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Sexual selection: harem size and the variance in male reproductive success.

Sexual selection is potentially stronger than natural selection when the variance in male reproductive fitness exceeds all other components of fitness variance combined. However, measuring the variance in male reproductive fitness is difficult when nonmating males are absent, inconspicuous, or otherwise difficult to find. Omitting the nonmating males inflates estimates of average male reproductive success and diminishes the variance, leading to underestimates of the potential strength of sexual selection. We show that, in theory, the proportion of the total variance in male fitness owing to sexual selection is approximately equal to H, the mean harem size, as long as H is large and females are randomly distributed across mating males (i.e., Vharem=H). In this case, mean harem size not only provides an easy way to estimate the potential strength of sexual selection but also equals the opportunity for sexual selection, I(mates). In nature, however, females may be overdispersed with Vharem H. We show that H+(k-1) is a good measure of the opportunity for sexual selection, where k is the ratio Vharem/H. A review of mating system data reveals that in nature the median ratio for Vharem/H is 1.04, but as H increases, females tend to become more aggregated across mating males with V(harem) two to three times larger than H.

Animals↗

Nuptial feeding of spermless spermatophores in the Hawaiian swordtail cricket, Laupala pacifica (Gryllidae: Triginodiinae).

Crickets in the genus Laupala (subfamily Trigonidiinae) have an elaborate courtship system, defined by a highly ritualized serial transfer of multiple spermatophores. Males produce multiple "micro" spermatophores followed by a final "macro" spermatophore during a single mating bout. Remarkably, the microspermatophores of L. cerasina, the first species whose mating system was studied in detail, were discovered to be spermless. However, in a study of another species, L. pacifica, sperm transfer was reported after every copulation suggesting that L. pacifica microspermatophores contain sperm. The presence or absence of sperm in the microspermatophore has important implications for the evolution of this exaggerated courtship system and the origin of nuptial gifts. In this study, we systematically examined L. pacifica spermatophore contents for sperm using a fluorescent nuclear stain. We detected sperm only in macrospermatophores. This finding suggests that spermless microspermatophores are typical for Laupala; thus, to determine the origin of this highly modified phenotype will require comparative analyses with closely related outgroups that exhibit less exaggerated courtship systems.

Animals↗

Female copying increases the variance in male mating success.

Theoretical models of sexual selection assume that females choose males independently of the actions and choice of other individual females. Variance in male mating success in promiscuous species is thus interpreted as a result of phenotypic differences among males which females perceive and to which they respond. Here we show that, if some females copy the behavior of other females in choosing mates, the variance in male mating success and therefore the opportunity for sexual selection is greatly increased. Copying behavior is most likely in non-resource-based harem and lek mating systems but may occur in polygynous, territorial systems as well. It can be shown that copying behavior by females is an adaptive alternative to random choice whenever there is a cost to mate choice. We develop a statistical means of estimating the degree of female copying in natural populations where it occurs.

Animals↗

Gibberella xylarioides sensu lato from Coffea canephora: a new mating population in the Gibberella fujikuroi species complex.

Gibberella xylarioides Heim & Saccas (presumed anamorph, Fusarium xylarioides Steyaert) is the causal agent of coffee wilt disease, an economically important tracheomycosis in Africa. In vitro crosses carried out with Congolese, Ugandan, and Tanzanian single-ascospore/conidial isolates originating from diseased Coffea canephora/excelsa demonstrated a heterothallic mating system, controlled by a single locus with two alleles, MAT-1 and MAT-2. Compatible isolates produced fertile perithecia within 2 to 8 weeks after mating. Mating type (MAT) was characterized by PCR with primer pairs previously developed for the Gibberella fujikuroi species complex (GFC) and for Fusarium oxysporum. All strains analyzed were morphologically identical and corresponded to Booth's description of the "female" F. xylarioides strain. Based on crossing results and MAT-2/translation elongation 1-alpha (tef) sequence data, G. xylarioides, as currently understood, is demonstrated to encompass at least three "groups": G. xylarioides sensu strictu Ia, defined hitherto by two "historical" West African strains originating from the severe 1930s to 1950s epidemic (CBS 25852 and CBS 74979); G. xylarioides sensu strictu Ib, defined by two "historical" Central African lowland strains (DSMZ 62457 and ATCC 15664); and G. xylarioides sensu lato II, containing Congolese, Ugandan, and Tanzanian C. canephora/excelsa isolates. Infertility of crosses between the coffee wilt pathogen and known GFC mating populations demonstrates that G. xylarioides sensu lato constitutes a new biological species within the G. fujikuroi complex. MUCL 44532/MUCL 43887 and MUCL 35223/MUCL 44549 are proposed as G. xylarioides sensu lato II MAT-1/MAT-2 reference mating type tester strains.

Citrus sinensis↗

Chemical and visual communication during mate searching in rock shrimp.

Mate searching in crustaceans depends on different communicational cues, of which chemical and visual cues are most important. Herein we examined the role of chemical and visual communication during mate searching and assessment in the rock shrimp Rhynchocinetes typus. Adult male rock shrimp experience major ontogenetic changes. The terminal molt stages (named "robustus") are dominant and capable of monopolizing females during the mating process. Previous studies had shown that most females preferably mate with robustus males, but how these dominant males and receptive females find each other is uncertain, and is the question we examined herein. In a Y-maze designed to test for the importance of waterborne chemical cues, we observed that females approached the robustus male significantly more often than the typus male. Robustus males, however, were unable to locate receptive females via chemical signals. Using an experimental set-up that allowed testing for the importance of visual cues, we demonstrated that receptive females do not use visual cues to select robustus males, but robustus males use visual cues to find receptive females. Visual cues used by the robustus males were the tumults created by agitated aggregations of subordinate typus males around the receptive females. These results indicate a strong link between sexual communication and the mating system of rock shrimp in which dominant males monopolize receptive females. We found that females and males use different (sex-specific) communicational cues during mate searching and assessment, and that the sexual communication of rock shrimp is similar to that of the American lobster, where females are first attracted to the dominant males by chemical cues emitted by these males. A brief comparison between these two species shows that female behaviors during sexual communication contribute strongly to the outcome of mate searching and assessment.

Analysis of Variance↗

Sow productivity comparisons for four breeds of swine: purebred and crossbred litters.

Data were collected from 366 litters produced in a four-breed (Duroc, Yorkshire, Spotted and Landrace) diallel mating system. Purebred and crossbred litters were farrowed during five consecutive (fall and spring) farrowing seasons (fall 1976 through fall 1978). Litters were produced by randomly mating boars of each breed to at least one female of each breed. Litter size and weight at birth, litter size and weight at 42 d and pig survival to 42 d were analyzed. There were significant parity effects for all traits except survival rate, and significant year and season effects for litter size at birth and 42 d and litter weight at 42 d. Large breed of dam differences were detected for most sow productivity traits. Litters from Spotted females were smaller and lighter in weight. Landrace females were most productive in terms of litter weight at weaning and proportion of their pigs raised from birth to weaning. Breed of sire differences were not significant for any trait. Significant heterosis estimates were obtained for litter size and weight at weaning and for survival rate. Crossbred litters had .79 more pigs, were 11.72 kg heavier at weaning and had a 5.56% greater survival rate than the average of all purebreds.

Animals↗

Natural variation in expression of self-incompatibility in Arabidopsis thaliana: implications for the evolution of selfing.

The switch from an out-crossing to a self-fertilizing mating system is one of the most prevalent evolutionary trends in plant reproduction and is thought to have occurred repeatedly in flowering plants. However, little is known about the evolution of self-fertility and the genetic architecture of selfing. Here, we establish Arabidopsis thaliana as a model for genetic analysis of the switch to self-fertility in the crucifer family, where the ancestral out-crossing mode of mating is determined by self-incompatibility (SI), a genetic system controlled by the S locus. We show that A. thaliana ecotypes exhibit S-locus polymorphisms and differ in their ability to express the SI trait upon transformation with S-locus genes derived from the obligate out-crosser Arabidopsis lyrata. Remarkably, at least one ecotype was reverted to a stable, self-incompatible phenotype identical to that of naturally self-incompatible species. These ecotype differences are heritable and reflect the fixation in different A. thaliana populations of independent mutations that caused or enforced the switch to self-fertility. Their continued analysis promises to identify the loci that were the targets of natural selection for selfing and to contribute to a mechanistic understanding of the SI response.

Arabidopsis↗

Scent marking in two western Amazonian populations of woolly monkeys (Lagothrix lagotricha).

We describe patterns of scent marking observed in two wild populations of lowland woolly monkeys that were the subjects of long-term studies in the westernmost portion of the Amazon basin. The woolly monkeys engaged primarily in two types of scent marking: chest rubbing and anogenital rubbing. In both study populations, males and females performed both types of scent marking, but males chest-rubbed more commonly than females, while females engaged in more anogenital rubbing. We evaluated two nonexclusive hypotheses for the function of scent marking by woolly monkeys: 1) that scent marking is used in sociosexual contexts, and 2) that scent marking is used to convey information about occupancy of or willingness to defend an area from conspecifics in other social groups. We found no association between the occurrence of scent-marking behavior and location within the home range, but did find that scent marking occurred more commonly than expected on days when copulations, mating solicitations, and intergroup encounters were observed. Additionally, mating activity and chest rubbing were highly correlated across the yearly cycle, even when the potentially confounding variable of ripe fruit availability was controlled for. In woolly monkeys, overt male-male competition is rare and female choice is an important part of the mating system. Our results are most consistent with the idea that scent marking plays a role in advertising male quality or competitive ability, and perhaps in coordinating mating activity.

Animal Communication↗

Quantitative genetics of female choice in an ultrasonic pyralid moth, Achroia grisella: variation and evolvability of preference along multiple dimensions of the male advertisement signal.

The mating system of Achroia grisella (Lepidoptera: Pyralidae) is characterized by male ultrasonic advertisement signalling to which females orientate. Although males provide no direct, somatic benefits to their mates, females prefer males whose signal characters are more exaggerated than the population means. Previous studies showed that the signal characters influencing mate attraction are highly repeatable and heritable. We measured the phenotypic and additive genetic variances (heritability) of female preference in A. grisella, as this additive genetic variance is one of the genetic assumptions of indirect models of sexual selection. We determined the preference index of female A. grisella by repeated phonotaxis trials in which a choice of simulated male signals was presented. These playback experiments showed that female preference indices varied but were repeatable within individuals. Specifically, females differ in the relative importance of the several signal characters during mate assessment. A subsequent half-sib breeding design revealed an amount of additive genetic variance for the female preference index (h(s)2 = 0.212, SE = 0.1347, P = 0.0611; CVA = 0.1826). Our study highlights the importance of careful preparation of test signals and experimental design for quantifying individual variation in (female) preference along multiple signal dimensions.

Animals↗

High levels of genetic monogamy in the group-living Australian lizard Egernia stokesii.

The Australian lizard Egernia stokesii lives in spatially and temporally stable groups of up to 17 individuals. We have recently shown that these groups are comprised of breeding partners, their offspring and, in some cases, highly related adults, providing the first genetic evidence of a family structure in any lizard species. Here we investigated the mating system of E. stokesii using data from up to eight polymorphic microsatellite loci and tested the hypothesis that breeding partners are monogamous both within and between mating seasons. Among 16 laboratory-born litters from field collected gravid females from two sites in South Australia, 75% had a single male parent and no male contributed to more than one litter, indicating a high level of genetic monogamy within a season. Additional analyses of field caught individuals, captured between 1994 and 1998, enabled assignment of parentage for 70 juveniles and subadults. These data showed that most young (88.6%) had both parents from within the same group and that high proportions of males (88.9%) and females (63.6%) have multiple cohorts of offspring only with the same partner. Our results suggest that monogamy both within and between seasons is a common mating strategy of E. stokesii and that breeding partners maintain stable associations together and with multiple cohorts of their offspring over periods of up to at least 5 years.

Animals↗

Mating types and sexual development in filamentous ascomycetes.

The progress made in the molecular characterization of the mating types in several filamentous ascomycetes has allowed us to better understand their role in sexual development and has brought to light interesting biological problems. The mating types of Neurospora crassa, Podospora anserina, and Cochliobolus heterostrophus consist of unrelated and unique sequences containing one or several genes with multiple functions, related to sexuality or not, such as vegetative incompatibility in N. crassa. The presence of putative DNA binding domains in the proteins encoded by the mating-type (mat) genes suggests that they may be transcriptional factors. The mat genes play a role in cell-cell recognition at fertilization, probably by activating the genes responsible for the hormonal signal whose occurrence was previously demonstrated by physiological experiments. They also control recognition between nuclei at a later stage, when reproductive nuclei of each mating type which have divided in the common cytoplasm pair within the ascogenous hyphae. How self is distinguished from nonself at the nuclear level is not known. The finding that homothallic species, able to mate in the absence of a partner, contain both mating types in the same haploid genome has raised more issues than it has resolved. The instability of the mating type, in particular in Sclerotinia trifolorium and Botrytinia fuckeliana, is also unexplained. This diversity of mating systems, still more apparent if the yeasts and the basidiomycetes are taken into account, clearly shows that no single species can serve as a universal mating-type model.

Amino Acid Sequence↗

Effects of prenatal and postnatal fraternity size on long-term reproduction in mice.

Effects of prenatal and postnatal fraternity size (size of litter in which an animal develops prior to birth or is reared following birth) on long-term reproduction were studied by rearing 178 female ICR mice in standardized prenatal and postnatal fraternities. Three levels of prenatal and postnatal fraternity sizes were used in a 3 x 3 factorial experiment. Prenatal fraternity size was standardized by selectively terminating fetal development in pregnant females carrying at least 14 conceptuses. Prenatal fraternities were standardized to either 6, 10 or 14 fetuses, and postnatal fraternities were standardized by randomly assigning individuals to nurse litters of 5, 10 or 15 pups. Prenatal fraternity size negatively affected average pup weight at birth (P less than .05) but had little subsequent effect on growth or reproduction. Postnatal fraternity size negatively affected weight at weaning (P less than .01), with mice reared in smaller postnatal fraternities being heavier than those reared in larger fraternities. Following weaning, mice reared in smaller fraternities gained weight less rapidly (P less than .01) but still tended to be heavier at maturity (P = .11). Vaginal opening occurred at older ages in females reared in larger postnatal litters (P less than .01). An interval mating system was used to examine fraternity size effects on long-term reproduction. Females were exposed to males six times at 8-wk intervals with initial mating at 7 wk of age. Postnatal fraternity size and age at mating jointly affected litter size (P less than .05).(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Patterns of mating and mitochondrial DNA inheritance in the agaric Basidiomycete Coprinus cinereus.

Patterns of mating and mitochondrial DNA (mtDNA) inheritance were investigated for the Basidiomycete, Coprinus cinereus in order to better understand the relationship of reproductive biology and mtDNA evolution in fungi. Results showed that the unique mating system of basidiomycetes can lead to the formation of mitochondrial mosaics (i.e., colonies composed of sectors differing in mtDNA). Mitochondria do not migrate along with nuclei during mating. Intracellular mixed or recombinant mtDNA molecules were not observed. Interestingly, it was found that mating asymmetry, caused by nonreciprocal nuclear migration, may be an important part of the reproductive biology of C. cinereus.

Agaricales↗

Resource defense and juvenile hormone: the "challenge hypothesis" extended to insects.

As predicted for vertebrates by the challenge hypothesis, the endocrine system of invertebrates can respond to social stimuli to modulate aggression. Testosterone (T) is generally considered to moderate aggression associated with reproduction, i.e. the establishment of breeding territories, mate guarding and offspring defense; juvenile hormone (JH) serves an analogous function in burying beetles. Hemolymph titers of JH increase significantly in Nicrophorus orbicollis, a species with facultative biparental care, when challenged by an intruder to defend their resource. During the first 12 h after the discovery of a carcass, the necessary breeding resource, competition is intrasexual, and JH of males responds only to a challenge by males, and JH of females responds only to a challenge by female intruders. After this period, competition is intersexual, and JH increases significantly in both males and females challenged by an intruder of either sex. In contrast, JH titers in a nonparental species are much higher throughout the breeding season, and neither males nor females respond hormonally to an intruder. These findings support the challenge hypothesis and suggest that mating systems and breeding strategies can promote plastic responses in insect, as well as vertebrate, endocrine systems.

Aggression↗

Genetic exchange in Trypanosoma brucei.

The discovery of genetic exchange in African trypanosomes belonging to the Trypanosoma brucei group is an important development in our understanding of these organisms. Genetic exchange is a feature of major importance in relation to population structure and speciation. Furthermore, a convenient laboratory-based mating system would be of considerable value as a tool in trypanosomiasis research. It is now known that although cyclical development of trypanosomes within the tsetse fly does not require mating to occur, genetic exchange may take place under Conditions in which genetically distinct trypanosomes develop within the same fly. During the past few years there has been a considerable body of research on laboratory crosses, and a number of controversial and apparently contradictory models of the mechanism of genetic exchange and the ploidy of different life cycle stages have been proposed. In this article, Andy Tait and Mike Turner review the present state of knowledge regarding gene exchange in T. brucei, and attempt to reconcile the various observations and models available.

Journal Article↗

Male chimpanzees prefer mating with old females.

Cross-cultural studies indicate that women's sexual attractiveness generally peaks before motherhood and declines with age. Cues of female youth are thought to be attractive because humans maintain long-term pair bonds, making reproductive value (i.e. future reproductive potential) particularly important to males. Menopause is believed to exaggerate this preference for youth by limiting women's future fertility. This theory predicts that in species lacking long-term pair bonds and menopause, males should not exhibit a preference for young mates. We tested this prediction by studying male preferences in our closest living relative, the chimpanzee (Pan troglodytes). We show that despite their promiscuous mating system, chimpanzee males, like humans, prefer some females over others. However, in contrast to humans, chimpanzee males prefer older, not younger, females. These data robustly discriminate patterns of male mate choice between humans and chimpanzees. Given that the human lineage evolved from a chimpanzee-like ancestor, they indicate that male preference for youth is a derived human feature, likely adapted from a tendency to form unusually long term mating bonds.

Age Factors↗

A major evolutionary transition to more than two sexes?

Two recently discovered cases of genetic caste determination in social insects might provide the first example of a major evolutionary transition from two to more than two sexes. I argue here that the system can be interpreted as comprising primarily individuals requiring gametes from three parental types and having four sexes from the perspective of demographic extinction. Additionally, I show how this mating system can be seen as a major evolutionary transition. For these populations, it is apparent that the mechanism for a three- or four-sex system does not lie within the myriad of possible arrangements of chromosomes within individuals, but at the next level of evolutionary complexity, with the arrangement of chromosomes among individuals within a social system.

Journal Article↗