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Effect of cortisol on luteinizing hormone release during sexual behavior in the boar.

The effect of inhibiting the rise in cortisol concentrations that occurs at copulation upon luteinizing hormone release was studied in seven adult boars. Plasma samples were collected for assay of luteinizing hormone, testosterone and cortisol on a control day and before, during and after exposure to an estrous sow. The area under the curve was used to evaluate hormone production and treatment effects were evaluated by a paired Student's t-test. The 11 beta-hydroxylase inhibitor, metyrapone, was used to suppress glucocorticoid hormone production. Cortisol concentrations increased significantly (p less than 0.05) after breeding compared to values on the control day while treatment with metyrapone prior to breeding prevented the cortisol increase (p greater than 0.05). Although luteinizing hormone production increased significantly after copulation in both breeding experiments, metyrapone pretreatment resulted in a reduction of luteinizing hormone secretion (p less than 0.05). Testosterone production was also reduced in boars pretreated with metyrapone. The results suggest that the increased levels of cortisol occurring at copulation may enhance luteinizing hormone release in boars.

Animals↗

Social olfaction in male brown lemmings (Lemmus sibiricus = trimucronatus) and collared lemmings (Dicrostonyx groenlandicus): II. Discrimination of mated and unmated females.

When tested in a Y-maze olfactometer, male brown and collared lemmings (Lemmus sibiricus = trimucronatus and Dicrostonyx groenlandicus) preferred the odor of unmated receptive females to the odor of females with which they had just copulated. Similarly, sexually satiated males preferred the odor of an unmated receptive female to that of a strange female that had recently copulated with another male. Sexually experienced males without recent copulatory experience also demonstrated this preference, but sexually naive males did not. Sexually satiated collared lemmings preferred the odor of bedding from a novel estrous female to bedding from the female with which they had just copulated even when the bedding was collected before mating occurred. These results suggest that discrimination between prior mates and unmated females may be based on individual recognition as well as recognition of subclasses of females (i.e., mated vs. unmated, familiar vs. unfamiliar).

Animals↗

Reproduction in wild gorillas and some comparisons with chimpanzees.

Information was collected over a period of almost 12 years on the gorillas of the Virunga Volcanoes region of Rwanda and Zaire, most of it collected since September 1972. Comparisons were made with the Gombe Stream chimpanzee population (values in parentheses). Gorilla females matured at about 8 years (compared with 9-10 years for the chimpanzee) and first bred at 10-11 years (11-12 years). Males started to breed later, possibly at 15 years (about 13 years). Oestrous periods of female gorillas lasted for about 2 days (10 days) and oestrous cycles for about 30 days (36 days). Gestation in the gorilla lasts 255 days (228 days). Intervals between surviving offspring in the Virunga study groups was about 4 1/2 (5 1/2 years) but in the whole Virunga population was nearer 8 years. Lactational amenorrhoea lasted about 2 1/2 years (3 1/2 years). In gorillas and chimpanzees there were about 3 cycles to conception after parturition and females probably produce in their lifetime about 3 offspring that survive to adulthood. A successful male's productivity is greater. Mortality of immature gorillas was about 40% (50%). Initiation of courtship is generally by the female in gorillas but the male in chimpanzees. Copulation in gorillas lasts for about 1 1/2 min (7 sec) and occurs at a rate of once every 3 h when a female is in oestrus (once in 2 h). Interference in copulation is more common in the chimpanzee than the gorilla, but competition between individual males is more intense in gorilla populations. Females of both species can clearly exercise their preferences for particular males. The observed differences between the species in courtship and mating behaviour can be related to differences in the number of males available to and competing for oestrous females: in the loose multi-male chimpanzee community there is more advantage to males in initiating copulation and mating frequently and efficiently, and to females in advertising oestrus, than in the relatively stable one-male mating system of the gorilla.

Animals↗

[The sexual behavior of the African ostrich (Struthio camelus)].

The African ostrich (Struthio camelus) is different from other birds not only because it is a running bird (Ratitae), but also because it urinates and the male bird has a penis. On a specialized farm in Israel the sexual behaviour of ostriches was studied. Courtship behaviour and mating have a very refined pattern. The copulation, which starts with an immission of the penis and ends with an orgasm, lasts about one minute. Most of the copulations take place during the morning morning hours. Occasionally, female ostriches demonstrate courtship behaviour to other females. Sometimes female ostriches mount other females as in the case of copulation.

Animals↗

Dynamic changes in nucleus accumbens dopamine efflux during the Coolidge effect in male rats.

The Coolidge effect describes the reinitiation of sexual behavior in a "sexually satiated" animal in response to a novel receptive mate. Given the role of the mesolimbic dopamine (DA) system in the initiation and maintenance of motivated behavior, microdialysis was used to monitor nucleus accumbens (NAC) DA transmission during copulation, sexual satiety, and the reinitiation of sexual behavior. In agreement with earlier reports, the presentation of an estrous female behind a screen and copulation were associated with significant increases in NAC DA efflux. Return of NAC DA concentrations to baseline values coincided with a period of sexual satiety, although concentrations of the DA metabolites, dihydroxyphenylacetic acid and homovanillic acid, remained elevated. The presentation of a novel receptive female behind a screen resulted in a slight increase in NAC DA, which was augmented significantly during renewed copulation with the novel female. The present data suggest that the stimulus properties of a novel receptive female may serve to increase NAC DA transmission in a sexually satiated male rat, and this, in turn, may be related to the reinitiation of sexual behavior.

3,4-Dihydroxyphenylacetic Acid↗

Lesions in medial preoptic area and bed nucleus of stria terminalis: differential effects on copulatory behavior and noncontact erection in male rats.

The goal of these studies was to assess the regulatory roles of the medial preoptic area (MPOA) and the bed nucleus of the stria terminalis (BST) in sexual arousal, inferred from noncontact erection (NCE) evoked in male rats by remote cues from estrous females. NCE and copulatory behavior were recorded before and after quinolinic acid or radiofrequency (RF) lesions were made in the MPOA (Experiments 1-3) or RF lesions were made in the BST (Experiment 4). All males with MPOA lesions, particularly in the rostral region, displayed severe deficits in copulation but little or no decrement in NCE. In contrast, BST lesions caused relatively moderate deficits in copulation, but they severely impaired NCE. Animals with larger BST lesions, including rostral and caudal medial regions, had more deficits in both copulatory behavior and NCE than did males with smaller lesions confined to the rostral medial BST. These results suggest that (1) the MPOA is critical for copulatory behavior but not for NCE, (2) males that stop copulating after MPOA lesions are still sexually aroused by estrous females, and (3) the BST plays an important role in mediating NCE.

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Multi-messenger innervation of the male sexual system of Lymnaea stagnalis.

The male copulation behaviour of the hermaphrodite pond snail Lymnaea stagnalis is under the control of five groups of central neurons that produce a variety of neuropeptides and a classical transmitter, 5-hydroxy tryptamine (5HT). In this article, we describe how the male sexual organs of this snail are innervated by axons from these central neurons. We carried out immunocytochemistry with antisera against the tetra peptide Ala-Pro-Gly-TRP-NH2 (APGWamide), the Lymnaea form of neuropeptide tyrosine (LNPY), conopressin, pedal peptide, the FRMFamide copeptide SEEPLY, the GDPFLRFamide co-peptide DEILSR, myomodulin, Lymnaea inhibitory peptide, and 5HT on tissue sections of the following male sexual organs that receive input from the penis nerve: the prostate gland, vas deferens, preputium, and penis. The results demonstrate that the axons of the separate muscle systems contain particular combinations of transmitters. In addition, two networks of peripheral neurons were revealed. In the tip of the everted preputium lies what appears to be a network of conopressin-containing sensory neurons, which is possibly involved in probing; probing is the part of copulation behaviour in which the male searches for the female genital pore. The other network of peripheral neurons surrounds the most proximal part of the vas deferens and is most likely involved in the pacemaker control of vas deferens motility. On the basis of the data obtained, we hypothesize how the preputium and penis are everted during copulation and which transmitters and central neurons might be involved.

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Reproductive inefficiency in male black-footed ferrets (Mustela nigripes).

The black-footed ferret (Mustela nigripes), once considered extinct, has benefited from captive breeding and subsequent reintroduction into native habitat. A high proportion of females (>90%) exhibit estrus in captivity during the spring breeding season. However, many males considered to be prime-breeding age (1-3 years old) fail to sire offspring. Breeding records in 1995 revealed that 40 of 73 males (55%) managed under the Black-Footed Ferret Species Survival Plan did not reproduce, despite being provided opportunity. The present study was conducted to determine the incidence and etiology of male reproductive failure in 1996 and 1997. In 1996, 38 of 69 (55%) 1- to 3-year-old males failed to sire offspring. Likewise, 35 of 60 (58%) males did not reproduce in 1997. Overall, 21% of adult males failed to sire young in three consecutive breeding seasons (1995-1997). Electroejaculate traits (ejaculate volume, sperm concentration, motility, morphology, and acrosomal integrity) from 29 proven breeder males were not different (P > 0.05) from those of 23 males that did not sire young (nonproven breeders). However, six categories of reproductive failure were identified for the 73 prime-breeding age, nonproven males: 1) underdeveloped testes (22%); 2) improper breeding position with the female (25%); 3) excessive aggression toward estrous females (9%); 4) copulation with no sperm detected at postcoital lavage (19%); 5) copulation with sperm in the vaginal lavage but no resulting pregnancy (18%); and 6) copulation with no vaginal lavage performed and no resulting pregnancy (7%). These data indicate that combined behavioral and physiologic factors, but not overall sperm quality, influence reproductive performance in male black-footed ferrets managed in captivity. Zoo Biol 19:517-528, 2000. Copyright 2000 Wiley-Liss, Inc.

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Patterns of behavior across reproductive states of free-ranging female black-handed spider monkeys (Ateles geoffroyi).

Reports on the behavior of spider monkeys (genus Ateles) describe a suite of 5-6 behaviors that are indicative of an estrous female. This study presents hormonal data as an independent measure of reproductive state to determine if these behaviors are associated with any particular reproductive state or stage of the ovarian cycle. Fecal samples collected from 6 free-ranging female black-handed spider monkeys (Ateles geoffroyi) over the course of 11 months were assayed for estrogen (E1C) and progesterone (PdG) metabolites, using an enzyme immunoassay. Behavioral data collected from the same females were analyzed for patterns associated with different reproductive states. A more detailed analysis of behavioral data associated with reproductively cycling periods determined whether each behavior clustered with copulations or a particular phase in the ovarian cycle. Observations of place-sniffing were more frequent for 4 of 5 females when reproductively cycling. Of all copulations observed, 80% occurred when the female participant was reproductively cycling. In addition, the rates of self-clitoral hold and self-clitoral rub behaviors were significantly higher for 2 of 4 and 4 of 5 of females, respectively, when reproductively cycling. No behavior consistently occurred in association with copulations or with any particular phase in the ovarian cycle. Results from this study suggest that the behaviors tested in this paper can provide some information regarding female spider monkey reproductive state (whether or not she is reproductively cycling), but that they are not indicators of estrus in female spider monkeys.

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Mating biology of the leaf-cutting ants Atta colombica and A. cephalotes.

Copulation behavior has often been shaped by sexually selected sperm competition or cryptic female choice. However, manipulation of previously deposited ejaculates is unknown in the social Hymenoptera and the degree to which sperm competes after insemination or is actively selected by females has remained ambiguous. We studied the mating process in the leaf-cutting ants Atta colombica and A. cephalotes, which belong to one of the few derived social insect lineages where obligate multiple mating has evolved. As copulations often occur at night and in remote places, direct observations were impossible, so we had to reconstruct the sequential copulation events by morphological analysis of the male and female genitalia and by tracking the process of sperm transfer and sperm storage. We show that Atta male genitalia have two external rows of spiny teeth, which fit into a specialized pouch organ in the female sexual tract. Reconstruction of the sperm storage process indicated that sperm is transferred to the spermatheca during or immediately after ejaculation and without being mixed with sperm and seminal fluids from other males. A convergent mechanism of direct sperm transfer to the spermatheca of queens is known from two species of dwarf honeybees. Direct sperm transfer may restrict female control over the sperm storage process and the number of males that contribute to the stored sperm.

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Extra-pair mating effort of male hooded warblers, Wilsonia citrina

Few researchers have been able to quantify the time or effort expended by male birds in seeking extra-pair copulations with neighbouring females. In this study, we radio-tracked male hooded warblers, to determine the frequency and duration of intrusions onto neighbouring territories. Extra-pair fertilizations are common in hooded warblers, with 35% of females producing extra-pair young. Males intruded onto territories where females were nest building more often than expected by chance and approached females during most intrusions (78%), suggesting that these intrusions were extra-pair copulation attempts. Almost all intrusions (96%) were to adjacent territories, and males made an average+/-se of 0.4+/-0.2 forays/h and spent 4.7+/-2.6% of their time off-territory. Males were involved in fights or chases with the resident male in 21% of intrusions, but were apparently undetected during most intrusions. Males with high intrusion effort onto neighbouring territories had the lowest rate of intrusions onto their own territory by other radio-tracked males. Males varied extensively in their intrusion effort, but this was not significantly correlated with male body mass, age or song rate. The percentage of time spent intruding onto other territories was small (0-8%), so males probably do not face strong trade-offs between making extra-pair copulation forays and other activities like mate guarding and feeding offspring. Copyright 1998 The Association for the Study of Animal Behaviour Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Certainty of paternity and paternal investment in eastern bluebirds and tree swallows.

Extra-pair paternity is common in many socially monogamous passerine birds with biparental care. Thus, males often invest in offspring to which they are not related. Models of optimal parental investment predict that, under certain assumptions, males should lower their investment in response to reduced certainty of paternity. We attempted to reduce certainty of paternity experimentally in two species, the eastern bluebird, Sialia sialis, and the tree swallow, Tachycineta bicolor, by temporarily removing fertile females on two mornings during egg laying. In both species, experimental males usually attempted to copulate with the female immediately after her reappearance, suggesting that they experienced the absence of their mate as a threat to their paternity. Experimental males copulated at a significantly higher rate than control males. However, contrary to the prediction of the model, experimental males did not invest less than control males in their offspring. There was no difference between experimental and control nests in the proportion of male feeds, male and female feeding rates, nestling growth and nestling condition and size at age 14 days. We argue that females might have restored the males' confidence in paternity after the experiment by soliciting or accepting copulations. Alternatively, males may not reduce their effort, because the fitness costs to their own offspring may outweigh the benefits for the males, at least in populations where females cannot fully compensate for reduced male investment. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

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Sex-role reversal and the absence of extra-pair fertilization in Wilson's phalaropes.

Extra-pair fertilizations are common in birds, which has led to great interest in how this phenomenon is regulated at a proximate level and how extra-pair fertilizations, and extra-pair fertilization avoidance, shape avian social systems. In Wilson's phalaropes, Phalaropus tricolor, incubation and brood-rearing is performed exclusively by males. Males are able to rear only a single brood of four during a breeding season. This suggests that males have a high level of paternity in clutches and broods under their care and, thus, that extra-pair fertilizations are infrequent. In contrast, female social dominance, lack of territoriality and frequent interactions among breeding adults suggest that both males and females have the opportunity to engage in extra-pair copulations. Using DNA fingerprint band-sharing between putative parents and offspring, we found no evidence of extra-pair fertilizations among 51 offspring from 17 families of phalaropes. Copulation disruption by non-copulatory adults, ability of females to reject copulation attempts and potential fitness benefits to females by avoiding extra-pair fertilizations were sufficient to explain the absence of extra-pair fertilizations in Wilson's phalaropes. We propose that sex-role reversal affects the relative costs and benefits to females of seeking extra-pair fertilizations. At the time of clutch completion, females have invested particularly heavily in their clutches due to intense competition among females to gain and keep a mate during the pre-laying and laying periods. After clutch completion, nest success requires significant male parental care. Benefits to females in gaining extra-pair fertilizations may be offset by the risk of losing male parental care. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

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Female sexual preferences differ in Mus spicilegus and Mus musculus domesticus: the role of familiarization and sexual experience.

Mating systems correspond to particular ecological conditions and result from proximate interactions between individuals. We compared the mating preferences of female mice of two species: the house mouse, Mus musculus domesticus, and the mound-builder mouse, Mus spicilegus. Because of differences in their habitat, we expected to observe differences in their sexual preferences. We studied female preferences for a familiar or an unfamiliar male and the occurrence of copulation with the unfamiliar male, during two states of female sexual activity: (1) the postpartum oestrus of paired females, to evaluate the stability of their sexual partnership; and (2) the oestrus of females familiarized with a male, to study the mechanisms underlying their sexual preferences. In the polygamous house mouse, postpartum oestrous females did not show a clear preference between their familiar male and the unfamiliar one. Moreover, oestrous females, familiarized with a male (without sexual interactions), preferred an unfamiliar male and copulated with him. In contrast, postpartum oestrous females and oestrous females of M. spicilegus preferred their familiar male and rarely copulated with the unfamiliar male. This study indicates a strong pair bond in established breeding pairs in M. spicilegus and shows that this bond can be established by familiarization, which is not the case in M. m. domesticus. Our study suggests the existence of monogamous traits in M. spicilegus in contrast to the polygamous M. m. domesticus. (c) 1998 The Association for the Study of Animal Behaviour.

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Postcopulatory sexual selection in Mediterranean fruit flies: advantages for large and protein-fed males.

Previous laboratory studies of Mediterranean fruit flies, Ceratitis capitata (medflies), have identified large size and protein feeding as positive influences on the ability of males to secure copulations. In this study, we investigated whether large and protein-fed males experience additional advantages in terms of amount and distribution of sperm stored by mates. We also examined relationships between copula duration and sperm storage. Mates of large and protein-fed males were more likely to store sperm and to store more sperm than mates of small and protein-deprived males. Probability of sperm storage was associated with copula duration; all copulations lasting less than 100 min failed whereas 98% lasting longer than 100 min succeeded. Copulations involving sperm storage were longer if males were small or protein deprived or if the female was large, although there was no evidence of a relationship between copula duration and total sperm storage. Evidence from related studies suggests that variation in latency until sperm transfer, caused by size and diet, is a likely explanation for varying copula duration. Sperm tended to be stored asymmetrically between the female's two spermathecae, consistent with a mating system in which females maintain isolated populations of sperm from different males and later select between them. Storage was less asymmetric when large numbers of sperm were stored but there was little evidence that male size or diet affected this asymmetry. It is uncertain whether postcopulatory advantages of large and protein-fed male medflies arise from female preferences or male dominance through coercion or force. Copyright 1999 The Association for the Study of Animal Behaviour.

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Male mating behaviour and sperm production characteristics under varying sperm competition risk in guppies.

Since natural populations of guppies, Poecilia reticulata, often differ from one another in social structure, the intensity of sperm competition is likely to vary between localities. Guppies are promiscuous, with female choice for colourful males playing a central role in the mating system. In addition, male guppies use forced copulations to circumvent female choice. Both methods of copulation are used interchangeably by individual males, but the degree to which either is used may depend on the social environment into which males are born. Here we show that male mating behaviour varies according to the rearing sex ratio: when reared in male-biased groups, males performed more forced copulations and fewer courtship displays but showed the opposite pattern of behaviour when reared in female-biased groups. Our prediction, based on sperm competition theory, that stripped sperm number would reflect social structure was not supported by our results. Instead, the overall level of sexual activity (gonopodial thrusts+sigmoid displays) was a better predictor of sperm number in the different groups of males. Rearing density, where sex ratio was controlled, did not significantly affect male mating behaviour or sperm traits. Males reared under the different sex ratios continued to show their characteristic behaviour patterns when placed in equal sex ratio tanks. We conclude, therefore, that males adopt mating strategies to suit their social environment, and that these strategies remain fixed, for short periods at least, if population structure changes. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗

Population density influences male-male competition in guppies.

This study tested the general prediction that population density affects male-male competition, female mate choice and the opportunity for sexual selection. By manipulating the density of guppies, Poecilia reticulata, while keeping the sex ratio constant, I found that male mating tactics were phenotypically plastic with respect to density. As density increased, males decreased their courtship displays. Male-male competition and mate searching were highest at intermediate densities. Population density had no significant effect on the total number of copulations, copulatory tactics or the percentage of postcopulatory guarding. Female preference for males with a higher percentage of orange coloration was similar at all density levels. The 'opportunity for sexual selection', which estimates the upper limit to which a selected trait can shift if directional selection is operating and was calculated as the variance in number of copulations per male divided by the square of the mean number of copulations, was negatively associated with population density. This may be due to the decrease in male-male competition at high density rather than female preference which was similar across density treatments. Copyright 1999 The Association for the Study of Animal Behaviour.

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Size and stability of vertebrate leks.

Lek size varies greatly among lekking species. At present there is no explicit theoretical explanation for this diversity. We extend an existing model of optimal lek size that incorporates female mating preferences and male-male contest competition. The model shows that variation in lek size is predicted by the interaction between lek size, overall copulation rate and the proportion of copulations accruing to males of different rank. In species where females prefer to mate on the largest leks and high-ranking males are able to monopolize females irrespective of the size of the lek, the maximum lek size will be large. Conversely, in species where females show weak preference for mating on large leks or increasing lek size quickly results in scramble competition, the maximum lek size will be smaller. Thus, differences between species in lek size may be due largely to differences in the extent to which high-ranking males can monopolize mating opportunities. Leks become unstable and break down when high-ranking males can no longer get their 'expected' copulation success. Therefore, the mechanism that generates male clustering, that is, sexual parasitism of high-ranking males by subordinates, also sets a limit to the largest stable lek size. Copyright 1999 The Association for the Study of Animal Behaviour.

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