David M. Shuker

SEX-RATIO ADJUSTMENT WHEN RELATIVES INTERACT: A TEST OF CONSTRAINTS ON ADAPTATION

Abstract Studies of sex allocation offer excellent opportunities for examining the constraints and limits on adaptation. A major topic of debate within this field concerns the extent to which the ability of individuals to adaptively manipulate their offspring sex ratio is determined by constraints such as the method of sex determination. We address this problem by comparing the extent of sex‐ratio adjustment across taxa with different methods of sex determination, under the common selective scenario of interactions between relatives. These interactions comprise the following: local resource competition (LRC), local mate competition (LMC), and local resource enhancement (LRE). We found that: (1) species with supposedly constraining methods of sex determination showed consistent sex‐ratio adjustment in the predicted direction; (2) vertebrates with chromosomal sex determination (CSD) showed less adjustment then haplodiploid invertebrates; (3) invertebrates with possibly constraining sex‐determination mechanisms (CSD and pseudo‐arrhenotoky) did not show less adjustment then haplodiploid invertebrates; (4) greater sex‐ratio adjustment was seen in response to LRC and LMC than LRE; (5) greater sex‐ratio adjustment was seen in response to interactions between relatives (LRC, LMC, and LRE) compared to responses to other environmental factors. Our results also illustrate the problem that sex‐determination mechanism and selective pressure are confounded across taxa because vertebrates with CSD are influenced primarily by LRE whereas invertebrates are influenced by LRC and LMC. Overall, our analyses suggest that sex‐allocation theory needs to consider simultaneously the influence of variable selection pressures and variable constraints when applying general theory to specific cases.

Cooperation and the Scale of Competition in Humans

Explaining cooperation is one of the greatest challenges for evolutionary biology. It is particularly a problem in species such as humans, where there is cooperation between nonrelatives. Numerous possible solutions have been suggested for the problem of cooperation between nonrelatives, including punishment, policing, and various forms of reciprocity. Here, we suggest that local competition for resources can pose a problem for these hypotheses, analogous to how it can select against cooperation between relatives. We extend the prisoners dilemma (PD) game to show that local competition between interacting individuals can reduce selection for cooperation between nonrelatives. This is because, with local competition, fitness is relative to social partners, and cooperation benefits social partners. We then test whether nonrelated humans adjust their level of cooperation facultatively in response to the scale of competition when playing the PD for cash prizes. As predicted, we found that individuals were less likely to cooperate when competition was relatively local. Cooperation between humans will therefore be most likely when repeated interactions take place on a local scale between small numbers of people, and competition for resources takes place on a more global scale among large numbers of people.

The Coevolution of Virulence: Tolerance in Perspective

Coevolutionary interactions, such as those between host and parasite, predator and prey, or plant and pollinator, evolve subject to the genes of both interactors. It is clear, for example, that the evolution of pollination strategies can only be understood with knowledge of both the pollinator and the pollinated. Studies of the evolution of virulence, the reduction in host fitness due to infection, have nonetheless tended to focus on parasite evolution. Host-centric approaches have also been proposed—for example, under the rubric of “tolerance”, the ability of hosts to minimize virulence without necessarily minimizing parasite density. Within the tolerance framework, however, there is room for more comprehensive measures of host fitness traits, and for fuller consideration of the consequences of coevolution. For example, the evolution of tolerance can result in changed selection on parasite populations, which should provoke parasite evolution despite the fact that tolerance is not directly antagonistic to parasite fitness. As a result, consideration of the potential for parasite counter-adaptation to host tolerance—whether evolved or medially manipulated—is essential to the emergence of a cohesive theory of biotic partnerships and robust disease control strategies.

Sex Ratios under Asymmetrical Local Mate Competition: Theory and a Test with Parasitoid Wasps

Sex ratio theory allows unparalleled opportunities for testing how well animal behavior can be predicted by evolutionary theory. For example, Hamilton’s theory of local mate competition (LMC) is well understood and can explain variation in sex allocation across numerous species. This allows more specific predictions to be developed and tested. Here we extend LMC theory to a situation that will be common in a range of species: asymmetrical LMC. Asymmetrical LMC occurs when females lay eggs on a patch asynchronously and male offspring do not disperse, leading to relatively weaker LMC for males emerging from later broods. Varying levels of LMC then lead to varying optimal sex ratios for females, depending on when and where they oviposit. We confirm the assumptions of our theory using the wasp Nasonia vitripennis and then test our predictions. We show that females adjust their offspring sex ratios in the directions predicted, laying different sex ratios on different hosts within a patch. Specifically, there was a less female‐biased sex ratio when ovipositing on an unparasitized host if another host on the patch had previously been parasitized and a less female‐biased sex ratio on parasitized hosts if females also oviposited on an unparasitized host.

THE NICHE CONSTRUCTION PERSPECTIVE: A CRITICAL APPRAISAL

Niche construction refers to the activities of organisms that bring about changes in their environments, many of which are evolutionarily and ecologically consequential. Advocates of niche construction theory (NCT) believe that standard evolutionary theory fails to recognize the full importance of niche construction, and consequently propose a novel view of evolution, in which niche construction and its legacy over time (ecological inheritance) are described as evolutionary processes, equivalent in importance to natural selection. Here, we subject NCT to critical evaluation, in the form of a collaboration between one prominent advocate of NCT, and a team of skeptics. We discuss whether niche construction is an evolutionary process, whether NCT obscures or clarifies how natural selection leads to organismal adaptation, and whether niche construction and natural selection are of equivalent explanatory importance. We also consider whether the literature that promotes NCT overstates the significance of niche construction, whether it is internally coherent, and whether it accurately portrays standard evolutionary theory. Our disagreements reflect a wider dispute within evolutionary theory over whether the neo‐Darwinian synthesis is in need of reformulation, as well as different usages of some key terms (e.g., evolutionary process).

Genetic structure of natural Nasonia vitripennis populations: validating assumptions of sex-ratio theory

The parasitic wasp Nasonia vitripennis has been used extensively in sex allocation research. Although laboratory experiments have largely confirmed predictions of local mate competition (LMC) theory, the underlying assumptions of LMC models have hardly been explored in nature. We genotyped over 3500 individuals from two distant locations (in the Netherlands and Germany) at four polymorphic microsatellite loci to validate key assumptions of LMC theory, in terms of both the original models and more recent extensions to them. We estimated the number of females contributing eggs to patches of hosts and the clutch sizes as well as sex ratios produced by individual foundresses. In addition, we evaluated the level of inbreeding and population differentiation. Foundress numbers ranged from 1 to 7 (average 3.0 ± 0.46 SE). Foundresses were randomly distributed across the patches and across hosts within patches, with few parasitizing more than one patch. Of the hosts, 40% were parasitized by more than one foundress. Clutch sizes of individual foundresses (average 9.99 ± 0.51 SE) varied considerably between hosts. The time period during which offspring continued to emerge from a patch or host correlated strongly with foundress number, indicating that sequential rather than simultaneous parasitism is the more common. Genetic differentiation at the regional level between Germany and the Netherlands, as estimated by Slatkins private allele method (0.11) and Hedricks corrected G′LT (0.23), indicates significant substructuring between regions. The level of population inbreeding for the two localities (FIL = 0.168) fitted the expectation based on the average foundress number per patch.

Kin discrimination and sex ratios in a parasitoid wasp

Sex ratio theory provides a clear and simple way to test if nonsocial haplodiploid wasps can discriminate between kin and nonkin. Specifically, if females can discriminate siblings from nonrelatives, then they are expected to produce a higher proportion of daughters if they mate with a sibling. This prediction arises because in haplodiploids, inbreeding (sib‐mating) causes a mother to be relatively more related to her daughters than her sons. Here we formally model this prediction for when multiple females lay eggs in a patch, and test it with the parasitoid wasp Nasonia vitripennis. Our results show that females do not adjust their sex ratio behaviour dependent upon whether they mate with a sibling or nonrelative, in response to either direct genetic or a range of indirect environmental cues. This suggests that females of N. vitripennis cannot discriminate between kin and nonkin. The implications of our results for the understanding of sex ratio and social evolution are discussed.

Relationships between student characteristics and self‐, peer and tutor evaluations of oral presentations

There are many influences on how assessors grade themselves and others. Oral presentations are useful for exploring such factors in peer, self‐ and tutor marked assessments, being rapidly completed and assessed, commonly used in HE and very difficult to anonymize. This opportunistic study examined the effects of gender and level of attainment on the triangulation of marks awarded to student presenters. Grades generated by peer assessment were associated more strongly with tutor‐awarded marks than those from self‐assessment. For self‐assessment there was a strong effect of gender (female students undervalued their performance compared with tutor grades). Peer assessment produced higher marks than from tutors, perhaps because of the close‐knit community developed during residential courses. For tutor marks, the greatest variability was at the lower end of the scale, whereas peer assessors were most variable when marking students who self‐evaluated or peer assessed highly. Students awarded a narrower range of marks to peers compared with tutors, but when self‐assessing used a larger range. Presentations by students who admitted to little sleep the night before received lower grades from both peers and tutors, but this was not reflected by self‐assessments, suggesting they were unaware of their poorer performances. Sessions with fewer talks (four rather than seven) reduced the ‘dip’ in marks previously observed in the middle of sessions. Findings are discussed in the context of bias in this mode of assessment.

Facultative Sex Ratio Adjustment in Natural Populations of Wasps: Cues of Local Mate Competition and the Precision of Adaptation

Sex ratio theory offers excellent opportunities to examine the extent to which individuals adaptively adjust their behavior in response to local conditions. Hamilton’s theory of local mate competition, which predicts female‐biased sex ratios in structured populations, has been extended in numerous directions to predict individual behavior in response to factors such as relative fecundity, time of oviposition, and relatedness between cofoundresses and between mates. These extended models assume that foundresses use different sources of information, and they have generally been untested or have only been tested in the laboratory. We use microsatellite markers to describe the wild oviposition behavior of individual foundresses in natural populations of the parasitoid wasp Nasonia vitripennis, and we use the data collected to test these various models. The offspring sex ratio produced by a foundress on a particular host reflected the number of eggs that were laid on that host relative to the number of eggs that were laid on that host by other foundresses. In contrast, the offspring sex ratio was not directly influenced by other potentially important factors, such as the number of foundresses laying eggs on that patch, relative fecundity at the patch level, or relatedness to either a mate or other foundresses on the patch.

Patterns of male sterility in a grasshopper hybrid zone imply accumulation of hybrid incompatibilities without selection

It is now widely accepted that post-zygotic reproductive isolation is the result of negative epistatic interactions between derived alleles fixed independently at different loci in diverging populations (the Dobzhansky–Muller model). What is less clear is the nature of the loci involved and whether the derived alleles increase in frequency through genetic drift, or as a result of natural or sexual selection. If incompatible alleles are fixed by selection, transient polymorphisms will be rare and clines for these alleles will be steep where divergent populations meet. If they evolve by drift, populations are expected to harbour substantial genetic variation in compatibility and alleles will introgress across hybrid zones once they recombine onto a genetic background with which they are compatible. Here we show that variation in male sterility in a naturally occurring Chorthippus parallelus grasshopper hybrid zone conforms to the neutral expectations. Asymmetrical clines for male sterility have long tails of introgression and populations distant from the zone centre show significant genetic variation for compatibility. Our data contrast with recent observations on ‘speciation genes’ that have diverged as a result of strong natural selection.