Receptivity of female Neohelice granulata (Brachyura, Varunidae): different strategies to maximize their reproductive success in contrasting habitats

The extent of the receptive period may determine the mating strategies employed by female crabs to obtain mates. Here, we studied the receptivity of female Neohelice granulata (Dana, 1851) in the laboratory, including the form of the vulvae and the anatomy of the seminal receptacle (SR). We examined the factors that influence the duration of receptivity by comparing two populations inhabiting contrasting habitats: Mar Chiquita Coastal lagoon (MCL), which is an oligo-polyhaline estuary, and San Antonio Oeste (SAO), which is an eu-hyperhaline marine bay. Non-receptive females have immobile vulva opercula, while receptive females have mobile opercula. Histological sections of the SR showed that the degree of epithelium secretions was associated with the receptive stage of females, and they may be involved in the maintenance of viable sperm and in the dehiscence of spermatophores. The existence of a special tissue at the junction of the oviduct and the SR was described and proposed as an internal mechanism influencing the timing of ovulation. The duration of receptivity was dependent on the SR load and the capacity to lay eggs. Thus, females with empty SR exhibited longer receptivity and did not lay eggs, while those with full SR exhibited shorter receptivity and always laid eggs. Interpopulation differences showed that females from SAO had shorter receptivity and heavier SR and laid eggs more frequently than females from MCL. Based on our results, we suggest that N. granulata females can adjust the duration of their receptivity and control the moment of fertilization according to different internal mechanisms related to the morphology of the vulvae, the fullness of the SR and anatomical attributes of the SR. An important consequence of this control is greater sperm competition. The extent of the receptive period and the number of times that a female could become receptive in a single reproductive season may also depend on the habitat characteristics.     

Receptivity of female remating and sperm number in the sperm storage organ in the bean bug,Riptortus clavatus (Heteroptera: Alydidae)

This study examines the relationship between the number of sperm in the seminal receptacle (spermatheca) and the receptivity of female remating in the bean bugRiptortus clavatus Thunberg. On the 21 st day after the first mating when receptivity to remating was > 70%, females receptive to remating had significantly fewer sperm ( < 40 on average) in the spermathecae than females reluctant to do (about 150 on average). However, averages of the number of eggs laid by receptive and reluctant females within 21 days were almost same. The proportion of fertilized eggs for receptive females at 15–21 days after copulation was significantly lower than that for reluctant females. Spermatozoa transferred from a male to a female’s spermatheca were detected 5 min after copulation and then increased continuously to about 500 with the first hour. When copulation durations were manipulated artificially, the shorter the copulation period (=females had less sperm in their spermathecae), the higher the remating rate became. Females may perceive the number of sperm in their seminal receptacles and then determine whether they copulate or not. These results support the hypothesis that females mate multiply in order to replenish inadequate sperm supplies to fertilize all eggs produced.     

Ovulation and fertilization in the vole, Pitymys subterraneus

The females of Pitymys subterraneus bred in laboratory conditions have an irregular sexual cycle and induced ovulation. The first freshly ovulated eggs, surrounded by dense cumulus cells, appear in oviducts 10 h after copulation. Administering exogenous gonadotropins: pregnant mare's serum (PMS), human chorionic gonadotropin (hCG) or luteinizing hormone releasing hormone (LHRH), also induces ovulation in mature females of Pitymys subterraneus. In these experimental conditions females ovulate a similar number of eggs as after copulation. Dual stimulation (PMS and hCG plus copulation) does not result in the ovulation of a large number of normal ova, however, it does cause a release of degenerated oocytes.     

Factors influencing the onset and duration of receptivity of female purple rock crabs, Hemigrapsus sexdentatus (H. Milne Edwards, 1837) (Brachyura: Grapsidae)

The effects of males, field, and laboratory conditions on the receptivity of females were tested in the New Zealand purple rock crab Hemigrapsus sexdentatus. Onset and duration of female receptivity is of interest because it influences the time available for mating and therefore the operational sex ratio (OSR), male-male competition, and the extent of sperm competition. Females were receptive once a year for a short time prior to oviposition. The breeding season was highly synchronised and lasted for about 3 weeks (from the end of March to mid-April; southern autumn), after which, almost all females carried eggs. We found few receptive females in the field (0% to 4.9%) during the breeding season despite a large number of crabs examined (935 in 1999 and 555 in 2000), suggesting that females are receptive for less than a day. The onset of the breeding season was the same for the wild crabs and those held in field cages, but the duration of receptivity increased to several days for caged females. The onset of the breeding season of females in the laboratory was earlier compared to females in the field and had, overall, a longer breeding season. Females isolated from males stayed receptive significantly longer (5.5 days) than females caged with males (3.3 days), suggesting that the duration of female receptivity is adjusted according to the presence or absence of males. Our results suggest that females have some control over their receptivity in relation to male presence, and this could influence the outcome of sexual selection.     

Prior oviposition, female receptivity and last-male sperm precedence in the cosmopolitan pest Callosobruchus maculatus (Coleoptera: Bruchidae)

Sperm competition studies have shown that P₂ (the proportion of ova fertilized by the last male to mate) increases as the interval between inseminations is experimentally increased. Variation in the number of sperm in storage is associated with sperm use (or loss) from the female's sperm stores between copulations (fewer sperm from previous mates at the time of the last copulation) and with the extent of prior oviposition and female receptivity to further copulation: females that lay many eggs tend to have few remaining sperm in storage and to be more receptive to further copulation. Using the bruchid beetle Callosobruchus maculatus, we examined the effect of prior oviposition and female receptivity to further copulation on the extent of last-male sperm precedence (measured as P₂). Extent of prior oviposition was experimentally manipulated independently of the intermating interval by altering the availability of oviposition sites between inseminations. Females given few or no oviposition sites laid fewer eggs, were less receptive and had a lower P₂ than females given abundant oviposition sites. To examine the effect of female receptivity on P₂ independently of prior oviposition, we examined the outcome of sperm competition experiments using (1) females from lines that had been selected for different latencies to copulation and (2) natural variation in female latency to receptivity. Female receptivity to further copulation had no detectable effect on P₂. When oviposition resource is abundant, female receptivity may be a poor predictor of current sperm load.     

Stimulus requirements for pregnancy initiation in the golden hamster (Mesocricetus auratus) change with time of mating during the receptive period

When maintained under a 14L:10D photoperiod, the duration of behavioural receptivity in female golden hamsters was about 18-21 h depending on age and/or parity. The effectiveness of mating stimuli in initiating pregnancy was shown to be a function of when in the receptive period (early, middle, late) that mating occurred. During the 9-h period before ovulation, 5 ejaculatory series were sufficient to produce a nearly 100% pregnancy rate and maximum litter size. During the ovulation period, however, high pregnancy rates were achieved only when mating continued to satiety (12-15 ejaculatory series plus 10-24 long intromissions). Late in the receptive period even mating to satiety failed to result in a pregnancy. In general, pregnancy rates were significantly higher for young virgin than for older multiparous females when mating occurred during or after the ovulation period. The reduced fecundity of females mating during or after ovulation was due to insufficient vaginocervical stimulation to induce functional luteal activity and not to lack of spermatozoa. Females mating late in the receptive period did not show a cessation of oestrous cycles which characteristically follows the induction of a luteal phase. Greater amounts of vaginocervical stimulation during this time increased the number of females which delivered litters but had no significant effect on litter size. These results suggest that levels of male copulatory behaviour considered 'excessive' when mating occurs early in the receptive period are essential for pregnancy initiation when mating occurs later.     

Relationship between protein ingestion and sexual receptivity in females of the Australian sheep blowfly Lucilia cuprina

The receptivity of females of Luciliu cuprina to mating attempts is increased by protein feeding. Females became maximally receptive after consuming a quantity of protein which was insufficient to allow full ovarian development in any individual, and which was 25 % of that needed to produce full ovarian development in most flies. Receptivity increased progressively over 3 days following a protein meal taken on the day after emergence, but was almost maximal within 24 h when females were given a protein meal on the fifth day after emergence. Topical application of the insect growth regulator Altosid caused a marked increase in the sexual receptivity of non-protein fed females.     

Embryo transport through the mare's oviduct depends upon cleavage and is independent of the ipsilateral corpus luteum.

Two experiments were conducted using 14 mares. In Exp. 1, mares were inseminated with semen treated with TEPA, which, in other species, has been shown to lead to an arrest in ovum cleavage at 2--4 cells. The oviducts and/or uterus were then flushed 7--10 days after ovulation in 6 mares (Group A) or 2--6 days after ovulation in 5 mares (Group B). Fresh eggs were found in the oviduct flushes of 5 Group A and 5 Group B mares: 9 of the 10 eggs appeared to have cleaved, but none had developed beyond 16-cells. Seven eggs contained spermatozoa and 3 of 4 eggs from each group showed evidence of fertilization when examined ultrastructurally. Group A mares had thus retained fertilized eggs in the oviduct beyond the time at which they would normally have entered the uterus (6 days), indicating that development beyond at least the 2- to 4-cell stage is necessary for normal transport. In Exp. 2, 5 attempts were made to recover the embryo within 4 days of ovulation and transfer it to the contralateral oviduct. A single pregnancy resulted, indicating that a unilateral interaction with the corpus luteum was not necessary for the transport of the embryo to the uterus.     

Fertility of the post-partum bank vole (Clethrionomys glareolus)

Analysis of records of a bank vole breeding colony suggests that fertility is high immediately post partum, declines during established lactation and rises after weaning of young. Mating tests with lactating females and females whose young had been removed at birth showed that receptivity is reduced during lactation, although amongst the females which did mate there was no difference between lactating and non-lactating animals in the proportion which produced litters. However, average size of litters at birth was significantly larger for the lactating than for the non-lactating females. There is some evidence suggesting that this difference may arise after ovulation has occurred. Virgin females were no more receptive or fertile than lactating females.     

Material Benefits from Multiple Mating in Female Mealworm Beetles (Tenebrio molitor L.)

In yellow mealworm beetles (Tenebrio molitor), females are sexually receptive throughout their adult lives. We examined how access to mates affected female fecundity by varying the number of matings per female and quantifying cumulative egg production. Also, we dissected females at successive intervals after a single mating to assess the relationship among time since mating, sperm supplies, egg load, and oviposition rate. Females that mated at intervals greater than 2 days did not produce as many eggs as females that mated every 2 days or were allowed to mate ad libitum. Dissections showed that the amount of sperm remaining in a female spermatheca was correlated with the number of eggs she had laid recently, which suggests sperm replenishment as the material benefit gained through multiple mating. However, females mate more frequently than necessary for sperm replenishment, and therefore material benefits alone may not fully explain the continuous receptivity of T. molitor females.     

Capacitation status of hamster spermatozoa in the oviduct at various times after mating

Female hamsters were mated shortly after the onset of oestrus or immediately after ovulation. At various times after mating, spermatozoa were flushed from the isthmus of the oviduct using a modified Tyrode's medium supplemented with 20% hamster serum. Cumulus oophorus-free eggs were introduced into the suspensions of isthmic spermatozoa. Some eggs were removed every 30 min and examined for evidence of fertilization. For females mated shortly after the onset of oestrus, spermatozoa recovered from the oviducts 8 h after mating (about 1.5 h after ovulation) could penetrate eggs within 30 min and were considered fully capacitated. When spermatozoa were recovered at earlier times (1, 2, 4 and 6 h after mating) they required additional time (2, 1.5, 1 and 1 h respectively) in vitro before penetrating eggs. Therefore, when mating occurs shortly after the onset of oestrus, spermatozoa in the oviduct do not appear to become fully capacitated until about the time of ovulation. For females mated immediately after ovulation, spermatozoa recovered from the oviducts at 4 h after mating could penetrate eggs within 30 min. Spermatozoa recovered at 1 and 3 h after mating required 2 and 1 h respectively in vitro before penetrating eggs. These results suggest that sperm capacitation proceeds at a faster rate when mating occurs after ovulation.     

Spawning kinetics in the gilthead sea-bream, Sparus aurata L. after low doses of human chronic gonadotropin

This paper reports the temporal pattern of hormonally induced spawning in Sparus aurata females reared in captivity. A low dose of human chorionic gonadotropin (100–400 i.u. kg⁻¹ body weight), administered to females with oocytes in the last stages of vitello genesis, induced a daily cycle of maturation, ovulation and spawning of fertilizable eggs. This cyclic pattern lasted for a period of 4–100 days. In fish injected with saline and most of the untreated fish, all vitellogenetic eggs underwent rapid atresia.     

Environmental Cues, Endocrine Factors, and Reproductive Diapause in Male Insects

Environmental cues, mostly photoperiod and temperature, mediated by effects on the neuroendocrine system, control reproductive diapause in female insects. Arrest of oocyte development characterizes female reproductive diapause, which has two major adaptive functions: It improves chances of survival during unfavorable season(s), and/or it confines oviposition to that period of the year that is optimal for survival of the eggs and progeny. Although reproductive diapause is less well studied in male insects, there may be no sex-dependent differences in regard to the first of these functions. The second one, however, is not valid for the male; instead, selection pressure directs the male's reproductive strategy toward maximum chances of fertilization of the female's eggs with minimum waste of energy. Therefore, in species with female reproductive diapause, the males may or may not exhibit diapause, but if they do, their diapause must be adapted to that existing in conspecific females. Male reproductive diapause is defined as a reversible state of inability of the male to inseminate receptive females. In relation to reproductive diapause, there are several patterns of coadaptations between male reproductive strategy and timing of female receptivity, (a) In some insects, the females are receptive in the early part of their diapause; mating occurs during this period and there is no diapause in the male. The male dies shortly after copulation and the female stores the sperms to fertilize the eggs that develop after termination of the female's diapause, (b) In some species, as in the grasshopper Anacridium aegyptium, females are receptive during diapause; though oocyte development is arrested, copulation occurs and the stored sperms fertilize the eggs when the female's diapause ends. Males were claimed to have no diapause, but recent studies have revealed the presence of a reproductive diapause in a proportion of the males. This and other cases show that female receptivity during reproductive diapause may or may not be accompanied by male reproductive diapause. If there is a reproductive diapause in the male, it is controlled by the same endocrine mechanism, the corpora allata (CA), as in the females, (c) In many species females are refractory during their diapause. In these cases, males exhibit reproductive diapause, which may be light, as in the beetle Oulema melanopus, or well established, as in certain grasshoppers, butterflies, and beetles. In the latter cases, male diapause is controlled by similar environmental cues (photoperiod, temperature) and by the same intrinsic mechanism (neuroendocrine system, especially CA) as female diapause. Nevertheless, male diapause is less intense; the environmental cues leading to its termination are less complex and/or less extreme, so male diapause terminates before that of the females. Presumably, male diapause is under two antagonistic selection pressures: A male should not waste energy by courting dia-pausing refractory females, but he should be ready to copulate as soon as the females become receptive, otherwise he may lose in the competition between males for females. Some further strategies, which do not seem to fit the above patterns, are also outlined.     

Mate Guarding in the Androdioecious Clam Shrimp Eulimnadia texana: Male Assessment of Hermaphrodite Receptivity

Precopulatory mate guarding primarily occurs when males encounter receptive females at a low enough rate that such females become a valuable resource once encountered. Such circumstances are common in aquatic crustaceans wherein females are only receptive for a short period directly after molting. In these species, males commonly mate guard by physically attaching themselves to their prospective mates for hours to days at a time. To be effective in mate guarding, males must be able to assess the time to receptivity in their mates, which is commonly via chemical cues associated with molting. Clam shrimp in the genus Eulimnadia exhibit mate guarding, but with an important variation: these species are mixtures of males and hermaphrodites (androdioecy) rather than males and females. Nonetheless, the mate guarding behaviors of these shrimp are much the same as in other aquatic crustaceans. In this study, three projects were undertaken to determine the ability of Eulimnadia texana males to assess hermaphroditic receptivity. Males were found to be unable to assess receptivity without physically contacting hermaphrodites. However, after physical contact, males spent a significantly greater amount of time guarding receptive relative to non‐receptive hermaphrodites. Additionally, male interest in mate guarding was highest during the period between the dropping of one clutch of eggs and the extrusion of the following clutch. Because this period is also associated with hermaphroditic molting, it is consistent with the notion that males cue into chemicals associated with molting to determine hermaphroditic receptivity. These findings are consistent with previous studies of mating behavior in this species, and we discuss their importance to future tests of optimal mate guarding planned for these shrimp.     

Detrimental effect on availability of buserelin acetate administered in seminal doses in rabbits

The study evaluated a seminal effect on the ability to induce ovulation of a synthetic GnRH analogue, buserelin acetate, administered by vaginal mucosa in rabbit does. In a first experiment, 751 receptive nulliparous and multiparous non-lactating does were randomly assigned to groups of different seminal doses (6, 12, 24, 50, and 100 million total sperm in 0.5 mL). All seminal doses contained 5μg of buserelin acetate to induce ovulation by vaginal mucosa absorption. Two hundred and six does from 751 were laparoscopized at 12^th days of gestation to evaluate ovulation induction, ovulation rate and implanted embryos, while pregnancy rate and total and live born were noted in all females. Results showed that the pregnancy rate was significantly affected by the seminal dose used (0.82 vs 0.72, 0.50, and 0.45, for 6, 24, 50, and 100 million of spermatozoa, respectively). Data from laparoscopized does showed significant differences between the group of 6 and 50 million sperm dose in the ovulation induction and consequently in the pregnancy rate (0.79 vs 0.52, 0.79 vs 0.48, respectively). Does from all groups had similar implanted embryos and litter sizes irrespective of seminal dose used. In a second experiment, inseminations were done without spermatozoa, 0.5 mL of two dilutions of seminal plasma (1/4 and 1/20) with 5μg of buserelin acetate were introduced into vagina from 71 receptive females and its results were compared to a control group (35 does) induced to ovulate with 1μg of buserelin acetate administered intramuscularly. Only 40% of females from 1/4 plasma dilution group became to ovulate. Consequently, the dilution rate of seminal plasma may reduce the availability rate of the GnRH analogue and the concentration needed to provoke the ovulation induction.     

Influence of mating on sexual receptivity and oviposition in the mosquito, Culex tarsalis

ABSTRACT. Mated females of Culex tarsalis Coq. (Diptera, Culicidae) were receptive to additional insemination after gonotrophic activity. Sexual receptivity was not renewed in mated, aged nulliparous females. Insemination was determined by the detection in females of sperm labelled with adenine-³H(G) after matings with radio-labelled males. When females were deprived of males after their first insemination, the production of fertile eggs was reduced and oviposition activity was altered throughout successive gonotrophic cycles.     

Seasonal characteristics and tetrodotoxin-producing ability of bacteria in gastropod Oliva hirasei

Lycoteuthis lorigera is among the most abundant mesopelagic squid on the continental slope of the south Atlantic. It is the dominant prey in some commercially harvested groundfish. Despite its abundance and importance in the mesopelagic foodweb, its general biology is poorly known. In this study, the spawning pattern and fecundity of L. lorigera from southern African waters were investigated. Both histological examination and length frequency analysis of ovaries in various stages of development revealed that the ovulation pattern of L. lorigera is asynchronous, which indicates a spawning strategy where batches of eggs are spawned repeatedly over time. More specifically, ovulation is group synchronous, where distinct batches of 1,200–2,400 eggs ripen in the ovary and accumulate in the oviducts. The potential fecundity was estimated to be 20,000–50,000 in immature females (n = 6) and 8,000–25,000 in mature females (n = 21). The number of ripe eggs in the oviducts suggests that batches of spawned egg masses contain between 1,000 and 4,000 eggs. The reproductive strategy of L. lorigera is discussed and compared to the reproductive strategies of oceanic squid inhabiting the continental slope of southern Africa.     

Maturation-associated changes in the rat zona pellucida

Rat follicular oocytes, arrested at prophase I, cannot be fertilized in vitro. This capacity is acquired following resumption of meiosis and a series of changes involving both the oocyte and the cumulus cells surrounding it. Oocytes exposed to sperm at different hours before ovulation show a gradual increase in the permeability of their zona pellucida (ZP). Our study examined whether the ZP, in response to the physiological stimulus for maturation and concomitant with the other oocyte--cumulus components, undergoes maturational changes. Two ZP characteristics were assessed, sensitivity to proteolysis and sperm binding. ZP surrounding oocytes and eggs were collected from five sources: 1) germinal vesicle (GV)-intact oocytes, 2) preovulatory eggs, 3) ovulated eggs isolated from oviducts of immature females, 4) fertilized eggs, 5) ovulated eggs isolated from oviducts of mature females. All ZP surrounding oocytes/eggs from groups 1-5 were dissolved by trypsin. When solubility by pronase and alpha-chymotrypsin was examined, a large variation between groups was found. All ZP from group 2 were dissolved by 0.001% pronase, compared to 0% solubility in group 4. Only 10% of the ZP surrounding GV-intact oocytes (group 1) were dissolved by this enzyme, compared to 82% in group 3. Solubility in 0.01% alpha-chymotrypsin showed a similar pattern. Capacitated sperm were incubated with eggs from groups 1 and 3. The number of sperm binding to ZP in group 3 was repeatedly higher than that in group 1. In both tests it was found that the ZP surrounding the mature eggs differ in their characteristics from ZP of GV-intact oocytes.(ABSTRACT TRUNCATED AT 250 WORDS)     

Analyses of Oviductal Pars Recta-Induced Fertilizability of Coelomic Eggs in Xenopus laevis

The acquisition of fertilizability in coelomic eggs of Xenopus laevis has been shown to be correlated with the physical, biochemical, and ultrastructural alterations of the egg envelope [coelomic envelope (CE)] induced during the passage of eggs through the pars recta portion of the oviduct. However, no direct evidence that the pars recta renders eggs fertilizable has yet been presented. In this study, we show that coelomic eggs are highly fertilizable when they are incubated with continuous shaking for 4 h at 15 degrees C in pars recta extract (PRE) derived from females prestimulated by pregnant mare serum gonadotropin. The PRE from pituitary-stimulated Bufo japonicus was as potent as homologous PRE in rendering Xenopus eggs fertilizable. Incubation of coelomic eggs in PRE for 30 min induced a dramatic increase in the rates of sperm binding to the envelope to a level equivalent to that exhibited by the envelope from uterine eggs (VEs). The CE-to-VE ultrastructural conversion and a 43k-to-41k hydrolysis of the envelope glycoprotein component started 5 min after, and were completed by 15 min after, the start of incubation in PRE and were accompanied by an exposure of a new N-terminal sequence typical to gp41. Thus, the biochemical and ultrastructural conversions and the sperm-binding activity of the envelope induced by PREs, although being prerequisite, were not sufficient to render coelomic eggs fully accessible to fertilizing sperm.     

Chromosomal Polymorphism and Female Receptivity in a Natural Population of DROSOPHILA PERSIMILIS

In a natural population of Drosophila persimilis (McDonald Ranch, Napa Valley, California), KL and MD chromosomal arrangement frequencies undergo a seasonal cycle, with MD common in spring and KL common in summer. Samples collected from spring and summer provided isofemale strains established as homozygous KL and MD pairs (kinlines) with each pair derived from a single heterokaryotype wild progenitor. Haploid doses of chromosomes 2 and 4 were controlled by marker-cross derivations of kinlines. Percentage onset of female receptivity was measured from cultures at 25° and at 15°, using ten kinlines from spring and nine from summer collections, with fast-mating hybrid males as standard testers. Mating tests consisted of 20 tester males x 20 females of specific age, karyotype, and kinline observed for 30 min. At 25° females became receptive at 48 hr after eclosion: parental line (KL_i/KL_i and MD_i/MD_i) females were approximately equal at 55 to 60% receptive, while among hybrids, MD_i/MD_j homokaryotype females were significantly less receptive (68%) than all other outbred combinations (73 to 77%). At 15°, females became receptive at four days of age, with increases on the fifth and sixth days: both parental line and outbred MD/MD females were significantly more receptive (28% at four days and 62% at six days) than all heterokaryotype females (20 to 26% at four days and 55 to 59% at six days), which in turn were more receptive than KL/KL parental and outbred females (10% at four days and 40% at six days). Heterosis was expressed at 25°, but not at 15°. Thus, dominance for female receptivity was temperature dependent. Females polymorphic for these third chromosomal karyotypes possess differential temperature sensitivity for onset of receptivity and are likely to contribute in a significant way to the observed seasonal frequency cycle in the natural population from which they have been derived.