Anisogamy,sexual selection,and the evolution and maintenance of sex

Summary In the present paper we distinguish between two aspects of sexual reproduction. Genetic recombination is a universal features of the sexual process. It is a primitive condition found in simple, single-celled organisms, as well as in higher plants and animals. Its function is primarily to repair genetic damage and eliminate deleterious mutations. Recombination also produces new variation, however, and this can provide the basis for adaptive evolutionary change in spatially and temporally variable environments.The other feature usually associated with sexual reproduction, differentiated male and female roles, is a derived condition, largely restricted to complex, diploid, multicellular organisms. The evolution of anisogamous gametes (small, mobile male gametes containing only genetic material, and large, relatively immobile female gametes containing both genetic material and resources for the developing offspring) not only established the fundamental basis for maleness and femaleness, it also led to an asymmetry between the sexes in the allocation of resources to mating and offspring. Whereas females allocate their resources primarily to offspring, the existence of many male gametes for each female one results in sexual selection on males to allocate their resources to traits that enhance success in competition for fertilizations. A consequence of this reproductive competition, higher variance in male than female reproductive success, results in more intense selection on males.The greater response of males to both stabilizing and directional selection constitutes an evolutionary advantage of males that partially compensates for the cost of producing them. The increased fitness contributed by sexual selection on males will complement the advantages of genetic recombination for DNA repair and elimination of deleterious mutations in any outcrossing breeding system in which males contribute only genetic material to their offspring. Higher plants and animals tend to maintain sexual reproduction in part because of the enhanced fitness of offspring resulting from sexual selection at the level of individual organisms, and in part because of the superiority of sexual populations in competition with asexual clones.     

Differential selection between the sexes and selection for sex

Anisogamy is known to generate an important cost for sexual reproduction (the famous "twofold cost of sex"). However, male-female differences may have other consequences on the evolution of sex, due to the fact that selective pressures may differ among the sexes. On the one hand, intralocus sexual conflict should favor asexual females, which can fix female-beneficial, male-detrimental alleles. On the other hand, it has been suggested repeatedly that sexual selection among males may help to purge the mutation load, providing an advantage to sexual females. However, no analytical model has computed the strength of selection acting on a modifier gene affecting the frequency of sexual reproduction when selection differs between the sexes. In this article, we analyze a two-locus model using two approaches: a quasi-linkage-equilibrium (QLE) analysis and a local stability analysis, whose predictions are verified using a multilocus simulation. We find that costly sex can be maintained when selection is stronger in males than in females, but acts in the same direction in both. Complete asexuality, however, evolves under any other form of selection. Finally, we discuss how experimental measurements of fitness variances and covariances between sexes could be used to determine the overall direction and strength on selection for sex arising from differences in selection between males and females.     

Evidence for the equal resilience of Triodia spp. (Poaceae), from different functional groups, to frequent fire dating back to the late Pleistocene

Species with different regenerative responses to fire are hypothesised to coexist by utilising the different temporal and spatial niche opportunities created by the stochasticity of the fire regime. This is strongly supported by observations of instability of species'' presence and abundance at the local scale while these are stable at the community scale. However, observations of species coexistence in fire-prone communities are limited to several decades only. To improve the robustness of this hypothesis, coalescent analysis, using chloroplast microsatellites, was undertaken on three sympatric species of Triodia from different functional groups in the fire-prone Kimberley region of Western Australia. The results inferred that T. bitextura, an obligate resprouter, Triodia sp., an obligate seeder, and T. epactia, a facultative resprouter, had mean T_mrca values of 65k, 40k and 111k generations, respectively. Using a mutation rate of 3.2 × 10⁻⁵ and a generation time of 5 years gave T_mrca values of 436k, 203k and 556 k years, respectively. These results provide evidence for the coexistence of these species to the same fire regime dating back to the late Pleistocene. It also demonstrates the long-term resilience of an obligate seeder, Triodia sp., in a frequently burnt environment at the community scale.     

Relationship between leaf traits and fire-response strategies in shrub species of a mountainous region of south-eastern Australia

Background and Aims

Resprouting and seed recruitment are important ways in which plants respond to fire. However, the investments a plant makes into ensuring the success of post-fire resprouting or seedling recruitment can result in trade-offs that are manifested in a range of co-occurring morphological, life history and physiological traits. Relationships between fire-response strategies and other traits have been widely examined in fire-prone Mediterranean-type climates. In this paper, we aim to determine whether shrubs growing in a non-Mediterranean climate region exhibit relationships between their fire-response strategy and leaf traits.

Methods

Field surveys were used to classify species into fire-response types. We then compared specific leaf area, leaf dry-matter content, leaf width, leaf nitrogen and carbon to nitrogen ratios between (a) obligate seeders and all other resprouters, and (b) obligate seeders, facultative resprouters and obligate resprouters.

Key Results

Leaf traits only varied between fire-response types when we considered facultative resprouters as a separate group to obligate resprouters, as observed after a large landscape-scale fire. We found no differences between obligate seeders and obligate resprouters, nor between obligate seeders and resprouters considered as one group.

Conclusions

The results suggest that facultative resprouters may require a strategy of rapid resource acquisition and fast growth in order to compete with species that either resprout, or recruit from seed. However, the overall lack of difference between obligate seeders and obligate resprouters suggests that environmental factors are exerting similar effects on species'' ecological strategies, irrespective of the constraints and trade-offs that may be associated with obligate seeding and obligate resprouting. These results highlight the limits to trait co-occurrences across different ecosystems and the difficulty in identifying global-scale relationships amongst traits.     

Intralocus sexual conflict and offspring sex ratio

Males and females frequently have different fitness optima for shared traits, and as a result, genotypes that are high fitness as males are low fitness as females, and vice versa. When this occurs, biasing of offspring sex-ratio to reduce the production of the lower-fitness sex would be advantageous, so that for example, broods produced by high-fitness females should contain fewer sons. We tested for offspring sex-ratio biasing consistent with these predictions in broad-horned flour beetles. We found that in both wild-type beetles and populations subject to artificial selection for high- and low-fitness males, offspring sex ratios were biased in the predicted direction: low-fitness females produced an excess of sons, whereas high-fitness females produced an excess of daughters. Thus, these beetles are able to adaptively bias sex ratio and recoup indirect fitness benefits of mate choice.     

Sex-limited expression of ornamental feathers in birds

Extravagant secondary sexual characters show sexual size dimorphismin some species but are completely sex limited in others. Sexualornamentation has been hypothesized to benefit mainly malesthrough sexual selection, but the costs of secondary sexualcharacters initially would be experienced by both sexes. Theevolution of sexual size dimorphism of ornaments and, eventually,the complete sex-limited expression of these characters, willdepend on the effects of sexual and natural selection on thetwo sexes. A phylogenetic analysis controlling for similaritiesdue to common ancestry of 60 independent evolutionary originsof feather ornamentation in birds was used to investigate ecologicalfactors correlated with sexual size dimorphism and sex-limitedexpression of secondary sexual characters. When the size ofan ornament is large relative to body size, the trait willbe particularly costly for females, resulting in selectionfor increased sexual size dimorphism of the ornament. Indeed,sexual size dimorphism of ornaments was positively relatedto the relative size of male ornaments but was unrelated torelative size of female ornaments. Species with polygynousand lekking mating systems with little or no male parentalcare (in particular nest building and incubation) demonstratedsex-limited expression of ornaments as compared to monogamousspecies. Species with no food provisioning of offspring by themale showed a trend for increased sexual size dimorphism ofornaments. Therefore, large natural selection costs duringreproduction imposed by the expression of secondary sexualcharacters are related to the evolution of sexual size dimorphismof ornaments and eventually their complete loss from females.     

The kin structure of sexual interactions

The origin of sexual reproduction involved the evolution of zygotes from separate genomes and, like other social processes, should therefore be amenable to analysis using kin selection theory. I consider how kin structure affects sexual interactions in three contexts—the evolution of sexual reproduction, sex allocation and sexual conflict. Kin structure helps explain the even-handed replication of paternal and maternal genes under outbreeding. Under inbreeding, it predicts altruistic failure to replicate by one half of the diploid genome. Kin structure predicts optimal sex ratios and potential conflicts over sex ratio within social groups and individuals. Sexual conflict predictably occurs as a function of (i) the probability that current sexual partners will reproduce together in future and (ii) between-partner relatedness. I conclude that systematically analysing the kin structure of sexual interactions helps illuminate their evolution.     

The effect of costly information in diet choice

Summary We distinguish three cases which consider the effect of information on animal behaviour: static information, obligate information and facultative information. Static information deals with the case in which the animal does not acquire additional information; it starts with enough information to discriminate options. Obligate information deals with the case in which the animal acquires information at no additional cost. Facultative information is when the animal may choose to pay a cost in order to acquire information. We illustrate the differences among these three situations by analysing the optimal diet problem subject to the different information regimes. Compared to the case with static information, obligate recognition time narrows the range of prey densities over which an optimal forager feeds selectively, and facultative recognition time reduces it further still. The three models yield qualitatively different predictions regarding how the optimal diet varies with relative abundances of alternative resources. In the space of resource densities, the line separating the optimal behaviours of selectivity and opportunism is straight for both the perfect and obligate information cases. In the case of facultative recognition time this line or isoleg is part of a quadratic curve. This non-linearity yields two completely new predictions: a less profitable resource may be lost from the diet after becoming more abundant and the poor resource may be included in the diet as a result of the rich resource becoming more common.     

Sex-biased survival predicts adult sex ratio variation in wild birds

Adult sex ratio (ASR) is a central concept in population demography and breeding system evolution, and has implications for population viability and biodiversity conservation. ASR exhibits immense interspecific variation in wild populations, although the causes of this variation have remained elusive. Using phylogenetic analyses of 187 avian species from 59 families, we show that neither hatching sex ratios nor fledging sex ratios correlate with ASR. However, sex-biased adult mortality is a significant predictor of ASR, and this relationship is robust to 100 alternative phylogenetic hypotheses, and potential ecological and life-history confounds. A significant component of adult mortality bias is sexual selection acting on males, whereas increased reproductive output predicts higher mortality in females. These results provide the most comprehensive insights into ASR variation to date, and suggest that ASR is an outcome of selective processes operating differentially on adult males and females. Therefore, revealing the causes of ASR variation in wild populations is essential for understanding breeding systems and population dynamics.