The evolution of genome size: what can be learned from anuran development?

Differences in nuclear DNA content in vertebrates have been shown to be correlated with cell size, cell division rate, and embryonic developmental rate. We compare seven species of anuran amphibians with a three-fold range of genome sizes. Parameters examined include the number and density of cells in a number of embryonic structures, and the change in cell number in the CNS during development. We show that genome size is correlated with cell proliferation rate and with developmental rate at different stages of embryonic development, but that the correlation between genome size and cell size is only evident at later stages. We discuss the evolution of genome size in amphibians. Our discussion takes into account data that reportedly support two conflicting hypotheses: the "skeletal DNA" hypothesis, which claims a selective role for differences in genome size, and the "junk DNA" hypothesis, which claims that differences in genome size are a random result of the accumulation of noncoding DNA sequences. We show that these supposedly conflicting hypotheses can be integrated into a more complex and inclusive model for the evolution of genome size.     

The contribution of small individuals to density–body size relationships

The relationship between density and body size is central to our understanding of species assemblages. The greatest challenge in sampling complete assemblages is obtaining reliable estimates of all taxa regardless of body size. We therefore examined the density-body size relationship in a coral reef fish assemblage using a novel sampling method which permits reliable quantification of the small/cryptic reef fish fauna. We found a negative linear relationship between density and adult body size. This is in marked contrast to the polygonal relationship previously described for other local scale assemblage studies. Our linear relationship may be a consequence of the larger differences in body size among taxa. Spanning over five orders of magnitude, the range of body sizes appears to be an important factor in shaping density-body size relationships.     

The evolution of growth trajectories: what limits growth rate?

According to life‐history theory, growth rates are subject to strong directional selection due to reproductive and survival advantages associated with large adult body size. Yet, growth is commonly observed to occur at rates lower than the maximum that is physiologically possible and intrinsic growth rates often vary among populations. This implies that slower growth is favoured under certain conditions. Realized growth rate is thus the result of a compromise between the costs and advantages of growing rapidly, and the optimal rate of growth is not equivalent to the fundamental maximum rate. The ecological and evolutionary factors influencing growth rate are reviewed, with particular emphasis on how growth might be constrained by direct fitness costs. Costs of accelerating growth might contribute to the variance in fitness that is not attributable to age or size at maturity, as well as to the variation in life‐history strategies observed within and among species. Two main approaches have been taken to study the fitness trade‐offs relating to growth rate. First, environmental manipulations can be used to produce treatment groups with different rates of growth. Second, common garden experiments can be used to compare fitness correlates among populations with different intrinsic growth rates. Data from these studies reveal a number of potential costs for growth over both the short and long term. In order to acquire the energy needed for faster growth, animals must increase food intake. Accordingly, in many taxa, the major constraint on growth rate appears to arise from the trade‐off between predation risk and foraging effort. However, growth rates are also frequently observed to be submaximal in the absence of predation, suggesting that growth trajectories also impact fitness via other channels, such as the reallocation of finite resources between growth and other traits and functions. Despite the prevalence of submaximal growth, even when predators are absent, there is surprisingly little evidence to date demonstrating predator‐independent costs of growth acceleration. Evidence that does exist indicates that such costs may be most apparent under stressful conditions. Future studies should examine more closely the link between patterns of resource allocation to traits in the adult organism and lifetime fitness. Changes in body composition at maturation, for example, may determine the outcome of trade‐offs between reproduction and survival or between early and late reproduction. A number of design issues for studies investigating costs of growth that are imposed over the long term are discussed, along with suggestions for alternative approaches. Despite these issues, identifying costs of growth acceleration may fill a gap in our understanding of life‐history evolution: the relationships between growth rate, the environment, and fitness may contribute substantially to the diversification of life histories in nature.     

The evolution of large size: how does Cope's Rule work?

Cope's Rule is the tendency for organisms in evolving lineages to increase in size over time. The concept is detailed in many textbooks, but has rarely been demonstrated. Many suggestions of the benefits of large body size exist, but none has yet been confirmed empirically. Using a large-scale analysis of recent studies, Kingsolver and Pfennig have now shown how size benefits survival, mating success and fecundity, and they provide convincing arguments for a mechanism that is capable of driving Cope's Rule.     

How accurately can retention of benthic macrofauna by a particular mesh size be predicted from body size of organisms?

Size of organisms is frequently the prime criterion in selecting a mesh size to sample benthic assemblages. This study quantified the accuracy in estimating the sampling efficiency of screens from body size of macrozoobenthos in the upper, sandy, reaches of a small, shallow estuary, where the community consisted primarily of peracarid crustaceans and polychaetes. Body size of organisms retained by a 0.25, 0.5 and 1.0 mm screen was used to predict the retention efficiency of each gear by multiple discriminant analysis (MDA), or by simply assuming that an animal of known size will be retained by a screen of the same aperture size (‘body aperture match’). MDA-models classified about 70% of all cases correctly, whereas matching of body-to screen-size gave at best spurious, and often seriously wrong estimates of retention efficiency for all tested mesh sizes. Regardless of the method employed, body size was found to be an extremely poor predictor of mesh retention. Consequently, sampling performance of each mesh size in a particular habitat × community combination should be determined experimentally and not from body size measurements obtained during pilot studies.     

How did big brains evolve? The role of neonatal body size

The hypothesis that a mammalian species’ neonatal body size plays a role in determining its adult brain size is examined. This hypothesis is suggested by the observed correlation among mammals between neonatal body weight and adult brain weight. It is argued that there is no direct developmental linkage between these variables; rather, they are associated in evolution because of their opposing effects on the maturity level of the neonate. The evolution of neonate size in the hominids is also discussed, and consideration is given to trade-offs in pelvic design between locomotor and obstetrical functions. It is concluded that there was strong selection for brain enlargement in the hominids and that neonatal enlargement, rather than being intrinsically adaptive, was a direct response to the maturity-reducing effect of adult brain enlargement.     

A phylogenetic analysis of body size evolution in the Anolis roquet group (Sauria: Iguanidae): character displacement or size assortment?

The important role that competition plays in structuring communities is well documented; however, the role of competition in an evolutionary context remains unclear. Evolutionary investigations into the role of competition have often focused on the process of character displacement, and a good example of this is the evolution of body size in the Anolis lizards of the Caribbean islands. Previous work on the A. roquet species group has taken a phylogenetic approach and concluded that patterns of body size differences are not caused by character displacement but are a result of size assortment. Using a phylogenetic reconstruction based on the sequence of the cytochrome b gene (cyt-b) and ancestral character-state reconstruction methods, we investigated the roles of character displacement and size assortment. Our results indicated that size assortment alone was insufficient to explain the observed patterns of body size differences. Furthermore, we found that change in body size was associated with a change in allopatry/sympatry, thus supporting the character-displacement hypothesis. We conclude that patterns of body size differences in the A. roquet species group appear to be the result of a combination of character displacement and size assortment because character displacement was only found to be possible on three occasions.     

What keeps insects small? Time limitation during oviposition reduces the fecundity benefit of female size in a butterfly

Laboratory studies of insects suggest that female fecundity may increase very rapidly with adult size and that mass may often increase close to exponentially with time during larval growth. These relationships make it difficult to see how realistic levels of larval mortality can outweigh the fecundity benefit of prolonged growth. Hence, it is unclear why many insects do not become bigger. In this study, we experimentally explore the relationship between female size and fecundity in the butterfly Pararge aegeria and show that thermally induced time limitation during oviposition may substantially reduce the fecundity benefit of larger females. We model time-limited oviposition under natural temperature conditions and show that fecundity is also likely to increase asymptotically with female size in the field. With realistic estimates of juvenile mortality, the model predicts optimal body sizes within the observed range even when larvae grow exponentially. We conclude that one important reason for why insects with a high capacity of larval growth do not evolve toward larger sizes may be that the fecundity benefit is in fact relatively limited under natural conditions. If so, these results may help resolve some of the inconsistencies between theory and empirical patterns in explaining optimal size in insects.     

The evolution of ADHD: a disorder of communication?

Attention deficit hyperactivity disorder (ADHD) is the most commonly diagnosed psychiatric condition. Many believe that the central disability is impaired inhibition, which leads to reduced abilities in social skills, self-control, organization and time management. The behaviors identified by clinicians as problematic--inattention, hyperactivity and impulsivity--have been incorporated into several evolutionary models as selectively adaptive cognitive skills for surviving the challenges of a variable Pleistocene environment. We propose that the "disabilities" exhibited by individuals with ADHD are maladaptive, and we concur with Barkley that there is a central impairment in the behavioral inhibition system. The underlying neural anatomy and physiology support the possibility that neurotransmitter pathology may have an impact on other interlinked systems (including language), and may also account for the frequent comorbidity of aggression, anxiety, depression, and learning disabilities (many of which are language-related). Language skills compete with other cognitive activities for the attentional system, and thus the evolution of language could not in fact be independent of the evolution of attention. If language represents the ultimate expression of the attentional system, and some individuals with ADHD are seriously impaired in the coordination of interlinked neural systems (including language), then ADHD fits Jerome Wakefield's definition of "harmful dysfunction," and communication impairments should be investigated more thoroughly by clinicians.     

The ontology of organisms: Mechanistic modules or patterned processes?

Though the realm of biology has long been under the philosophical rule of the mechanistic magisterium, recent years have seen a surprisingly steady rise in the usurping prowess of process ontology. According to its proponents, theoretical advances in the contemporary science of evo-devo have afforded that ontology a particularly powerful claim to the throne: in that increasingly empirically confirmed discipline, emergently autonomous, higher-order entities are the reigning explanantia. If we are to accept the election of evo-devo as our best conceptualisation of the biological realm with metaphysical rigour, must we depose our mechanistic ontology for failing to properly “carve at the joints” of organisms? In this paper, I challenge the legitimacy of that claim: not only can the theoretical benefits offered by a process ontology be had without it, they cannot be sufficiently grounded without the metaphysical underpinning of the very mechanisms which processes purport to replace. The biological realm, I argue, remains one best understood as under the governance of mechanistic principles.     

ATP synthase: what dictates the size of a ring?

Recent data suggest the source of F(0)F(1) ATP synthase determines a significant and surprising difference in the size of a putative rotating ring of integral membrane subunits of F(0); this can be correlated with biochemical data suggesting there is variation in the number of protons translocated per ATP synthesised.     

Investigating yellow dung fly body size evolution in the field: Response to climate change?

Uncovering genetic responses to selection in wild populations typically requires tracking individuals over generations and use of animal models. Our group monitored the body size of one Swiss Yellow Dung Fly (Scathophaga stercoraria; Diptera: Scathophagidae) field population over 15 years, including intermittent common‐garden rearing in the laboratory to assess body size with minimized environmental and maximized genetic variation. Contrary to expectations based on repeated heritability and phenotypic selection assessments over the years (reported elsewhere), field body sizes declined by >10% and common‐garden laboratory sizes by >5% from 1993 to 2009. Our results confirm the temperature‐size rule (smaller when warmer) and, albeit entirely correlational, could be mediated by climate change, as over this period mean temperature at the site increased by 0.5°C, although alternative systematic environmental changes cannot be entirely excluded. Monitoring genetic responses to selection in wild invertebrate populations is thus possible, though indirect, and wild populations may evolve in directions not consistent with strongly positive directional selection favoring large body size.     

The evolution of flightlessness: Is history important?

Summary Though most birds and insects are capable of flight (volant) some species are flightless. In this paper I test the hypothesis that phylogenetic constraints have played a role in the evolution of flightlessness. If speciation occurred after the evolutionary transition to flightlessness, inferences concerning the importance of particular aspects of the environment on the probability of the evolution of flightlessness may be statistically spurious because of the inflation of the sample size. Among birds, ratites and penguins illustrate the phenomenon of considerable speciation subsequent to the transition to the evolution of flightlessness. In contrast, the rails represent a group in which each flightless species probably represents a separate evolutionary transition. There are many more flightless insect species than bird species and several orders are monomorphically flightless, the sometimes enormous speciation within the order following and possibly being a consequence of the evolution of flightlessness. While it can be shown in insects that flightlessness has evolved independently many times, there are at least as many cases in which the question cannot be resolved. Therefore, in both birds and insects phylogenetic effects should not be ignored, for the number of evolutionary transitions may be much less than the number of species. The effect of incorporating phylogenetic (or at least taxonomic) constraints into the analysis of habitat factors associated with flightlessness is considered.     

The evolution of regeneration: adaptive or inherent?

If regeneration were adaptive, it would have arisen autonomously by natural selection from non-regenerative antecedents. Unless each episode coincidentally reinvented the same method of regeneration independently, one would expect the various lineages to differ basically from each other, which they do not. On the other hand, if regeneration were inherent to metazoan life, a derivative of embryogenesis, its various expressions should be as much like each other as they resemble the development of embryonic appendage buds, which they do. It follows that the uneven distribution of regeneration must have been due to its extinction here and there, not as a negative adaptation by natural selection but as a pleiotropic epiphenomenon linked to more useful adaptations with which it was incompatible. In vertebrate evolution, these adaptations have included the transition from aquatic to terrestrial habitats and the modification of poikilothermic to homeothermic metabolism. The former advance rendered the regeneration of weight-bearing limbs impractical; the latter favored rapid wound healing and scar formation which effectively precluded blastema formation. If the latent capacity for regeneration persists in non-regenerative appendages, as would seem to be the case, then the restoration of its overt expression should be possible if the mechanisms of its inhibition could be discovered and eventually rendered ineffectual.     

Egg size in relation to body size in rotifers: an indication of reproductive strategy?

Egg sizes and body sizes of 43 egg-bearing rotifer species of numerous, mostly tropical, general have been recorded. Larger absolute egg volumes have been found for larger rotifer species, but the increase was lower than expected in proportion to body size, i.e. the relative egg volume decreased with increasing body size. Obviously the relative investment per offspring is smaller in larger rotifer species.     

The evolution of large‐scale body size clines in Plethodon salamanders: evidence of heat‐balance or species‐specific artifact?

A major goal in macroecology is to determine how body size varies geographically, and explain why such patterns exist. Recently, a grid‐cell assemblage analysis found significant body size trends with latitude and temperature in Plethodon salamanders, and support for the heat‐balance hypothesis as a possible explanation for these trends. Here we demonstrate that the heat‐balance hypothesis is unlikely to have generated this pattern, and that there is no overall body size trend with temperature in Plethodon. Using data from 3155 local Plethodon assemblages, we find no support for body size clines with latitude, and no relationship between body size and temperature. We also found that body size did not covary with elevation, in contrast to what was predicted by heat‐balance. We then examined the various scenarios under which body size clines across grid‐cell assemblages could evolve via heat‐balance, and found that none were tenable in light of the existing data. Instead, a single, widely distributed species was responsible for the pattern across grid‐cell assemblages. Finally, we examined why phylogenetic eigenvector regression does not account for phylogenetic non‐independence among taxa, and should not be used to account for shared evolutionary history in assembly‐level analyses. Assemblage‐level patterns are a useful means of assessing biogeographic trends, and are an important complement to within‐species and cross‐species patterns. However, while the use of grid‐cell assemblage approaches from digital databases is expedient, their results must be examined critically, and whenever possible, compared with data obtained from local species assemblages (particularly for ecological mechanisms that operate at the level of individuals). Finally, our results emphasize the importance of using corroborative data to evaluate alternative hypotheses, so that potential mechanisms that explain bioegeographic patterns are properly assigned.     

The role of ecological feedbacks in the evolution of host defence: what does theory tell us?

Hosts have evolved a diverse range of defence mechanisms in response to challenge by infectious organisms (parasites and pathogens). Whether defence is through avoidance of infection, control of the growth of the parasite once infected, clearance of the infection, tolerance to the disease caused by infection or innate and/or acquired immunity, it will have important implications for the population ecology (epidemiology) of the host-parasite interaction. As a consequence, it is important to understand the evolutionary dynamics of defence in the light of the ecological feedbacks that are intrinsic to the interaction. Here, we review the theoretical models that examine how these feedbacks influence the nature and extent of the defence that will evolve. We begin by briefly comparing different evolutionary modelling approaches and discuss in detail the modern game theoretical approach (adaptive dynamics) that allows ecological feedbacks to be taken into account. Next, we discuss a number of models of host defence in detail and, in particular, make a distinction between 'resistance' and 'tolerance'. Finally, we discuss coevolutionary models and the potential use of models that include genetic and game theoretical approaches. Our aim is to review theoretical approaches that investigate the evolution of defence and to explain how the type of defence and the costs associated with its acquisition are important in determining the level of defence that evolves.