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Reproductive toxicity assessment by continuous breeding in Sprague-Dawley rats: a comparison of two study designs.

The protocol for Reproductive Assessment by Continuous Breeding (RACB) studies was originally designed to use mice as the test species. However, rats are commonly used for reproductive toxicity research and could be used in the basic RACB design. One of the studies reported below evaluated a standard murine RACB design using rats, which rears the fifth litter to test second generation fertility. The second design tested the logistics and feasibility of rearing the second litter for second generation fertility testing. The standard fifth litter design (L5) was modified slightly for rats by increasing the time allowed for gestation and delivery. Compared to rats rearing their second litter (L2), rats in this L5 design had more litters per pair during continuous breeding and maintained this fertility better over time, as evidenced by producing more pups per litter during the crossover mating segment. Both L2 and L5 rats gave sufficient pups to conduct the second generation fertility evaluation. In addition, the L5 design was easier to conduct and produced pups from gametes exposed to chemical throughout spermatogenesis, making it the preferred design for using rats in Continuous Breeding studies.

Animals↗

Analyzing correlations of three types in selected lines of Drosophila melanogaster that have evolved stable extreme geotactic performance.

The behavior-genetic analysis of Drosophila melanogaster with geotactic performance as the phenotype is an ideal model system with which to investigate the complex relations between heredity and behavior. As part of a long-term, 38-year study, we report 4 experiments that identify and analyze trait correlations in the selected high- and low-geotaxis lines. We performed F2 correlational analyses and backcrosses to examine 3 types of correlations: (a) genotype-genotype (alcohol dehydrogenase [Adh]-amylase [Amy]), (b) genotype-phenotype (Adh and Amy-geotaxis), and (c) phenotype-phenotype (mate preference-geotaxis). Only the Adh-geotaxis correlation survived meiosis and reappeared in the F2 generation, which indicates a genotype-phenotype correlation, whereas the others did not. The importance of hybrid correlational analysis to the behavior-genetic analysis of a species is discussed.

Alcohol Dehydrogenase↗

Male and female mate choice affects offspring quality in a sex-role-reversed pipefish.

Where both sexes invest substantially in offspring, both females and males should discriminate between potential partners when choosing mates. The degree of choosiness should relate to the costs of choice and to the potential benefits to be gained. We measured offspring quality from experimentally staged matings with preferred and non-preferred partners in a sex-role-reversed pipefish, Syngnathus typhle L. Here, a substantial male investment in offspring results in a lower potential reproductive rate in males than in females, and access to males limits female reproductive success rather than vice versa. Thus, males are choosier than females and females compete more intensely over mates than do males. Broods from preferred matings were superior at escaping predation, when either males or females were allowed to choose a partner. However, only 'choosing' females benefited in terms of faster-growing offspring. Our results have important implications for mate-choice research: here we show that even the more competitive and less choosy sex may contribute significantly to sexual selection through mate choice.

Animals↗

Olfactory receptor-like genes are located in the human major histocompatibility complex.

The murine major histocompatibility complex (MHC) includes sequences that are responsible for haplotype-specific odor types that, in turn, influence mating preference. We report that there are several olfactory receptor genes or pseudogenes in the Class I region of the human MHC. At least one of these genes is intact, appears to encode an mRNA, and is quite homologous to a previously reported murine olfactory receptor.

Amino Acid Sequence↗

Removal of the preputial glands alters the individual odors of male MHC-congenic mice and the preferences of females for these odors.

Inbred mice of MHC-congenic strains prefer to mate with mice of different MHC types. Mate selection based on MHC-type occurs through the urinary odors specific to each MHC-congenic mouse strain. Both urine and preputial gland odors of male mice have been postulated to act as female attractants and the present study examined the attraction of MHC-congenic female mice (C57BL/6J and B6-AKR.H-2k) to the urine and preputial gland odors of males. Females of both strains showed a greater attraction to the urine odors of intact males of the other MHC type, but did not show a preference for the urine odors of these males after their preputial glands were removed. There was, however, a preference for the preputial fluid of males of the other MHC type. These results indicate that the MHC-specific urinary odors of mice may arise from the preputial glands, and that it is the MHC-specific preputial gland odors of MHC-conspecific males that are used for mate selection by females.

Animals↗

Attractivity and social preferences in mice (Mus musculus domesticus): the role of prepubertal sexual segregation and of precocious weaning.

Mice (Mus musculus domesticus) were raised (Postnatal Day 15 to 25) in single- or mixed-sex litters and precociously (Day 15) or regularly (Day 25) weaned. When they were faced as adults with a basic social choice--between two stimulus mice raised in litters of different sex composition but both of the same sex as the chooser--mice raised in mixed-sex litters were preferred. In the sociosexual choice-between a male and a female, both from the single- or the mixed-sex group--the opposite-sex preference was expressed. Both these preferences were abolished by the sexual segregation of the choosers. This variable hardly affected potential mate choice--between two stimulus mice both of the opposite sex of the chooser but raised in litters of different sex composition. Data indicate that socially mediated behavioral plasticity has a major role in the early shaping of adult individual differences both in attractive stimulus properties and in sociosexual preferences.

Animals↗

Beyond reinforcement: the evolution of premating isolation by direct selection on preferences and postmating, prezygotic incompatibilities.

The evolution of premating isolation after secondary contact is primarily considered in the guise of reinforcement, which relies on low hybrid fitness as the driving force for mating preference divergence. Here I consider two additional forces that may play a substantial role in the adaptive evolution of premating isolation, direct selection on preferences and indirect selection against postmating, prezygotic incompatibilities. First, I argue that a combination of ecological character displacement and sensory bias can cause direct selection on preferences that results in the pattern of reproductive character displacement. Both analytical and numerical methods are then used to demonstrate that, as expected from work in single populations, such direct selection will easily overwhelm indirect selection due to low hybrid fitness as the primary determinant of preference evolution. Second, postmating, prezygotic incompatibilities are presented as a driving force in the evolution of premating isolation. Two classes of these mechanisms, those increasing female mortality after mating but before producing offspring and those reducing female fertility, are shown to be identical in their effects on preference divergence. Analytical and numerical techniques are then used to demonstrate that postmating, prezygotic factors may place strong selection on preference divergence. These selective forces are shown to be comparable if not greater than those produced by the low fitness of hybrids.

Animals↗

Antennal asymmetry and sexual selection in a cerambycid beetle.

Cerambycid beetles have exaggerated antennae that are usually sexually size-dimorphic. We investigated the relationship between antenna morphology and sexual selection in the species Stenurella melanura (L.) in which males on average have antennae that are 13% longer than those of females. Males and females aggregate at flowers near oviposition sites for feeding during June-August. We sampled both copulating and single individuals at these sites. Fluctuating asymmetry (a measure of developmental instability) in antennae was considerably larger than in tibia and elytra and males had larger degrees of asymmetry in their antennae than females. Mated individuals did not differ from unmated individuals with respect to any of three size variables, but antennal asymmetry was smaller in mated individuals of both sexes. When two males were released with a female on a flower, males with symmetric antennae more often won the fight over the female than expected by chance. When two females and a male were released on a flower, the male more often preferred the female with more symmetric antennae than expected by chance. These results suggest that antennal symmetry, but not length, is currently under sexual selection.Copyright 1997 The Association for the Study of Animal Behaviour

Journal Article↗

Olfactory discrimination of estrous condition by the male golden hamster (Mesocricetus auratus).

After repeated exposure to receptive and nonreceptive females, male golden hamsters were tested for olfactory preferences in a four-choice olfactometer. Males discriminated between the odor of anesthetized females in different stages of their estrous cycle when the airstreams carrying the stimulus odors were diluted. Previous failures to demonstrate such an ability were probably due to ceiling effects resulting from laboratory testing. Males preferred the odor of females on the day before receptivity (a day coincident with maximal scent marking by the female) and least preferred the odor of females on diestrus-1 (the day on which females attacked and chased males during pretesting encounters). Detection and quick response to an impending estrus would be especially important for males of a solitary and promiscuous species in which there is a first male mating advantage.

Animals↗

Chemoreception, symmetry and mate choice in lizards.

Research on fluctuating asymmetry (FA)-mediated sexual selection has focused almost exclusively on visual signals and ignored chemical communication despite the fact that many species rely on chemical signals for attracting mates. Female mate choice based on visual traits appears to be rare in lizards. However, the femoral glands of male lizards produce pheromones which might transmit chemical information about an individual's developmental stability. Therefore, we hypothesized that mate choice may be based on chemical cues. We analysed the effect of the developmental stability levels of males on the attractiveness of males' scents to females in a laboratory experiment with the lizard Lacerta monticola. When we offered two males of similar body size, females preferentially associated with the scents of males with low FA in their femoral pores and also with the scents of males with a higher number of femoral pores. This suggested that the females were able to discriminate the FA of the males by chemical signals alone and that the females preferred to be in areas marked by males of high quality, thus increasing their opportunities of mating with males of high quality. We suggest that the quality and/or amount of male pheromones could communicate the heritable genetic quality of a male to the female and thereby serve as the basis for adaptive female choice in lizards.

Animals↗

Thalamocortical mechanisms in odor-guided behavior. II. Effects of lesions of the mediodorsal thalamic nucleus and frontal cortex on odor preferences and sexual behavior in the hamster.

The role of the thalamocortical "olfactory" pathway in odor preferences and sexual behavior was examined in the hamster, a species which depends crucially upon olfaction for mating. Before and after ablation of primary or secondary olfactory structures, male hamsters were tested for threshold, interest, and preference for the odor of an estrus female hamster and other odors. Sexual competency was also observed in daily mating sessions. Disruption of the primary olfactory pathway resulted in an absence of interest in odors and mating. Lesions of the mediodorsal thalamic nucleus or frontal neocortex of the rhinal sulcus did not result in anosmia, but did eliminate or alter odor preferences and resulted in inappropriate, inefficient, precopulatory and copulatory behavior. It appears that substructures in the thalamofrontal pathway playa role in discriminative or cognitive aspects of processing adaptively significant stimuli.

Animals↗

Digit ratio varies with sex, egg order and strength of mate preference in zebra finches.

The steroid environment encountered by developing vertebrates has important organizational effects on physiology and behaviour that persist throughout an organism's lifetime. Optimal allocation of maternal steroids to zygotes may be difficult to achieve because of the sexually antagonistic effects of steroids; thus, for example, a hormone environment beneficial to a developing male may be much less beneficial to a developing female. Research into the important topic of how mothers might adaptively adjust steroid titres experienced by particular young has been constrained by the difficulty of measuring the steroid environment experienced by the embryo at critical times in development. A potential approach to this problem has been suggested by research on variation in digit ratios in humans, where the ratio of the length of the second and fourth digits reflects the steroid environment experienced by the foetus; notably, digit 4 lengthens in response to androgens. In light of the conservative nature of homeobox genes regulating early development in tetrapods, we questioned whether a sex difference in digit ratio exists in a passerine bird, the zebra finch, Taeniopygia guttata castanotis, and whether observed variation in the ratio is consistent with the previously reported pattern that androgen allocation to zebra finch egg yolk declines across laying order. We established an aviary population of outbred, wild-type zebra finches, and allowed them to breed freely. Hatchlings were marked to correspond to their egg order, and their digit ratios were measured after birds reached adulthood. We found that digit ratio increased across egg order, which is consistent with a pattern of decreasing androgen allocation. Moreover, digit ratios differed between the sexes. We also investigated whether variation in digit ratio among adult females predicted variation in their performance in mate-choice tests. Digit ratio accounted for almost 50% of the variance in strength of female preference for an attractive male trait: specifically, females with higher (presumably less 'androgenized') ratios had stronger preferences for attractive males. Digit ratio may prove to be an extremely useful tool for addressing a wide range of questions about vertebrate differentiation and behaviour.

Androgens↗

Ultraviolet vision and band-colour preferences in female zebra finches, Taeniopygia guttata

Zebra finches have previously been found to have preferences for particular colours of both natural and artificial traits among opposite sex conspecifics. For example, in some studies female zebra finches preferred males wearing red leg bands to orange-banded and unbanded birds and rejected light green-banded males. In other studies, females also preferred males with red beaks to orange-beaked males. However, several authors have failed to replicate these results. We show that females may fail to show a colour preference because of the absence or removal of ultraviolet light under experimental conditions. In mate-choice trials, females observing males through filters that transmitted ultraviolet preferred red-banded males but where females viewed males through ultraviolet-blocking filters, no such preference was observed. Further investigation revealed that the lack of a colour preference when ultraviolet was absent was probably due to the change in overall appearance of the bird, rather than the change in appearance of the rings themselves. This work highlights the importance of proper consider-ation of the sensory capabilities of animals in experimental design, particularly with regard to the role of ultraviolet light in avian colour perception.Copyright 1997 The Association for the Study of Animal BehaviourCopyright 1997The Association for the Study of Animal Behaviour.

Journal Article↗

Consistency of female choice in the túngara frog: a permissive preference for complex characters

Previous phonotaxis experiments in the túngara frog, Physalaemus pustulosus, indicated a permissiveness in female preference that allows sexually selected complex call characters to be replaced with various alternative characters. Although they prefer complex to simple calls, females as a group did not discriminate between several alternative complex characters appended to the simple conspecific call. However, these studies did not address the possibility that the apparent permissiveness in female preference occurred because of an averaging of population data. The observed patterns in female preference could result from all females finding a certain set of call variants equally attractive, or from a polymorphism in female call preference. To discriminate between these two alternatives, consistency of mate choice was determined for three pairs of calls that elicited no phonotactic bias in population studies. Individual females did not repeatedly choose one stimulus of a pair over the other, demonstrating that the patterns of permissiveness observed in the population are paralleled by similar patterns within females. A broad preference for complex calls in the P. pustulosus species group would permit the evolution of sexually selected call variation through sensory exploitation. Copyright 1998 The Association for the Study of Animal Behaviour Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Deleterious mutations as an evolutionary factor. III. Mating preference and some general remarks.

This paper is concerned with the evolution of female mating choice under truncation selection against deleterious mutations. It is shown that with genome degradation rates necessary to maintain crossing-over, the advantage of choice is of the order of several per cent, which is in good accord with some experimental data. Some other consequences of intensive selection against mutations are also discussed.

Animals↗

Special relationships instead of female dominance for redfronted lemurs, Eulemur fulvus rufus.

Most lemurs yet studied in detail exhibit some mode of adult female social dominance over males. The known exception, a brown lemur subspecies known as rufous or redfronted lemurs (Eulemur fulvus rufus), forms multimale-multifemale social groups within which unambiguous dominance relations are not observed among adults. Resting groups of redfronted lemurs consistently include huddling adult male-female pairs whose males selectively scentmark and rub their heads in the scentmarks of their female huddling partners. Quantitative observations confirmed that some of these male-female pairs maintain special relationships satisfying all criteria originally developed in research on cercopithecine monkeys. Observations before, during, and after mating season, intergroup encounters, male transfers, and changes in male-female affiliations illuminated developmental and functional aspects of male-female partnerships. Each adult female in two semi-free-ranging study groups shared high rates of association, grooming, and agonistic support and low rates of agonistic interaction with one unrelated or distantly related adult male partner. Such affinity characterized small proportions of adult male-female relationships. Several males directed not only support but also aggression toward adult females with whom they sought to affiliate. All bonded males sought to copulate with their partners, and some appeared to ignore estrus in nonpartners. All females accepted copulation attempts from partners and some seemed to prefer their partners as mates. Partial synchronization of brief estrus periods together with concealed ovulation appeared to minimize chances for polygynous mating. Results support the view that the male-female pair is the fundamental social unit of E. fulvus and suggest that female partnership with individual males obviates dominance behavior, including female dominance, in this lemurid primate.

Agonistic Behavior↗

Reproductive stage and history affect the phonotactic preferences of female midwife toads, Alytes muletensis.

We investigated whether the reproductive stage or reproductive history of females affects their preferences for male calls that differ in frequency. We repeatedly tested gravid female midwife toads prior to and immediately after mating, and during ovulation. When females were ovulating they were much more consistent in their choice of call than when they were gravid but not ovulating. When females had recently mated they still showed positive phonotaxis, but did not reliably discriminate between alternative calls. This study is the first to test gravid female anurans repeatedly at different stages in their reproductive cycle (ovulating or not; pre- and postmating). It highlights the need to treat with caution the results from population-based preference studies in which individuals are tested only once, and in which variation in reproductive state is not taken into account. Copyright 2000 The Association for the Study of Animal Behaviour.

Journal Article↗

Female sticklebacks count alleles in a strategy of sexual selection explaining MHC polymorphism.

The origin and maintenance of polymorphism in major histocompatibility complex (MHC) genes in natural populations is still unresolved. Sexual selection, frequency-dependent selection by parasites and pathogens, and heterozygote advantage have been suggested to explain the maintenance of high allele diversity at MHC genes. Here we argue that there are two (non-exclusive) strategies for MHC-related sexual selection, representing solutions to two different problems: inbreeding avoidance and parasite resistance. In species prone to inadvertent inbreeding, partners should prefer dissimilar MHC genotypes to similar ones. But if the goal is to maximize the resistance of offspring towards potential infections, the choosing sex should prefer mates with a higher diversity of MHC alleles. This latter strategy should apply when there are several MHC loci, as is the case in most vertebrates. We tested the relative importance of an 'allele counting' strategy compared to a disassortative mating strategy using wild-caught three-spined sticklebacks (Gasterosteus aculeatus) from an interconnected system of lakes. Here we show that gravid female fish preferred the odour of males with a large number of MHC class-IIB alleles to that of males with fewer alleles. Females did not prefer male genotypes dissimilar to their own.

Alleles↗