Recent interest has focused on immune response in an evolutionary context, with particular attention to disease resistance as a life-history trait, subject to trade-offs against other traits such as reproductive effort. Immune defense has several characteristics that complicate this approach, however; for example, because of the risk of autoimmunity, optimal immune defense is not necessarily maximum immune defense. Two important types of cost associated with immunity in the context of life history are resource costs, those related to the allocation of essential but limited resources, such as energy or nutrients, and option costs, those paid not in the currency of resources but in functional or structural components of the organism. Resource and option costs are likely to apply to different aspects of resistance. Recent investigations into possible trade-offs between reproductive effort, particularly sexual displays, and immunity have suggested interesting functional links between the two. Although all organisms balance the costs of immune defense against the requirements of reproduction, this balance works out differently for males than it does for females, creating sex differences in immune response that in turn are related to ecological factors such as the mating system. We conclude that immune response is indeed costly and that future work would do well to include invertebrates, which have sometimes been neglected in studies of the ecology of immune defense.
Phenotypic plasticity can be broadly defined as the ability of one genotype to produce more than one phenotype when exposed to different environments, as the modification of developmental events by the environment, or as the ability of an individual organism to alter its phenotype in response to changes in environmental conditions. Not surprisingly, the study of phenotypic plasticity is innately interdisciplinary and encompasses aspects of behavior, development, ecology, evolution, genetics, genomics, and multiple physiological systems at various levels of biological organization. From an ecological and evolutionary perspective, phenotypic plasticity may be a powerful means of adaptation and dramatic examples of phenotypic plasticity include predator avoidance, insect wing polymorphisms, the timing of metamorphosis in amphibians, osmoregulation in fishes, and alternative reproductive tactics in male vertebrates. From a human health perspective, documented examples of plasticity most commonly include the results of exercise, training, and/or dieting on human morphology and physiology. Regardless of the discipline, phenotypic plasticity has increasingly become the target of a plethora of investigations with the methodological approaches utilized ranging from the molecular to whole organsimal. In this article, we provide a brief historical outlook on phenotypic plasticity; examine its potential adaptive significance; emphasize recent molecular approaches that provide novel insight into underlying mechanisms, and highlight examples in fishes and insects. Finally, we highlight examples of phenotypic plasticity from a human health perspective and underscore the use of mouse models as a powerful tool in understanding the genetic architecture of phenotypic plasticity.
Some eyespots are thought to deflect attack away from the vulnerable body, yet there is limited empirical evidence for this function and its adaptive advantage. Here, we demonstrate the conspicuous ventral hindwing eyespots found on Bicyclus anynana butterflies protect against invertebrate predators, specifically praying mantids. Wet season (WS) butterflies with larger, brighter eyespots were easier for mantids to detect, but more difficult to capture compared to dry season (DS) butterflies with small, dull eyespots. Mantids attacked the wing eyespots of WS butterflies more frequently resulting in greater butterfly survival and reproductive success. With a reciprocal eyespot transplant, we demonstrated the fitness benefits of eyespots were independent of butterfly behaviour. Regardless of whether the butterfly was WS or DS, large marginal eyespots pasted on the hindwings increased butterfly survival and successful oviposition during predation encounters. In previous studies, DS B. anynana experienced delayed detection by vertebrate predators, but both forms suffered low survival once detected. Our results suggest predator abundance, identity and phenology may all be important selective forces for B. anynana. Thus, reciprocal selection between invertebrate and vertebrate predators across seasons may contribute to the evolution of the B. anynana polyphenism.
In animals, sex differences in immunity are proposed to shape variation in infection prevalence and intensity among individuals in a population, with females typically expected to exhibit superior immunity due to life‐history trade‐offs. We performed a systematic meta‐analysis to investigate the magnitude and direction of sex differences in immunity and to identify factors that shape sex‐biased immunocompetence. In addition to considering taxonomic and methodological effects as moderators, we assessed age‐related effects, which are predicted to occur if sex differences in immunity are due to sex‐specific resource allocation trade‐offs with reproduction. In a meta‐analysis of 584 effects from 124 studies, we found that females exhibit a significantly stronger immune response than do males, but the effect size is relatively small, and became non‐significant after controlling for phylogeny. Female‐biased immunity was more pronounced in adult than immature animals. More recently published studies did not report significantly smaller effect sizes. Among taxonomic and methodological subsets of the data, some of the largest effect sizes were in insects, further supporting previous suggestions that testosterone is not the only potential driver of sex differences in immunity. Our findings challenge the notion of pervasive biases towards female‐biased immunity and the role of testosterone in driving these differences.
Mycoplasma gallisepticum is a well-known disease of poultry, but until 1994 it had not been observed in passerine birds. From 1994 to 1996, tens of millions of house ¢nches (Carpodacus mexicanus) are believed to have died in an epidemic of mycoplasmal conjunctivitis, similar to`pinkeye' in humans. The outbreak of Mycoplasma gallisepticum a¡ected ¢nches of both sexes, but disproportionately killed males, shifting the sex ratio from male-biased to female-biased. This di¡erential male mortality is consistent with a cost of testosterone, which is a key prediction of the immunocompetence handicap hypothesis. Males and females that survived the epidemic weighed signi¢cantly less and had signi¢cantly shorter wing chords, tarsi, and bills than did individuals before the epidemic. Male survivors also had signi¢cantly redder plumage than males that did not survive, supporting the idea that plumage brightness serves as an indicator of condition, as proposed by the honest advertisement model of sexual selection.
Summary 1.Experimental work in vertebrates shows that different forms of coloration -particularly carotenoid-based vs melanin-based ornaments -are differentially sensitive to stressors such as diet manipulations and parasitic infections. Comparative work also shows that historical patterns of ornament evolution depend on the proximate mechanisms of colour production. 2. Based on the results from vertebrates, it has been argued that melanin-based ornaments may not be good indicators of quality because, unlike carotenoid-based ornaments, melanin pigments can be synthesized from amino acid precursors, and therefore melanin synthesis is unlikely to be costly. 3. However, there is evidence for costly melanin synthesis in insects, which use melanin in some ways that vertebrates do not. Differences across taxa in the costs of melanin synthesis challenge the generality of the hypotheses relevant to these ornaments. 4. In general, all pigment-based ornaments are a function of precursor intake and processing. Differences in costs of melanin synthesis across taxa may reflect differences in one or both of these parameters. 5. I urge communication between researchers working on related questions in disparate taxa (vertebrates and invertebrates). This should facilitate the discovery of the most fundamental aspects of colour ornament evolution.
Immune defense often differs between the sexes, with males often having a weaker response, at least among many vertebrates. We examined encapsulation ability, a cell-mediated immune response, in laboratory and field populations of two species of field crickets, Teleogryllus oceanicus (Le Guillou, 1841) and Teleogryllus commodus (Walker, 1869), which have different life histories. In the seasonally breeding T. commodus, males show a stronger encapsulation response than females in both the laboratory and the field, although the difference is more marked under field conditions. The aseasonal T. oceanicus showed no sex difference in encapsulation in either field or laboratory samples fed ad libitum, but when food was experimentally reduced, the same pattern of stronger male response emerged. It is possible that this pattern may result from selection on females to increase investment in reproduction when time and energy for breeding are limited, as is more likely for seasonal breeders or animals under food restriction.
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