The importance of evolutionary constraints in ecological time scales

Summary The importance of constraints, defined as factors that retard or prevent a population from reaching its immediate adaptive peak on an ecological time scale is analysed. This is done by means of simple quantitative genetic models, which if anything underestimate the importance of constraints. The results show that even in the simplest case the response to selection will not generally be in the same direction as the selection vector, i.e. the direction to the nearest optimum. Adding complexity identifies cases where selection may lead the population in suboptimal directions. It is concluded that information about univariate genetic variances is not sufficient to predict evolutionary responses and may even be misleading. However, genetic covariances are not always acting as constraints, but can under certain circumstances promote evolution towards the nearest optimum. This can be understood by a spectral decomposition of the genetic variance—covariance matrix, where it is shown that the eigenvector associated with the largest amount of variance will to various degrees determine the outcome of selection. A literature survey of the pattern of character covariation in morphological characters in natural populations shows a wide variety of correlation patterns, but quite often shows a high level of covariance between traits. This suggests that constraints to short-term evolution may be more common than generally appreciated.     

The genetic covariance between life cycle stages separated by metamorphosis

Metamorphosis is common in animals, yet the genetic associations between life cycle stages are poorly understood. Given the radical changes that occur at metamorphosis, selection may differ before and after metamorphosis, and the extent that genetic associations between pre- and post-metamorphic traits constrain evolutionary change is a subject of considerable interest. In some instances, metamorphosis may allow the genetic decoupling of life cycle stages, whereas in others, metamorphosis could allow complementary responses to selection across the life cycle. Using a diallel breeding design, we measured viability at four ontogenetic stages (embryo, larval, juvenile and adult viability), in the ascidian Ciona intestinalis and examined the orientation of additive genetic variation with respect to the metamorphic boundary. We found support for one eigenvector of G (g_obsmax), which contrasted larval viability against embryo viability and juvenile viability. Target matrix rotation confirmed that while g_obsmax shows genetic associations can extend beyond metamorphosis, there is still considerable scope for decoupled phenotypic evolution. Therefore, although genetic associations across metamorphosis could limit that range of phenotypes that are attainable, traits on either side of the metamorphic boundary are capable of some independent evolutionary change in response to the divergent conditions encountered during each life cycle stage.     

Large body size in an island-dwelling bird: a microevolutionary analysis

Island races of passerine birds display repeated evolution towards larger body size compared with their continental ancestors. The Capricorn silvereye (Zosterops lateralis chlorocephalus) has become up to six phenotypic standard deviations bigger in several morphological measures since colonization of an island approximately 4000 years ago. We estimated the genetic variance-covariance (G) matrix using full-sib and 'animal model' analyses, and selection gradients, for six morphological traits under field conditions in three consecutive cohorts of nestlings. Significant levels of genetic variance were found for all traits. Significant directional selection was detected for wing and tail lengths in one year and quadratic selection on culmen depth in another year. Although selection gradients on many traits were negative, the predicted evolutionary response to selection of these traits for all cohorts was uniformly positive. These results indicate that the G matrix and predicted evolutionary responses are consistent with those of a population evolving in the manner observed in the island passerine trend, that is, towards larger body size.     

Strong environmental determination of a carotenoid-based plumage trait is not mediated by carotenoid availability

Carotenoid-based colours are recognized as having an important signalling function, yet the nature of the mechanisms that maintain their honesty is not well understood. By combining a carotenoid-feeding experiment with a quantitative genetic experiment in a wild population of blue tits (Parus caeruleus), we were able to test predictions that differentiate between proposed mechanisms. If variation in carotenoid ingestion underlies variation in carotenoid-based colour expression, then carotenoid-supplemented birds should have reduced variance in colour. In this study, carotenoid supplementation produced a small but significant change in plumage colouration, but no significant change in variation. These results suggest that variation in carotenoid acquisition is not an important source of variation for this colour trait, and that variation in post-ingestion processes are likely to be more important. The low heritability of this colour trait suggests environmental factors are likely to underlie the majority of variation in these processes.     

Testing the phenotypic gambit: phenotypic, genetic and environmental correlations of colour

Evolutionary theory is primarily concerned with genetic processes, yet empirical testing of this theory often involves data collected on phenotypes. To make this tenable, the implicit assumption is often made that phenotypic patterns are good predictors of genetic patterns; an assumption that coined the phenotypic gambit. Although this assumption has been validated for traits with high heritability, such as morphology, its generality for traits with low heritabilities, such as life-history and behavioural traits, remains controversial. Using a large-scale cross-fostering experiment, we were able to measure genetic, common environmental and phenotypic correlations between four colour traits and two skeletal traits in a wild population of passerine birds, the blue tit (Parus caeruleus). Colour traits had little heritable variation but common environment effects were found to be important; skeletal traits showed the opposite pattern. Positive correlations because of a shared natal environment were found between all traits, obscuring negative genetic correlations between some colour and skeletal traits. Consequently, phenotypic patterns were poor surrogates for genetic patterns and we suggest that this may be common if trade-offs or substantial parental effects exist. For this group of traits, the phenotypic gambit cannot be made and we suggest caution when inferring genetic patterns from phenotypic data, especially for behavioural and life-history traits.     

Competing Models of Stability in Complex, Evolving Systems: Kauffman vs. Simon

I criticize Herbert Simon's argument for the claim that complex natural systems must constitute decomposable, mereological or functional hierarchies. The argument depends on certain assumptions about the requirements for the successful evolution of complex systems, most importantly, the existence of stable, intermediate stages in evolution. Simon offers an abstract model of any process that succeeds in meeting these requirements. This model necessarily involves construction through a decomposable hierarchy, and thus suggests that any complex, natural, i.e., evolved, system is constituted by a decomposable hierarchy. I argue that Stuart Kauffman's recent models of genetic regulatory networks succeed in specifying processes that could meet Simon's requirements for evolvability without requiring construction through a decomposable hierarchy. Since Kauffman's models are at least as plausible as Simon's model, Simon's argument that complex natural systems must constitute decomposable, mereological or functional hierarchies does not succeed.     

Tumor necrosis factor-alpha in serum of macaques during SIVmac251 acute infection

TNF secretion was explored in sera during acute SIV-infection of cynomolgus macaques. A peak of TNF was detected in sera of animals in concomitance with SIV replication. Likewise, AZT treatment delayed and reduced peaks of viral replication and TNF production. Thus, SIVmac251-infected monkey could be an excellent model to explore the interdigitation existing between HIV and TNF in acute and chronic infection and to develop new therapeutic strategies that target the production of this cytokine or its inductive effects.