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Overview: For crying out loud--dangers and opportunities of communicating in the public domain.

Communication often extends into the public domain, inviting "eavesdropping" and other forms of exploitation by other species. Conflict is frequently the result, although interspecific communication can be mutually beneficial even when interests conflict fundamentally, such as between predator and prey. However, the most complex communication systems are likely to involve species with mutual rather than with conflicting interests. When interests do conflict, exploitation of the communication systems of others may impose high costs on signalers and select for signals that are less exploitable. Other costs of communication in the public domain arise from "mistakes" (e.g., mating with the wrong species) and select for more species-specific signal structure. The articles in this section illustrate the complexity of interspecific communication and yield important insights into the dynamic nature of communication systems.

Agonistic Behavior↗

Selection of broiler chickens for a high and low incidence of tibial dyschondroplasia with observations on spondylolisthesis and twisted legs (perosis).

Radiography was used to select normal birds and affected with tibial dyschondroplasia from a commercial stock of broiler chickens. Birds were selected for three generations and mated like to like in an attempt to establish low and high incidence strains. Incidence of tibial dyschondroplasia in the low incidence strain was reduced to negligible levels while it was increased in the high incidence strain. Incidence of the defect in the high incidence strain could be influenced by diet. Observations during the selection program on the incidence of spondylolisthesis and twisted legs in birds younger than eight weeks of age indicated that these other skeletal defects were unrelated to tibial dyschondroplasia.

Animals↗

Mating in a viscous universe: the race is to the agile, not to the swift.

Charles Darwin's theory of natural selection has at its focal point the mating success of organisms. Among male animals, large body size is widely seen as the principal determinant of mating success. However, where mating takes place in a three-dimensional arena such as water, the arboreal habitat or air, small size with its concomitant aerobatic advantages might be advantageous. Despite considerable interest, the relationship between aerobatic ability and mating success has not yet been demonstrated in a single animal species. Here, we test the hypothesis that the known mating success of small male midges is due to their greater aerobatic ability. To do this, male midges collected from leks in the wild were flown and their flight paths in free flight were recorded on high-speed cameras in the laboratory. Four flight parameters that would seem relevant to male mate acquisition in flight, i.e. acceleration, maximum speed, tortuosity and turn-rate, were analysed with respect to body size. We show that, while in terms of maximum speed there was no detectable difference between small and large males, small males outperformed larger ones with respect to acceleration, tortuosity and turn-rate. We conclude that the hypothesis that small males gain their mating advantage through aerobatic superiority is consistent with the observations reported here.

Animals↗

Selection for high gamete encounter rates explains the evolution of anisogamy using plausible assumptions about size relationships of swimming speed and duration.

A previous general model describing physical constraints on gamete encounter rates was modified to incorporate assumptions that increased size causes decreased swimming speed and increased fertile period (or other proportional enhancement to gamete fertility). The analysis indicates that with moderately strong size dependence of fertile period and a range of speed dependencies, selection for high encounter rates pressures mating systems that develop any heritable difference in size between the gametes of different mating types to exaggerate the difference and evolve from isogamy to anisogamy. The smaller gamete has an optimal size, but the larger faces continuing selection for increased size. This continues to a size that is estimated to be sufficient to make pheromone production of sperm attractants practical. This mechanism then bridges the missing link between isogametes and oogamy in a previous analysis of the effectiveness of pheromones in explaining the success of male-female mating systems. The evolution and success of anisogamy and oogamy can be explained solely on the basis of physical effects on the encounter process.

Animals↗

Open-nucleus breeding strategies compared with population-wide positive assortative mating: I. Equal distribution of testing efforts.

Positive assortative mating (PAM) can enhance the additive genetic variance in a breeding population(BP). This increases the potential for gains in the production population (PP, selected subset of the BP) for recurrent selection programs in forest trees. The assortment of mates can be either: (1) by individual tree rank across the whole BP (PAM), or (2) trees of similar rank can be merged into larger hierarchical groups and then mated randomly within group ("open"-nucleus breeding,NB). The objective of this study was to compare PAM and NB in quantitative terms. The NB simulation model assumed two tiers (nucleus, main) with unrestricted migration between the tiers. Clonal tests were used to predict breeding values and test resources per mate were kept constant for all mates. Both gain and diversity were combined into a single selection criterion, "group-merit selection." Alternatives were compared over five breeding cycles by considering genetic gain and diversity in a selected PP established in a seed orchard. The assortment of mates in both alternatives enhanced additive variance and increased the additive effect in the BP, leading to additional gain in the PP. Gains generated under PAM always exceeded gains under NB. Thus, the main message from this study is that PAM in both the short- and long-term results in more gain at any target level of diversity in the PP (the breeder's target) than is achieved by the NB alternative. The optimum size of the nucleus varies with the desired level of seed orchard diversity. At lower target diversity, smaller nucleus sizes are favorable, while larger sizes result in more gain when seed orchard diversity is considered more important.

Breeding↗

The dynamics of male brooding, mating patterns, and sex roles in pipefishes and seahorses (family Syngnathidae).

Modern theory predicts that relative parental investment of the sexes in their young is a key factor responsible for sexual selection. Seahorses and pipefishes (family Syngnathidae) are extraordinary among fishes in their remarkable adaptations for paternal care and frequent occurrences of sex-role reversals (i.e., female-female competition for mates), offering exceptional opportunities to test predictions of sexual selection theory. During mating, the female transfers eggs into or onto specialized egg-brooding structures that are located on either the male's abdomen or its tail, where they are osmoregulated, aerated, and nourished by specially adapted structures. All syngnathid males exhibit this form of parental care but the brooding structures vary, ranging from the simple ventral gluing areas of some pipefishes to the completely enclosed pouches found in seahorses. We present a molecular phylogeny that indicates that the diversification of pouch types is positively correlated with the major evolutionary radiation of the group, suggesting that this extreme development and diversification of paternal care may have been an important evolutionary innovation of the Syngnathidae. Based on recent studies that show that the complexity of brooding structures reflects the degree of paternal investment in several syngnathid species, we predicted sex-role reversals to be more common among species with more complex brooding structures. In contrast to this prediction, however, both parsimony- and likelihood-based reconstructions of the evolution of sex-role reversal in pipefishes and seahorses suggest multiple shifts in sex roles in the group, independent from the degree of brood pouch development. At the same time, our data demonstrate that sex-role reversal is positively associated with polygamous mating patterns, whereas most nonreversed species mate monogamously, suggesting that selection for polygamy or monogamy in pipefishes and seahorses may strongly influence sex roles in the wild.

Animals↗

Sexual selection, Machiavellian intelligence, and the origins of psychosis.

According to Darwin's theory of sexual selection some features that differentiate the two sexes evolve by a process of "male competition" and "female choice". The sex difference in age of onset of psychotic illness in man may relate to a sexual dimorphism in cerebral organisation (the male brain being more lateralised or asymmetrical than the female brain), a difference consistent with a role for sexual selection in the evolution of the human brain. Differing criteria (reflected in a cross-culturally stable difference in mean age at marriage) in males and females for selecting personality characteristics in a mate may generate diversity in the balance of growth between the hemispheres, and this could maintain the high and relatively constant rates of psychosis in human populations.

Brain↗

Viability selection at three early life stages of the tropical tree, Platypodium elegans (Fabaceae, Papilionoideae).

Given the enormous number and high mortality of fertilized ovules in plants, it is possible that selection during the earliest stages of the life cycle plays an important role in shaping the genetic composition of plant populations. Previous research involving selection component analyses found strong evidence for viability selection in annual plant species. Yet despite this evidence, few attempts have been made to identify the magnitude and timing of viability selection as well as the mechanisms responsible for mortality among genotypes. Platypodium elegans, a Neotropical tree with high rates of early fruit mortality, represents an opportunity to study viability selection at a level of discernment not previously possible. Microsatellite markers were used to analyze the genetic composition of aborted embryos, as well as mature seeds and seedlings of the same cohort. While selection resulted in an overall decrease in self-fertilized progeny across each life stage, the greatest change in the genetic composition of progeny occurred between mature seeds and established seedlings. This suggested that inbreeding depression, and not late-acting self-incompatibility, was responsible for early selection. An investigation of the mature seed stage revealed that self-fertilized seeds weigh significantly less than outcrossed seeds. The result of this early selection conceals the mixed-mating system and high levels of inbreeding depression in Platypodium elegans, resulting in an apparently outcrossed adult population that does not differ significantly from Hardy-Weinberg expectations.

Alleles↗

Genetic diversity at a single locus under viability selection and facultative apomixis: equilibrium structure and deviations from Hardy-Weinberg frequencies.

We extensively analyze the maintenance of genetic variation and deviations from Hardy-Weinberg frequencies at a diallelic locus under mixed mating with apomixis and constant viability selection. Analytical proofs show that: (1) at most one polymorphic equilibrium exists, (2) polymorphism requires overdominant or underdominant selection, and (3) a simple, modified overdominance condition is sufficient to maintain genetic variation. In numerical analyses, only overdominant polymorphic equilibria are stable, and these are stable whenever they exist, which happens for approximately 78% of random fitness and mating parameters. The potential for maintaining both alleles increases with increasing apomixis or outcrossing and decreasing selfing. Simulations also indicate that equilibrium levels of heterozygosity will often be statistically indistinguishable from Hardy-Weinberg frequencies and that adults, not seeds, should usually be censused to maximize detecting deviations. Furthermore, although both censuses more often have an excess rather than a deficit of heterozygotes, analytical sign analyses of the fixation indices prove that, overall, adults are more likely to have an excess and seeds a deficit at equilibrium.

Adult↗

The a-factor transporter (STE6 gene product) and cell polarity in the yeast Saccharomyces cerevisiae.

STE6 gene product is required for secretion of the lipopeptide mating pheromone a-factor by Saccharomyces cerevisiae MATa cells. Radiolabeling and immunoprecipitation, either with specific polyclonal antibodies raised against a TrpE-Ste6 fusion protein or with mAbs that recognize c-myc epitopes in fully functional epitope-tagged Ste6 derivatives, demonstrated that Ste6 is a 145-kD phosphoprotein. Subcellular fractionation, various extraction procedures, and immunoblotting showed that Ste6 is an intrinsic plasma membrane-associated protein. The apparent molecular weight of Ste6 was unaffected by tunicamycin treatment, and the radiolabeled protein did not bind to concanavalin A, indicating that Ste6 is not glycosylated and that glycosylation is not required either for its membrane delivery or its function. The amino acid sequence of Ste6 predicts two ATP-binding folds; correspondingly, Ste6 was photoaffinity-labeled specifically with 8-azido-[alpha-32P]ATP. Indirect immunofluorescence revealed that in exponentially growing MATa cells, the majority of Ste6 showed a patchy distribution within the plasma membrane, but a significant fraction was found concentrated in a number of vesicle-like bodies subtending the plasma membrane. In contrast, in MATa cells exposed to the mating pheromone alpha-factor, which markedly induced Ste6 production, the majority of Ste6 was incorporated into the plasma membrane within the growing tip of the elongating cells. The highly localized insertion of this transporter may establish pronounced anisotropy in a-factor secretion from the MATa cell, and thereby may contribute to the establishment of the cell polarity which restricts partner selection and cell fusion during mating to one MAT alpha cell.

ATP-Binding Cassette Transporters↗

Release of a potent polymorphonuclear leukocyte chemoattractant is regulated by white-opaque switching in Candida albicans.

Previous studies employing transmembrane assays suggested that Candida albicans and related species, as well as Saccharomyces cerevisiae, release chemoattractants for human polymorphonuclear leukocytes (PMNs). Because transmembrane assays do not definitively distinguish between chemokinesis and chemotaxis, single-cell chemotaxis assays were used to confirm these findings and test whether mating-type or white-opaque switching affects the release of attractant. Our results demonstrate that C. albicans, C. dubliniensis, C. tropicalis, C. parapsilosis, and C. glabrata release bona fide chemoattractants for PMNs. S. cerevisiae, however, releases a chemokinetic factor but not a chemoattractant. Characterization of the C. albicans chemoattractant revealed that it is a peptide of approximately 1 kDa. Whereas the mating type of C. albicans did not affect the release of chemoattractant, switching did. White-phase cells released chemoattractant, but opaque-phase cells did not. Since the opaque phase of C. albicans represents the mating-competent phenotype, it may be that opaque-phase cells selectively suppress the release of chemoattractant to facilitate mating.

Candida albicans↗

Molecular parentage analysis in experimental newt populations: the response of mating system measures to variation in the operational sex ratio.

Molecular studies of parentage have been extremely influential in the study of sexual selection in the last decade, but a consensus statistical method for the characterization of genetic mating systems has not yet emerged. Here we study the utility of alternative mating system measures by experimentally altering the intensity of sexual selection in laboratory-based breeding populations of the rough-skinned newt. Our experiment involved skewed sex ratio (high sexual selection) and even sex ratio (low sexual selection) treatments, and we assessed the mating system by assigning parentage with microsatellite markers. Our results show that mating system measures based on Bateman's principles accurately reflect the intensity of sexual selection. One key component of this way of quantifying mating systems is the Bateman gradient, which is currently underutilized in the study of genetic mating systems. We also compare inferences based on Bateman's principles with those obtained using two other mating system measures that have been advocated recently (Morisita's index and the index of resource monopolization), and our results produce no justification for the use of these alternative measures. Overall, our results show that Bateman's principles provide the best available method for the statistical characterization of mating systems in nature.

Animals↗

Selection for body weight at eight weeks of age in dwarf and normal meat-type chickens originating from a common control population background.

A study was conducted to evaluate responses to selection for 8-week body weight in normal and dwarf meat-type broiler lines originating from the same nonselected randombred population. Following three generations of selection, normal and dwarf dams were mated to males from a commercial broiler stock to compare strain cross performance of progeny from the two dam groups. The mean realized heritabilities across four generations for 8-week body weight, estimated by dividing the cumulative responses by the cumulative selection differentials, were .69 for normal lines and .59 for dwarf lines. Realized estimates from regression of population means on cumulative selection differentials were .78 +/- .10 and .74 +/- .02 for normal lines and .70 +/- .14 and .59 +/- .19 for dwarf lines. Although these estimates were not significantly different, realized heritabilities were consistently greater in normal than in dwarf lines across generations. The smaller heritability estimates observed in dwarf lines were compensated for by larger selection intensities (i), which resulted in similar selection responses in 8-week body weight for normal and dwarf genotypes. The fertility of normal and dwarf females mated to commercial broiler males was identical. However, the hatchability of fertile eggs was approximately 10% higher for dwarfs than for normal females. Body weights (8-week) of male progeny from dwarf dams mated to commercial males were 3% smaller than comparable males from normal dams. There were no differences between 8-week body weights of female progeny from normal and dwarf dams mated to commercial males.

Animals↗

The evolution of genes for pigmentation in African cichlid fishes.

The cichlid fishes in the East African Great Lakes exhibit a great diversity of color patterns, presumably as adaptations to species-specific habitats and/or due to the action of sexual selection on color for species discrimination or female mate choice. To elucidate the mechanisms underlying such pigment pattern diversity, we cloned the cichlid homologs of tyrosinase, endothelin receptor b1, mitf, and Aim1 that previously had been cloned and characterized from pigmentation mutants of zebrafish and medaka fish. Gene sequence analysis among five cichlid species from the Great Lakes shows that the evolutionary rate of amino acid replacement in mitf is the highest of these four genes. We then compared the mitf amino acid replacement rates between species from the lacustrine and tilapiine/steatocranus lineages, and between Lake Malawi- and Victoria-haplochromine cichlids and Lake Tanganyika Lamprologini. We show that the evolutionary rate within the lacustrine lineage is twice that of the tilapiine/steatocranus lineage, but that rates for the Malawi-Victoria haplochromine and Lamprologini lineages are almost the same. These results suggest that the accelerated evolution of mitf might have occurred concomitantly with pigment pattern diversification in Great Lakes species, but not necessarily correlated with species under intense sexual selection on male mating color via female mate choice. Finally, we characterized a novel alternatively spliced variant of cichlid mitf that is similar to a mammalian mitf splice variant generated using alternate splice sites. We suggest that this new variant in cichlids, like that in mammals, encodes an MITF transcriptional factor having higher relative DNA binding affinity. These data provide a novel example of functional convergence in which a particular splice variant is independently generated via alternative splicing of a specific gene in different lineages.

Alternative Splicing↗

Concentrations of fat and plasma 5alpha-androstenone and plasma testosterone in boars selected for rate of body weight gain and thickness of back fat during growth, sexual maturation and after mating.

The levels of 5alpha-androstenone and testosterone in peripheral plasma and 5alpha-androstenone in fat collected from 18 boars selected according to thickness of back fat and rate of gain were evaluated. Sampling was started when the animals were about 100 days of age and samples were collected every 14 days until the boars were about 240 days old. The steroid profiles varied considerably between boars, although an increase in steroid levels was found in most animals with the onset of adolescence. The regression line for the level of 5alpha-androstenone in fat and the age of the animals in days was found to be y = 0.02x-1-31. High coefficients of correlation were found between the concentration of 5alpha-androstenone in peripheral plasma and fat (r = 0-78) and between levels of 5alpha-androstenone and testosterone in peripheral plasma (r = 0-64). Levels of 5alpha-androstenone in peripheral plasma above about 15 ng/ml were usually accompanied by a heavy accumulation of 5alpha-androstenone in adipose tissue. The use of four of the boars for mating resulted in increased steroid levels in peripheral plasma and fat. Significantly higher levels of 5alpha-androstenone and testosterone were found in the peripheral plasma of boars selected for fatness and a low rate of gain than in boars selected for low back-fat thickness and a high rate of gain.

Adipose Tissue↗

Male parental care, differential parental investment by females and sexual selection.

Males play a variable parental role in reproduction, ranging from no male parental care to extensive male care. Females may acquire either direct or indirect fitness benefits from their mate choice, and direct fitness benefits include male parental care. Theoreticians have traditionally emphasized direct fitness benefits to females in species with extensive male parental care. We review the literature and show extensive variation in the patterns of male care, related to the attractiveness of males to females. At one extreme of this continuum, females invest differentially in parental care, investing more when paired with attractive males. The costs of female parental care and other aspects of parental investment may be balanced by benefits in terms of more attractive sons and/or more viable offspring. At the other extreme, in species with extensive direct fitness benefits, males with preferred sexual phenotypes provide the largest relative share of parental care. A comparative study of birds revealed that the extent of the differential female parental investment was directly related to the frequency of extra-pair paternity. Since extra-pair paternity may arise mainly as a consequence of female choice for indirect fitness benefits, this result supports our prediction that differential parental investment is prevalent in species where females benefit indirectly from their mate choice. The consequences for sexual selection theory of these patterns of male care in relation to male attractiveness are emphasized. Copyright 1998 The Association for the Study of Animal Behaviour. Copyright 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Nest building is a sexually selected behaviour in the barn swallow.

Females may use male nest building to assess male parental quality, and nest size would then be a sexually selected trait. In the barn swallow, Hirundo rustica, females select their partner by his tail length, a character believed to signal good genes. Both sexes participate in nest building, although male participation is negatively related to his attractiveness as reflected by tail length. We tested the hypothesis that nest building is a sexually selected trait: females paired with males of high parental quality (as shown by the male during nest building) may obtain a mate providing large amounts of parental investment, while, as has been shown previously, females mated to attractive (long-tailed) males will acquire mates with good genetic quality. Therefore, since nest building in barn swallows occurs after mating, we predicted a postmating sexual selection process by which the female invests differentially in reproduction depending on the male's nest-building effort (reflecting his willingness to invest in reproduction). The volume of material in a nest was related to the male's contribution to nest building and, in agreement with our hypothesis, in a multiple regression analysis, male tail length and nest material volume were negatively related to laying date and positively to female investment in reproduction (total number of eggs laid during the breeding season). Moreover, females paired with long-tailed males (which contribute very little to nest building), but using the same amount of nest material as females paired with short-tailed males, reduced the thickness of the nest and hence increased its capacity. Therefore, in the barn swallow two different traits appear to be sexually selected: tail length of males owing to the good genes process and nest-building ability owing to the good parent process. (c) 1998 The Association for the Study of Animal Behaviour.

Journal Article↗

Sexual selection in Drosophila silvestris of Hawaii.

Previous discovery that Drosophila melanogaster females tend to discriminate in mating against phenotypes of earliest courting males prompted a study of the Hawaiian species D. silvestris. Tibial bristle variation in males from opposite coasts of the island of Hawaii functions in courtship, and the possibility that females can distinguish males differing in the tibial trait is explored. Mating tests, designed to give each female and male an alternative choice between two individuals of opposite sex every 30 min, consisted of intrapopulation tests with a strain derived from an eastern (Kilauea) population and interpopulation tests between that strain and one derived from a western (Kahuku) population. Males were given initial combat tests, with "winners" then used in mating (except one test with "loser" males). Matings (52-55%) were classified into categories according to the readiness of the female to mate and sequence of courtship. Low-threshold females (accepting the first male after less than four courtship bouts) occurred at 30-35%. Among intrapopulational tests, females (with higher threshold) accepted first- and second-courting males about equally (25:36, respectively), but for male success in mating, the winning of initial intermale combats and the uniformity of courtship effort tended to be important criteria. Among interpopulation tests, homogamic matings were nearly equal (25% each), but heterogamic matings contrasted in that Kilauea females were reluctant to mate with Kahuku males (14%), while reciprocal matings occurred most frequently (34%). Females favored males second to court, particularly when a Kilauea male (with extra tibial bristles) was the second male. Thus a morphological feature likely to be influential in mating is demonstrated to be so; and sexual selection is operating via male-male combat plus discrimination in favor of particular opposite-sex individuals in this species.

Journal Article↗