Adaptive sex-specific life history plasticity to temperature and photoperiod in a damselfly

We investigated four predictions about how temperature, photoperiod and sex affect the life history plasticity and foraging activity of a damselfly. (i) As predicted, increased temperatures increased foraging activity and growth rates, but in contrast with the prediction, late photoperiod (high time stress) did not affect foraging activity and growth rate. (ii) Unexpectedly, the increase in growth rate at increasing temperatures was not larger under high time stress. (iii) As predicted, age and size at emergence decreased at higher temperatures and at the late photoperiod. Temperature-induced life history shifts were direct or the result of behavioural growth mediation depending on the temperature range. Photoperiod-induced life history shifts were direct. (iv) As predicted, males emerged before females but at a smaller size. The degree of sexual size dimorphism was influenced by the joint effects of temperature and photoperiod. We could only detect genetic variation in size plasticity to photoperiod. The match between the sex-specific life history responses to temperature and photoperiod and predictions by relevant optimality models suggests adaptive life history plasticity to these variables.     

The effects of perceived predation risk on pre- and post-metamorphic phenotypes in the common frog

Where organisms undergo radical changes in habitat during ontogeny, dramatic phenotypic reshaping may be required. However, physiological and functional interrelationships may constrain the extent to which an individual's phenotype can be equally well adapted to their habitat throughout the life cycle. The phenotypic response of tadpoles to the presence of a predator has been reported for several species of anuran but the potential post-metamorphic consequences have rarely been considered. We reared common frog Rana temporaria tadpoles in the presence or absence of a larval odonate predator, Aeshna juncea , and examined the consequences of the resulting phenotypic adjustment in the aquatic larval stage of the life cycle for the terrestrial juvenile phenotype. In early development tadpoles developed deeper tail fins and muscles in response to the predator and, in experimental trials, swam further than those reared in the absence of a predator. While the difference in swimming ability remained significant throughout the larval period, by the onset of metamorphosis we could no longer detect any differences in the morphological parameters measured. The corresponding post-metamorphic phenotypes also did not initially differ in terms of morphology. At 12 weeks post-metamorphosis, however, froglets that developed from predator-exposed tadpoles swam more slowly and less far than those that developed from tadpoles reared in the absence of predators, the opposite trend to that observed in the larval stage of the life cycle, and had narrower femurs. These results suggest that there may be long-term costs for subsequent life-history stages of tailoring the larval phenotype to prevailing environmental conditions.     

Compensatory growth and oxidative stress in a damselfly

Physiological costs of compensatory growth are poorly understood, yet may be the key components in explaining why growth rates are typically submaximal. Here we tested the hypothesized direct costs of compensatory growth in terms of oxidative stress. We assessed oxidative stress in a study where we generated compensatory growth in body mass by exposing larvae of the damselfly Lestes viridis to a transient starvation period followed by ad libitum food. Compensatory growth in the larval stage was associated with higher oxidative stress (as measured by induction of superoxide dismutase and catalase) in the adult stage. Our results challenge two traditional views of life-history theory. First, they indicate that age and mass at metamorphosis not necessarily completely translate larval stress into adult fitness and that the observed physiological cost may explain hidden carry-over effects. Second, they support the notion that costs of compensatory growth may be associated with free-radical-mediated trade-offs and not necessarily with resource-mediated trade-offs.     

Evolution of clutch size in insects. I. A review of static optimality models

Abstract This paper reviews the importance of constraint assumptions to the predictions of static optimality models of insect clutch size. This allows us to identify predictions that distinguish between models embodying different constraints on female oviposition behaviour and hence to determine which resources or other factors limit clutch size evolutionarily. We conclude that while some models may be distinguished using qualitative criteria, others require the testing of quantitative predictions. In a companion paper (Wilson 1994) these models are tested using the bruchid beetle Callosobruchus maculatus.     

Latitudinal and voltinism compensation shape thermal reaction norms for growth rate

Latitudinal variation in thermal reaction norms of key fitness traits may inform about the response of populations to climate warming, yet their adaptive nature and evolutionary potential are poorly known. We assessed the contribution of quantitative genetic, neutral genetic and environmental effects to thermal reaction norms of growth rate for populations of the damselfly Ischnura elegans. Among populations, reaction norms differed primarily in elevation, suggesting that time constraints associated with shorter growth seasons in univoltine, high-latitude as well as multivoltine, low-latitude populations selected for faster growth rates. Phenotypic divergence among populations is consistent with selection rather than drift as Q(ST) was greater than F(ST) in all cases. Q(ST) estimates increased with experimental temperature and were influenced by genotype by environment interactions. Substantial additive genetic variation for growth rate in all populations suggests that evolution of trait means in different environments is not constrained. Heritability of growth rates was higher at high temperature, driven by increased genetic rather than environmental variance. While environment-specific nonadditive effects also may contribute to heritability differences among temperatures, maternal effects did not play a significant role (where these could be accounted for). Genotype by environment interactions strongly influenced the adaptive potential of populations, and our results suggest the potential for microevolution of thermal reaction norms in each of the studied populations. In summary, the observed latitudinal pattern in growth rates is adaptive and results from a combination of latitudinal and voltinism compensation. Combined with the evolutionary potential of thermal reaction norms, this may affect populations' ability to respond to future climate warming.     

Evolution of clutch size in insects. II. A test of static optimality models using the beetle Callosobruchus maculatus (Coleoptera: Bruchidae)

Wilson and Lessells (1993) analysed the effect of constraint assumptions on the predictions of static optimality models for insect clutch size. They concluded that the models could be reliably distinguished between (and hence the main constraints identified) only after precise quantitative predictions had been examined. The present paper describes a series of laboratory experiments, using the bruchid beetle Callosobruchus maculatus, that allow these quantitative predictions to be made and tested. Experiments in which female encounter rate with hosts was altered gave qualitative support for 3 out of 6 basic (single oviposition) models, but the quantitative fit of them all was poor. However, when the (a priori) condition was included in these models that several other females would oviposit on the same hosts (the multiple oviposition models), the time limiting multiple oviposition model alone produced quantitative predictions that were supported by observations. In other words, the results suggest that the main constraints on bruchid oviposition behaviour are the amount of time available for laying eggs and the number of other females ovipositing. However, additional qualitative predictions indicate that the number of eggs available to the female may also constrain clutch size evolutionarily. The usefulness of static optimality models for examining clutch size decisions in insects is discussed in the context of these results.     

Neuroendocrine control of life histories: what do we need to know to understand the evolution of phenotypic plasticity?

Almost all life histories are phenotypically plastic: that is, life-history traits such as timing of breeding, family size or the investment in individual offspring vary with some aspect of the environment, such as temperature or food availability. One approach to understanding this phenotypic plasticity from an evolutionary point of view is to extend the optimality approach to the range of environments experienced by the organism. This approach attempts to understand the value of particular traits in terms of the selection pressures that act on them either directly or owing to trade-offs due to resource allocation and other factors such as predation risk. Because these selection pressures will between environments, the predicted optimal phenotype will too. The relationship expressing the optimal phenotype for different environments is the optimal reaction norm and describes the optimal phenotypic plasticity. However, this view of phenotypic plasticity ignores the fact that the reaction norm must be underlain by some sort of control system: cues about the environment must be collected by sense organs, integrated into a decision about the appropriate life history, and a message sent to the relevant organs to implement that decision. In multicellular animals, this control mechanism is the neuroendocrine system. The central question that this paper addresses is whether the control system affects the reaction norm that evolves. This might happen in two different ways: first, the control system will create constraints on the evolution of reaction norms if it cannot be configured to produce the optimal reaction norm and second, the control system will create additional selection pressures on reaction norms if the neuroendocrine system is costly. If either of these happens, a full understanding of the way in which selection shapes reaction norms must include details of the neuroendocrine control system. This paper presents the conceptual framework needed to explain what is meant by a constraint or cost being created by the neuroendocrine system and discusses the extent to which this occurs and some possible examples. The purpose of doing this is to encourage endocrinologists to take a fresh look at neuroendocrine mechanisms and help identify the properties of the system and situations in which these generate constraints and costs that impinge on the evolution of phenotypic plasticity.     

Life history perspectives on pest insects: What’s the use?

In his seminal 1954 paper on the ‘population consequences of life history phenomena’, Cole noted that ‘these computations may have practical value in dealing with valuable or noxious species’. In the present paper, the question is asked: ‘is research based on evolutionary perspectives in general, and life history theory specifically, really useful for dealing with insect pests?’ Perhaps such theory‐based research is rather a luxury: time and resources would be better spent on entirely applied aspects of the problem. The conclusion of the present discussion is that having an evolutionary perspective guiding research is actually a very cost‐effective way of dealing with applied problems, as it provides a clear basis for interpretations, generalizations and predictions. Life history theory is a very central and necessary part of both population ecology and general evolutionary theory, and its specific usefulness in pest forecasting and management are discussed. Nevertheless, our ability to predict insect population dynamics is still limited, and so is our ability to make use of an insect’s life history traits to predict its propensity to become a pest. I suggest that the former shortcoming is largely due to poor understanding of insect life history plasticity. This, in turn, may partly be due to a paucity of studies where reaction norms are investigated as putative adaptations. I suggest that the latter shortcoming is due to problems inherent with studying life history traits as adaptations, for example the lack of an independent fitness model and the fact that life histories tend to form syndromes of coadapted traits. These points are illustrated with examples from my own work on non‐pest butterflies and from insect–Eucalyptus systems.     

Differential elevational cline in the phenology and demography of two temporally isolated populations of a damselfly: Not two but one taxon?

1. Temporal isolation by cohort splitting is a life‐history mechanism that has been reported in many temperate insects, including those inhabiting freshwater habitats. Although the cohorts seem to maintain separate temporal niches in a specific location, the temporal isolation may be disrupted across a geographic gradient due to constraints imposed by seasonality. 2. This prediction was tested on two temporally isolated populations of the obligatory univoltine Lestes virens (Odonata, Lestidae) in north‐east Algeria. Although the two cohorts emerge at the same time in spring, one cohort reproduces in summer, while the second cohort estivates in summer and reproduces in autumn. A survey assessing the phenology and abundance was conducted on eight ponds across an elevational gradient (5–1012 m asl) using capture–mark–recapture and adult density sampling. 3. In all sites from low to high elevation, the species showed cohort splitting. The phenology of reproduction of both cohorts showed a delay with elevation, but the cline was 2.2 days for the summer cohort and 0.7 days for the autumn cohort per 100 m of elevation. Moreover, the density of adults in the autumn cohort was higher than that of summer cohort across the entire elevational range, and the difference increased with elevation. 4. These findings regarding the differential elevational cline in the phenology show that the temporal isolation of the two cohorts becomes narrower at high elevation, suggesting potential inter‐cohort temporal overlap at higher elevations. 5. The claim that the two cohorts of L. virens are true temporally isolated species needs further investigation.     

The evolution of predictive adaptive responses in human life history

Many studies in humans have shown that adverse experience in early life is associated with accelerated reproductive timing, and there is comparative evidence for similar effects in other animals. There are two different classes of adaptive explanation for associations between early-life adversity and accelerated reproduction, both based on the idea of predictive adaptive responses (PARs). According to external PAR hypotheses, early-life adversity provides a ‘weather forecast’ of the environmental conditions into which the individual will mature, and it is adaptive for the individual to develop an appropriate phenotype for this anticipated environment. In internal PAR hypotheses, early-life adversity has a lasting negative impact on the individual''s somatic state, such that her health is likely to fail more rapidly as she gets older, and there is an advantage to adjusting her reproductive schedule accordingly. We use a model of fluctuating environments to derive evolveability conditions for acceleration of reproductive timing in response to early-life adversity in a long-lived organism. For acceleration to evolve via the external PAR process, early-life cues must have a high degree of validity and the level of annual autocorrelation in the individual''s environment must be almost perfect. For acceleration to evolve via the internal PAR process requires that early-life experience must determine a significant fraction of the variance in survival prospects in adulthood. The two processes are not mutually exclusive, and mechanisms for calibrating reproductive timing on the basis of early experience could evolve through a combination of the predictive value of early-life adversity for the later environment and its negative impact on somatic state.     

动物生活史进化理论研究进展

综述了生活史性状、生活史对策、权衡、适合度及进化种群统计学等动物生活史进化领域的进展。权衡是生活史性状之间相互联系的纽带,分为生理权衡与进化权衡。适合度是相对的,与个体所处的特定环境条件有关,性状进化与适合度之间关系紧密。适合度是生活史进化理论研究的焦点。探讨动物生活史对策的理论很多,影响最大的是MacArthur和Wilson提出的r对策及K对策理论。随年龄的增长,动物存活率及繁殖率逐步下降的过程,称为衰老;解释衰老的进化理论主要有突变-选择平衡假设和多效对抗假设。进化种群统计学将种群统计学应用于生活史进化研究,为探讨表型适合度的进化提供了有效的手段。将进化种群统计学、数量遗传学及特定种系效应理论进行整合,建立完整的动物生活史进化综合理论体系,是当代此领域的最大挑战。