cycles and social dominance rank. Ovulations with a short luteal phase were exhibited by four females (seven cycles), with the likelihood of occurrence increasing as the breeding season progressed. The incidence of abnormal cycles was predicted from the linear combination of parity, body weight and luteal phase P of normal ovulatory cycles. These results suggest that during the seasonally delimited period of ovulation, females exhibit a range in the quality and quantity of ovulations which may be predicted by certain idiosyncratic physical and behavioral traits.     

Seasonal effects on fertility and ovarian follicular growth and maturation in camels (Camelus dromedarius).

Camels are said to be seasonal breeders, but the extent to which season interferes with food supply to affect ovarian function is not fully documented. Hence, the three aims of this study were: (1) to define the breeding season of camels maintained in semi-arid conditions in southern Morocco; (2) to relate the proportion of females with active ovaries (i.e., with follicles > 5 mm), with ovulatory (11-17 mm) or cystic (> 18 mm) follicles to age and body conditions score; (3) to study the consequences of the interactions between age and body conditions score on the proportion of females ovulating and conceiving; and (4) to compare follicular maturation, using in vitro steroidogenesis by intact follicles as a marker during the transition into the breeding season (October) and peak breeding season (March). There was a clear breeding season in the two flocks studied, since over 80-90% of the matings occurred during the period from mid-November to mid-April. Collection of ovaries at slaughter (n = 238) demonstrated a significant seasonal effect on the proportion of females with active ovaries (increasing from 73.5% in October-December to 89% in January-May), but no changes in the proportion of females with ovulatory follicles. Lean females (BCS < 2.5) had a delayed initiation of ovarian function in October-December. In addition, the proportion of females with cystic follicles was also affected by season (peaking during April-May). Neither age nor body condition modulated the frequency of cysts. Finally, the proportion of females conceiving increased steadily as season progressed (peaking at 57% in April-May). Body condition score did not affect this proportion, but young females (< or = 5 years old) had a low ability to conceive. Morphological features of large follicles were unaffected by season. Ovulatory follicles contained around 10(7) granulosa and theca cells. In vitro testosterone output by intact follicles was unrelated to follicle size and season. In vitro oestradiol output increased with increasing follicle size and was larger in follicles obtained during peak breeding season than at its initiation. This may indicate that early breeding season follicles display a low aromatase activity in their granulosa cells. Whether the low oestradiol output of early breeding season follicles is resulting in the low fertility observed at this period remains to be determined.     

Androgen resistance in squirrel monkeys (Saimiri spp.)

The goal of this study was to understand the basis for high androgen levels in squirrel monkeys (Saimiri spp.). Mass spectrometry was used to analyze serum testosterone, androstenedione, and dihydrotestosterone of male squirrel monkeys during the nonbreeding (n = 7) and breeding (n = 10) seasons. All hormone levels were elevated compared with those of humans, even during the nonbreeding season; the highest levels occurred during the breeding season. The ratio of testosterone to dihydrotestosterone in squirrel monkeys is high during the breeding season compared to man. Squirrel monkeys may have high testosterone to compensate for inefficient metabolism to dihydrotestosterone. We also investigated whether squirrel monkeys have high androgens to compensate for low-activity androgen receptors (AR). The response to dihydrotestosterone in squirrel monkey cells transfected with AR and AR-responsive reporter plasmids was 4-fold, compared with 28-fold in human cells. This result was not due to overexpression of cellular FKBP51, which causes glucocorticoid and progestin resistance in squirrel monkeys, because overexpression of FKBP51 had no effect on dihydrotestosterone-stimulated reporter activity in a human fibroblast cell line. To test whether the inherently low levels of FKBP52 in squirrel monkeys contribute to androgen insensitivity, squirrel monkey cells were transfected with an AR expression plasmid, an AR-responsive reporter plasmid, and a plasmid expressing FKBP52. Expression of FKBP52 decreased the EC50 or increased the maximal response to dihydrotestosterone. Therefore, the high androgen levels in squirrel monkeys likely compensate for their relatively low 5 alpha-reductase activity during the breeding season and AR insensitivity resulting from low cellular levels of FKBP52.     

Thyroid status of female rhesus monkeys and preliminary information on impact of perchlorate administration

Thyroid status was assessed in adult female rhesus monkey breeders at the California National Primate Research Center at the beginning of the breeding season. The 95% confidence intervals for thyrotropin (TSH), thyroxine (T(4)) and triiodothyronine (T(3)) (n = 66-80) were similar to those previously reported in smaller samples of macaque monkeys. Based on human criteria, 10 of 80 monkeys (12%) were hypothyroid (TSH > 2.0 μIU/mL). Because hypothyroxinaemia can be a risk factor in pregnancy, T(4) status was compared with past breeding history, breeding outcome for that season and general health records in a subset of 42 breeders. Age, weight and parity did not differ between monkeys in the lowest T(4) quartile as compared with those in the upper three quartiles. However, T(4) concentrations were significantly associated with the number of missed menstrual cycles during the previous breeding season. In additional work, three healthy lactating rhesus monkeys were given three different doses of environmental contaminant and thyroid iodine uptake inhibitor, ammonium perchlorate (0.006, 0.34, 12.8 mg/kg/day, respectively) in food for two weeks. Thyroid status variables (TSH, T(4), T(3), thyroid radioactive iodine uptake) were then measured. In the monkey receiving the highest perchlorate dose, iodine uptake was suppressed relative to baseline. The study shows the availability of tools to study thyroid status in rhesus monkeys, the variability of thyroid status in the breeder colony and the potential ability of environmental factors to influence thyroid status.     

Effects of melatonin implants in pony mares. 1. Acute effects

The effects of melatonin implant treatment over a four week period on LH, estradiol (E2) and progesterone (P4) secretion during the breeding season were studied in ovary-intact and ovariectomized pony mares. Mares with melatonin implants had significantly higher daytime melatonin concentrations than mares with sharm implants (P = 0.0065). In ovariectomized mares, LH secretion did not differ between mares with melatonin and sham implants. In ovary-intact mares, melatonin implants altered the pattern of LH secretion (P = 0.0023) in such a way that an increase in LH secretion was observed during the periovulatory period. Estradiol and P4 secretion were unaffected by melatonin implants. These results suggest that constant administration of melatonin may enhance the secretion of LH during the periovulatory surge but does not adversely affect E2, P4 or basal LH secretion in mares during the breeding season.     

Territorial aggression and hormones during the non-breeding season in a tropical bird

The hormonal control of territorial aggression in male and female vertebrates outside the breeding season is still unresolved. Most vertebrates have regressed gonads when not breeding and do not secrete high levels of sex steroids. However, recent studies implicate estrogens in the regulation of non-breeding territoriality in some bird species. One possible source of steroids during the non-breeding season could be the adrenal glands that are known to produce sex steroid precursors such as dehydroepiandrosterone (DHEA). We studied tropical, year-round territorial spotted antbirds (Hylophylax n. naevioides) and asked (1). whether both males and females are aggressive in the non-breeding season and (2). whether DHEA is detectable in the plasma at that time. We conducted simulated territorial intrusions (STIs) with live decoys to male and female free-living spotted antbirds in central Panama. Non-breeding males and females displayed robust aggressive responses to STIs, and responded more intensely to decoys of their own sex. In both sexes, plasma DHEA concentrations were detectable and higher than levels of testosterone (T) and 17beta-estradiol (E(2)). In males, plasma DHEA concentrations were positively correlated with STI duration. Next, we conducted STIs in captive non-breeding birds. Captive males and females displayed robust aggressive behavior. Plasma DHEA concentrations were detectable in both sexes, whereas T was non-detectable (E(2) was not measured). Plasma DHEA concentrations of males were positively correlated with aggressive vocalizations and appeared to increase with longer STI durations. We conclude that male and female spotted antbirds can produce DHEA during the non-breeding season and DHEA may serve as a precursor of sex steroids for the regulation of year-round territorial behavior in both sexes.     

Long-term monitoring of fecal steroid hormones in female snow leopards (Panthera uncia) during pregnancy or pseudopregnancy

Knowledge of the basic reproductive physiology of snow leopards is required urgently in order to develop a suitable management conditions under captivity. In this study, the long-term monitoring of concentrations of three steroid hormones in fecal matter of three female snow leopards was performed using enzyme immunoassays: (1) estradiol-17β, (2) progesterone and (3) cortisol metabolite. Two of the female animals were housed with a male during the winter breeding season, and copulated around the day the estradiol-17β metabolite peaked subsequently becoming pregnant. The other female was treated in two different ways: (1) first housed with a male in all year round and then (2) in the winter season only. She did not mate with him on the first occasion, but did so latter around when estradiol-17β metabolite peaked, and became pseudopregnant. During pregnancy, progesterone metabolite concentrations increased for 92 or 94 days, with this period being approximately twice as long as in the pseudopregnant case (31, 42, 49 and 53 days). The levels of cortisol metabolite in the pseudopregnant female (1.35 μg/g) were significantly higher than in the pregnant females (0.33 and 0.24 μg/g) (P<0.05). Similarly, during the breeding season, the levels of estradiol-17β metabolite in the pseudopregnant female (2.18 μg/g) were significantly higher than those in the pregnant females (0.81 and 0.85 μg/g) (P<0.05). Unlike cortisol the average levels of estradiol-17β during the breeding season were independent of reproductive success.The hormone levels may also be related to housing conditions and the resulting reproductive success in female leopards. The female housed with a male during the non-breeding season had high levels of cortisol metabolites and low levels of estradiol-17β in the breeding season, and failed to become pregnant. This indicates that housing conditions in snow leopards may be an important factor for normal endocrine secretion and resulting breeding success.     

Sexual competition in an inbreeding social spider,Stegodyphus dumicola (Araneae: Eresidae)

Summary Sexual competition is shown to occur in the social spiderStegodyphus dumicola (Eresidae). While the secondary sex ratio inS. dumicola was female-biased, the overall operational sex ratio (numbers of breeding males to breeding females over the season) showed no strong female bias. Males matured before females and had a shorter lifespan than the females. Mating took place in the natal colony. Males fought over access to the few mature females available early in the reproductive season, but females appeared to control the duration of mating. Later in the season, some adults of both sexes dispersed alone to breed elsewhere. We conclude that different rates of maturation between the sexes within a colony provide the opportunity for females that mature early in the season to be choosy in selecting a mate and this forces males to compete. Early reproduction may be beneficial for both females and males, because the offspring of females that reproduce early may have a competitive advantage over later (and smaller) offspring in the colony.     

Sex, age, experience and condition as factors affecting arrival date in prospecting common terns, Sterna hirundo

Several studies have shown that seabird colonies consist to a large extent of young nonbreeders (prospectors). These individuals appear at the colony later in the season than established breeders. The reasons for this late arrival have remained unclear in most cases, mainly because of technical difficulties in collecting sufficient data from nonbreeding individuals. We used a novel transponder system to identify remotely the members of a common tern colony, including nonbreeders, during eight breeding seasons and we combined the system with automatic balances. Ninety-two per cent of prospectors returned for the first time when 2 years old and 88.9% of recruits to the breeding population had spent at least one previous season at the colony as prospectors. In both sexes, most individuals prospected for one season, but more males than females prospected for more than one season, although a higher proportion of females started breeding without a previous prospecting phase. Terns arrived earlier in the season the older they were and the more experience of the colony they had, but experience proved to be more important than age. Prospectors gained about 3 weeks with a previous prospecting season whereas an additional year of age allowed birds to arrive only about 6 days earlier. Prospectors returning later in the season arrived with lower body masses. Males on average arrived earlier at the colony than females. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Effect of ovariectomy at two periods of the year on LH and FSH basal concentrations and pituitary response to LHRH in the brown hare (Lepus europaeus)

In the brown hare, fertile mating takes place from the beginning of December to September. Seasonal variations of basal concentrations of LH and FSH, and pituitary response to a monthly i.v. injection of LHRH were studied in intact control females and in females ovariectomized during the seasonal anoestrus (OVX1) or during the breeding season (OVX2). In intact females, both basal and LHRH-stimulated LH levels showed an annual variation, with minimal values during anoestrus. During the breeding season, the LH response to LHRH exhibited a biphasic pattern. In contrast, there was no clear seasonal variation in basal and LHRH-stimulated FSH concentrations. After ovariectomy during anoestrus, basal LH remained low for 2 months and began to increase in December. After ovariectomy during the breeding season, LH basal concentrations increased within a few days after the operation. Thereafter, LH values remained high in both groups of females until September, and decreased significantly as in intact females. The pattern of LH release after LHRH remained monophasic in the two groups of ovariectomized females. In OVX1 females, the LH response increased as early as October, was maximum from December to April and decreased progressively until October. IN OVX2 females, the LH response decreased regularly after ovariectomy to a minimum in October. In the 2 groups of ovariectomized females, basal FSH concentrations and pituitary response to LHRH rose rapidly after ovariectomy and did not vary significantly thereafter. These results showed a direct central effect of season on the regulation of basal concentrations of LH, modulated by a negative feed-back of ovarian secretions during the breeding season. In intact hares, the enhanced LH response after LHRH during the breeding season was related to an acute positive effect of ovarian secretions. The regulation of FSH was less dependent on season and remained under a negative control of the ovary throughout the year.