Effects of ventromedial nucleus lesions on estradiol induced ovulation in the rat

Small bilateral stereotaxic lesions were made in the hypothalamic ventromedial nucleus (SVMN) to determine: (i) whether estrogen would restore early receptivity in unreceptive SVMN lesioned female rats and (ii) whether SVMN lesions would suppress estrogen induced ovulation in the rat. SVMN lesions were shown to completely suppress spontaneous early receptivity and seriously impair estrous receptivity in 5-day cyclic female Wistar rats. A loss of early receptivity in response to 10 μg estradiol benzoate (EB) was also observed in SVMN lesioned females, in comparison to unoperated, sham VMN and dorsomedial nucleus (DMN) lesioned animals. Isolated SVMN lesioned females exhibited a weak ovulatory response to 10 μg EB, but, where shown to be unreceptive prior to estrogen injection, they never ovulated. On the contrary, ovulation occured in about 50% of cases in isolated unoperated and in sham VMN and DMN lesioned females following estrogen administration. The mechanisms whereby EB brought about precocious ovulation in 5-day cyclic female rats were therefore concluded to be dependent on VMN functional integrity and thereby on the degree of early sexual receptivity in the rat.     

The role of pheromones in the sexual behaviour of the goldfish

Male goldfish (Carassius auratus) discriminate between ovulated and unovulated females on the basis of chemical cues. Goldfish males made temporarily anosmic by occlusion of the nares, or permanently anosmic by section of the olfactory nerves showed a marked decrease in sexual behaviour. Preference tests in a Y-maze indicate that the pheromone is present in the ovarian fluid of an ovulated female. It is concluded that shortly after ovulation the female goldfish releases a pheromone from the ovary which attracts the male and elicits the persistent courtship that accompanies spawning.     

Olfactory bulbectomy blocks mating-induced ovulation in musk shrews (Suncus murinus)

In many species, reproductive function can be modified by olfactory inputs. We employed bilateral olfactory bulbectomy (BULBX) to examine the effects of disruption of olfactory inputs on mating behavior and ovulation in female musk shrews. On several measures, sexual behavior was delayed in BULBX females compared to controls. When females were mated on five consecutive days, the majority of unoperated and sham-operated (SHAM) shrews ovulated; only one female subjected to BULBX ovulated. Administration of GnRH induced ovulation in the majority of females. We performed immunocytochemistry to assess the effects of bulbectomy on mating-induced responses of the neural GnRH system. In BULBX and SHAM females, the numbers of cells containing proGnRH immunoreactivity in the medial septum (MS)/diagonal band (DB) were significantly elevated 1 h after mating. Bulbectomy increased the numbers of GnRH-immunoreactive peptide-containing cells in the preoptic area, but it reduced neuron numbers in the MS/DB, as compared with those in SHAM controls. In addition, the GnRH-immunoreactive fiber area in the median eminence was greater in BULBX than in SHAM females. In sum, female musk shrews can display receptivity and engage in copulation without olfactory inputs. However, the olfactory system is essential for mating-induced ovulation.     

Adrenalectomy: Effects on mating behavior,ovulation, and early gestation in the cycling rat

Adrenalectomy was performed 4 weeks prior to evaluating the effects on mating behavior, ovulation, and the first 10 days of gestation in 4-day cycling female rats. Mating behavior was essentially normal and occurred at times similar to sham-adrenalectomized animals on the afternoon of predicted proestrus. The number of ova ovulated was significantly reduced in mated adrenalectomized subjects in contrast to data reported by Rodgers (1971) for intact females mated at similar times in proestrus. Embryonic swellings were significantly fewer at day 10 of gestation than in the sham-adrenalectomized group providing additional evidence of reduced numbers of ova ovulated. Embryonic weights at day 10 of gestation were not affected by adrenalectomy. Removal of adrenocortical hormones significantly increased uterine weight on the morning of vaginal estrus. The data fail to show a significant delay in sexual receptivity as a result of removing adrenal progesterone, but suggest that adrenalectomy interferes with the neuroendocrine reflex mediating ovulation.     

Prostaglandin induction of spawning behavior in Cichlasoma bimaculatum (Pisces cichlidae)

Prostaglandin (PG) stimulates female spawning behavior in goldfish and in some other teleosts in which female reproductive behaviors consist of postovulatory oviposition acts. This study examined the effects of PG on female sexual behavior in a teleost fish, Cichlasoma bimaculatum, in which female reproductive behaviors involve both preovulatory courtship and substrate cleaning behaviors, and post-ovulatory oviposition behavior. In females of established pairs, PGF2 alpha injection (5 micrograms, im) at any stage of the spawning cycle, or in the parental phase, rapidly induced substrate cleaning which soon merged into oviposition behavior (without egg release). These results support a role for PG in oviposition behavior of Cichlasoma. However, indomethacin (1 mg, ip), a PG synthesis inhibitor, did not block oviposition in ovulated females which had begun to spawn. Indomethacin may not have lowered PG levels sufficiently. Alternatively, as shown by J.J. Polder (1971, Neth. J. Zool. 21, 265), oviposition behavior may be induced or maintained by other factors associated with the spawning situation.     

Androgen Induction of Male Sexual Behaviors in Female Goldfish

The effectiveness of testosterone (T) and 11-ketotestosterone (K) in inducing male-typical sex behaviors in goldfish was examined by implanting intact adult females with one empty (blank) Silastic implant (B females), one implant containing T or K, or one T and one K implant (T + K females). Behavior of the four female groups was compared to that of untreated males and males containing a blank implant. Male-typical behaviors (coutship, spawning) and associated behavioral changes (increased activity, reduced spontaneous feeding) were assessed 3.5 and 4.5 months after implant in 30-min tests in which the test female or male was allowed to interact with a stimulus female in which sexual receptivity and attractivity had been induced by acute prostaglandin F_2αinjection. Prostaglandin-induced female-typical spawning behavior in the test females and males was also assessed 4.5 months after implant in a 60-min test for female-typical behavior in which the test fish was injected with prostaglandin and placed immediately with a sexually active male. Blood samples 5 months postimplant showed that implants generated physiological levels of T and K. In both tests for male-typical behaviors, K and T + K females exhibited the full suite of behaviors shown by spawning males, e.g., male-typical courtship and spawning, increased swimming activity, and reduced spontaneous feeding. Although behaviors of K and T + K females did not differ, those of T + K females were more often equivalent to those of males and significantly different from those of B females. T females exhibited marginal male-typical behaviors which never differed significantly from those of B females. Androgen-treated females exhibited female-typical; spawning behaviors equivalent to that of males and B females. The results show that adult female goldfish can be behaviorally masculinzed without behavioral defeminization, and suggest that male-typical sex behaviors in goldfish are dependent on K, although other steroids also may be required. The inducible behavioral bisexuality of goldfish, a gonochoristic species, is discussed in terms of the prevalence of hermaphroditism in teleosts.     

Importance of intense male sexual behavior for inducing the preovulatory LH surge and ovulation in seasonally anovulatory female goats

The present study was carried out to determine whether the presence of photostimulated sedated male goats could stimulate the LH preovulatory surge and ovulation in seasonal anestrous goats. Sexually experienced male goats were treated with artificial long days (16 hours light per day) from 1 November to 15 January to stimulate their sexual activity in March and April, corresponding to the natural sexual rest. A female group of goats (n = 20) was exposed to non-sedated males who displayed an intense sexual behavior and provided strong odor (non-sedated group). Another female group of goats (n = 20) was exposed to the photo-stimulated male goats, but these males were sedated with Xylazine 2% to prevent the expression of sexual behavior (sedated group). The sedated males also provided a strong odor. Females of both groups had full physical and visual contact with non-sedated or sedated males. In both groups, the males remained with females during 4 days. The LH preovulatory surge of 10 female goats per group was measured by determination of LH plasma concentrations in samples taken every 3 hours. In addition, in all goats, (n = 20 by group), ovulation was determined by measuring plasma concentrations of progesterone. The proportion of female goats showing a preovulatory LH surge was higher in goats exposed to non-sedated (10/10) than in those exposed to sedated bucks (0/10; P < 0.0001). Similarly, most of does in contact with non-sedated males ovulated (19/20), but none of those in contact with sedated males did so (0/20; P < 0.0001). We conclude that the expression of an intense sexual behavior by male goats is necessary to induce LH preovulatory surge and ovulation in seasonally anovulatory goats.     

Effects of indomethacin and prostaglandins on the spawning behaviour of female goldfish

In goldfish, injection of ovulated eggs (from donor females) through the ovipore and into the ovarian lumen of females with vitellogenic oocytes induces spawning behaviour within several hours. Intraperitoneal (i.p.) injection of indomethacin (IM), 10 μg/g, either 10 h prior to, or coincident with, injection of ovulated eggs, completely inhibits the onset of spawning behaviour. IM injection similarly terminates ongoing spawning behaviour induced by egg injection. PGF_2α (5 μg/g; i.p. injection) restores spawning behaviour of egg-injected, IM-blocked fish; at the same dosage, PGE₂ is marginally effective and PGE₁ is without effect. As PGF_2α and PGE₂ also induce spawning behaviour in females which have not been injected with ovulated eggs, it is suggested that distension of the oviduct following ovulation or egg injection results in the release of PG which then acts in some way to induce spawning behaviour. The ability of PG to induce spawning behaviour is eliminated by hypophysectomy and restored by treatment with salmon gonadotropin: no steroid treatment was effective in restoring PG-induced spawning in fish which had been hypophysectomized for 3–4 months. The possible mode of action of PG in inducing spawning behaviour in female goldfish is discussed.     

Prostaglandin E2 induces receptive behaviors in female Xenopus laevis

The object of this study was to examine the effects of exogenous and endogenous prostaglandin E2 (PGE2) on the sexual behavior of female South African clawed frogs, Xenopus laevis. Ticking and leg extension, which communicate sexual unreceptivity to males, were studied in intact, ovariectomized, and ovariectomized-oviductectomized females. The onset of the PGE2 behavioral effect occurs within 30 sec to 3 min of injection for intact and ovariectomized females; for ovariectomized-oviductectomized females, the latency period for the effect ranges from 10-20 min. PGE2 induced receptivity in doses as low as 0.03 microgram/frog. Injection of the prostaglandin synthesis inhibitors, indomethacin and flurbiprofen (FBP), blocked chorionic gonadotropin- (HCG-) induced behavioral receptivity, suggesting that endogenous prostaglandin synthesis may have a role in regulating female sexual behavior. Flurbiprofen blockade of HCG-induced receptivity was reversed by PGE2 administration, suggesting that FBP's effects are PG synthesis-specific.     

Egg-oviduct interaction initiates reproductive behavior

The experiments reported in this paper provide evidence that eggs must pass through the oviducts in order for receptivity to occur after ovulation in the female frog, Rana pipiens. In one experiment, oviductectomized frogs remained unreceptive after ovulation was induced by administration of exogenous pituitary glands, while sham-operates became receptive within 48 hr. Another experiment had four groups of subjects: ovariectomized females, females with oviducts ligated at the ostial end, females with openings in the uteri that prevented eggs from accumulating there, and sham-operated females. Only the last two groups, groups in which eggs could pass through the oviducts, became receptive. In these experiments, receptivity was indicated by absence of the release call during manual clasping of the trunk. Earlier experiments have shown that eggs have to pass through the oviducts in order to become fertilizable. Thus, the passage of eggs through the oviducts provides a mechanism which links the onset of reproductive behavior to the availability of fertilizable gametes.     

Behavioral correlates to gonadal state in the lizard, Anolis carolinensis

Female Anolis carolinensis of unknown reproductive condition were introduced into cages containing a sexually active male and the resulting sexual behavior was noted. Female sexual receptivity was found to be correlated with the stages of follicular maturation. These results suggest that sexual receptivity in female A. carolinensis is restricted to a time immediately preceding the ovulation of each follicle and is dependent upon hormonal conditions normally arising during follicular development. Evidence is offered to suggest that during the breeding season, female sexual receptivity is rhythmical and contingent upon the cyclic ovarian changes peculiar to lizards of this genus.     

Comparison of the penetrability of the egg vestments in follicular oocytes, unfertilized and fertilized ova of the rabbit

Mixed populations of rabbit ovulated eggs and follicular oocytes, one labelled with a fluorescent marker, were transferred to the same tubal ampulla of an inseminated recipient female and were then recovered 3 hr later. There was no significant difference in the number spermatozoa penetrating to the perivitelline space or within the substance of the zona pellucida of follicular oocytes (immature or atretic) and mature ovulated ova. In contrast to mature ovulated ova, however, none of the spermatozoa reaching the perivitelline space of vesicular (dictyate) oocytes had attached to or penetrated the vitelline surface to enter the ooplasm.The same approach involving transfer of nonpenetrated eggs together with eggs penetrated previously in a donor female, demonstrated that prior entry of spermatozoa does not reduce the penetrability or receptivity of the rabbit zona pellucida to subsequent spermatozoa.These experiments indicate: (a) that the penetrability of the granulosa cell investment and/or zona pellucida of the rabbit follicular oocyte does not change from the time of antrum formation until the point at which follicular atresia ensures; (b) that between the time of initial LH stimulation and ovulation important changes mediating the onset of the fertizability of the dictyate oocyte of the rabbit probably occur at the vitelline surface; and (c) that in neither a qualitative nor quantitative sense has the demonstrably greater resistance of the rabbit zona pellucida to proteolysis following fertilization any physiological significance for sperm penetration.     

Spontaneous and gonadotropin-induced ovulation in the goldfish, Carassius auratus L: effects of external factors

Spontaneous ovulation in goldfish is synchronized with photoperiod and influenced by water temperature and aquatic vegetation. As the latency to ovulation from injection of HCG is highly temperature dependent, the finding that ovulation occurs at approximately the same time of day at temperatures from 12° to 26° C suggests the time of the endogenous preovulatory gonadotropin surge may change with the temperature. The time of spontaneous ovulation adjusts to a reversed light:dark cycle within 2 weeks; some 4–6 h shifts in a single light:dark cycle modify the time of ovulation.
Few sexually mature females kept under long photoperiod (16L:8D) and transferred from cold (13 ± 1°C) to warm (21 ± 1°C) water ovulated spontaneously. Exposure to artificial aquatic vegetation for as little as one light phase significantly increases the proportion offish ovulating in warm water. Fish kept in cold water without vegetation do not ovulate; the addition of vegetation induces ovulation, although the response latency is longer than in warm water. Aquatic vegetation may be an effective stimulus for ovulation in other teleosts which spawn on this substrate.     

New data on ovulatory action of estrogens during 5 day cycles in rats

A peculiar laboratory strain W1 of Wistar rats has 5 day cycles and can be made precociously receptive to a male by an injection of estradiol 10 mcg sc at 1500 of Cycle Day 2 (Diestrus III). In the course of repeating previous work the authors noted that receptivity increased from 25 to 63%, and they investigated the mechanism by checking ovaries of estrogen treated females histologically for ovulation, in comparison with ovaries of females exposed to males during the night of Diestrus III. In this experiment 56% of estrogen treated rats ovulated. 41 out of 60 (68%) of the paired rats accepted a male (verified by sperm in vaginal smear), and 93% of these ovulated. Of the 19 who were unreceptive, 1 (5.3%) ovulated (p.001). Thus estrogen treatment is much more effective than previously observed; 5 day cycling rats have a related precocity of sexual receptivity and capacity to ovulate under this stimulus.     

Female sexual arousal in amphibians

Rather than being a static, species specific trait, reproductive behavior in female amphibians is variable within an individual during the breeding season when females are capable of reproductive activity. Changes in receptivity coincide with changes in circulating estrogen. Estrogen is highest at the point when females are ready to choose a male and lay eggs. At this time female receptivity (her probability of responding to a male vocal signal) is highest and her selectivity among conspecific calls (measured by her probability of responding to a degraded or otherwise usually unattractive male signal) is lowest. These changes occur even though females retain the ability to discriminate different acoustic characteristics of various conspecific calls. After releasing her eggs, female amphibians quickly become less receptive and more choosy in terms of their responses to male sexual advertisement signals. Male vocal signals stimulate both behavior and estrogen changes in amphibian females making mating more probable. The changes in female reproductive behavior are the same as those generally accepted as indicative of a change in female sexual arousal leading to copulation. They are situationally triggered, gated by interactions with males, and decline with the consummation of sexual reproduction with a chosen male. The changes can be triggered by either internal physiological state or by the presence of stimuli presented by males, and the same stimuli change both behavior and physiological (endocrine) state in such a way as to make acceptance of a male more likely. Thus amphibian females demonstrate many of the same general characteristics of changing female sexual state that in mammals indicate sexual arousal.     

Induction of developmental abnormalities in larval goldfish, Carassius auratus L., under cool incubation conditions

Compared with incubation at a constant 22° C, exposure of goldfish embryos and larvae to 13° C, under a variety of thermal protocols, caused increased frequencies of abnormal development and, in some cases, reduced survival to hatching. The low-temperature incubation conditions were particularly deleterious when eggs were incubated at 13° C from the outset, regardless of the temperature at which the donor female ovulated and the eggs were fertilized. Significantly higher frequencies of developmental abnormalities were also noted when embryos were transferred from 22°C to 13°C at 6, 24, 128 and, in one case, 175 h after fertilization. In three of five experiments, subjecting embryos and larvae to diel fluctuations between 22 and 13° C, with a 5-h hold at the lower temperature, caused an increase in development abnormalities. These results demonstrate that the thermal requirements of goldfish embryos and larvae necessitate a delay in ovulation and spawning until water is sufficiently warm. Developmental abnormalities can be induced by exposure to cool (13° C) conditions, at least up to the time that swimbladder inflation occurs.     

F prostaglandins function as potent olfactory stimulants that comprise the postovulatory female sex pheromone in goldfish

This study establishes that ovulated female goldfish release F type prostaglandins (PGFs) to the water where they stimulate male spawning behavior and comprise the goldfish postovulatory pheromone. We first demonstrated that ovulated and prostaglandin-injected female goldfish release immunoreactive PGFs to the water. Next, using electro-olfactogram recording (EOG), we determined that waterborne prostaglandins function as potent olfactory stimulants for mature male goldfish. Prostaglandin F2 alpha (PGF2 alpha) and its metabolite 15-keto-prostaglandin F2 alpha (15K-PGF2 alpha) were the most potent prostaglandins; the former had a detection threshold of 10(-10) M and the latter a detection threshold of 10(-12) M. Studies of prostaglandin-injected fish indicated that PGF metabolites are an important component of the pheromone. Cross-adaptation experiments using the EOG demonstrated that goldfish have separate olfactory receptor sites for PGF2 alpha and 15K-PGF2 alpha that are independent from those that detect other olfactory stimulants. Finally, we established that male goldfish exposed to low concentrations of waterborne PGFs exhibit reproductive behaviors similar to those elicited by exposure to the odor of ovulated fish. Together with our recent discovery that a steroidal maturational hormone functions as a preovulatory "priming" pheromone for goldfish, these findings suggest that hormones and their metabolites may commonly serve as reproductive pheromones in fish.     

Male stimulation of luteinizing hormone surge, progesterone secretion and ovulation in spontaneously persistent-estrous, aging rats

In aging, persistently estrous (PE) female rats, there are no estrous cycles or cyclic increases in luteinizing hormone (LH) secretion, but the sexual receptivity to the male is consistently maintained. We recently reported that caging and mating with fertile males elicits an LH surge followed by ovulation in aging PE rats. The present study examined the relationship between the LH surge, the increase in progesterone (P) secretion and ovulation in PE females exposed to males, and assessed whether intromission was essential for the male-induced pre-ovulatory LH surge. PE rats were implanted with intra-atrial cannulae. Six to eight days later, these females were individually caged with a fertile male and repeatedly sampled (once every 30 or 60 min) between 1400 and 1900 h for assays of plasma LH and P. Sexual behavior of the female was recorded and correlated with the changes in plasma LH and P values. Similar experiments were also performed on cannulated PE rats with their vaginal orifice blocked with adhesive tape during the caging and sampling session. In both experiments, over 90% of the PE females displayed a high degree of lordosis response to mounting by the male, and over 60% of those sexually receptive PE females exhibited an LH surge followed by ovulation. The male-induced preovulatory LH surge occurred in PE females without actual intromission. Caging with fertile males also elicited a marked increase in plasma P concentrations in PE rats and in PE females prevented from experiencing intromission.(ABSTRACT TRUNCATED AT 250 WORDS)     

Ovulatory rhythms and over-ripening of eggs in cultivated turbot, Scophthalmus maximus L.

The aim of this study was to investigate ageing of ovulated eggs retained in the ovarian lumen of captive turbot, Scophthalmus maximus L., prior to handstripping. The ovulation times of egg-batches were determined by catheterization, handstripping and plotting the percentage fertilizations and hatches of resulting eggs against time. The catheterization experiments showed that eggs age and change appearance rapidly after ovulation. The stripping method demonstrated that freshly-ovulated eggs showed greater than 90% fertilization and up to 97% hatch, but ovulated eggs retained in the ovary lumen for 1 day before stripping gave 0% hatch. The turbot showed precise ovulatory rhythms. The time between ovulations varied between individuals but was constant for any one female, making it possible to predict future ovulation times. Three of the females studied had ovulatory periods of alternating length.