Defence of bipedalism

The long-unresolved and much publicized puzzle of how human ancestors could have ventured upon habitual bipedalism without making themselves highly vulnerable to predation derives from a misunderstanding of the principles upon which predator-prey relations work. Erect stature, regardless of locomotion proficiency, in fact enhances passive, and not simply active, defence against predators. This was the necessary condition, the security ‘cover’, for selection for the elaboration of bipedal behavior. As illustration, I analyse the defence systems ofPan troglodytes and the hunting techniques of their predators, known and potential, in order to hypothesize the defence impact of bipedal selection on the former; the relevance of human experience with felid predators, in particular, is also then explored. Despite known major differences in the late Miocene mammalian predator guild, the predator-prey framework is likely to have been much the same as to-day, making the findings applicable to the situation of proto-hominids. Once it is accepted selection pressure for bipedalism was originally focused on posture and not locomotion, the mystery of its emergence disappears.     

Trajectories and Constraints in Brain Evolution in Primates and Cetaceans

A negative allometric relationship between body mass (BM) and brain size (BS) can be observed for many vertebrate groups. In the past decades, researchers have proposed several hypotheses to explain this finding, but none is definitive and some are possibly not mutually exclusive. Certain species diverge markedly (positively or negatively) from the mean of the ratio BM/BS expected for a particular taxonomic group. It is possible to define encephalization quotient (EQ) as the ratio between the actual BS and the expected brain size. Several cetacean species show higher EQs compared to all primates, except modern humans. The process that led to big brains in primates and cetaceans produced different trajectories, as shown by the organizational differences observed in every encephalic district (e.g., the cortex). However, these two groups both convergently developed complex cognitive abilities. The comparative study on the trajectories through which the encephalization process has independently evolved in primates and cetaceans allows a critical appraisal of the causes, the time and the mode of quantitative and qualitative development of the brain in our species and in the hominid evolutionary lineage.     

Testing some hypotheses on Human Evolution and sexual dimorphism

Research on human evolution and sexual dimorphism motivates an interesting test problem. In studying hominid phylogeny it is of interest to test whether parallel evolution plays a role. With regard to sexual dimorphism it is of interest to know whether the directions of sexual dimorphism in the populations being compared are the same. We show that testing these two problems gives rise to the same type of hypothesis testing, viz. the problem of testing the hypothesis that the means of independent, normally distributed random vectors with unit covariance matrices are situated on a straight line through the origin. A test is proposed and applied to study the sexual dimorphism of 20 recent skull populations. In this example the hypothesis of equal directions of sexual dimorphism is rejected. The classical theory of constructing multiple discriminant functions (canonical variates) is adapted to the problem of comparing sexual dimorphisms.     

Choosing among alternative trees of multigene families

Estimation of gene trees is the first step in testing alternative hypotheses about the evolution of multigene families. The standard practice for inferring gene family history is to construct trees that meet some objective criteria based on the fit of the character state changes (nucleotide or amino acid changes) to the gene tree. Unfortunately, analysis of character state data can be misleading. In addition, this approach ignores information about the relationships of the species from which the genes have been sampled. In this paper I explore using statistics of fit between the character data and gene trees and the reconciliation of the gene and species trees for choosing among alternative evolutionary hypotheses of gene families. In particular, I advocate a two-pronged strategy for choosing among alternative gene trees. First, the character data are used to define a set of acceptable gene trees (i.e., trees that are not significantly different from the minimum length tree). Next, the set of acceptable gene trees is reconciled with a known species tree, and the gene tree requiring the fewest number of gene duplications and losses is adopted as the best estimate of evolutionary history. The approach is illustrated using three gene families: BMP, EGR, and LDH.     

Increased brain size in mammals is associated with size variations in gene families with cell signalling,chemotaxis and immune-related functions

Genomic determinants underlying increased encephalization across mammalian lineages are unknown. Whole genome comparisons have revealed large and frequent changes in the size of gene families, and it has been proposed that these variations could play a major role in shaping morphological and physiological differences among species. Using a genome-wide comparative approach, we examined changes in gene family size (GFS) and degree of encephalization in 39 fully sequenced mammalian species and found a significant over-representation of GFS variations in line with increased encephalization in mammals. We found that this relationship is not accounted for by known correlates of brain size such as maximum lifespan or body size and is not explained by phylogenetic relatedness. Genes involved in chemotaxis, immune regulation and cell signalling-related functions are significantly over-represented among those gene families most highly correlated with encephalization. Genes within these families are prominently expressed in the human brain, particularly the cortex, and organized in co-expression modules that display distinct temporal patterns of expression in the developing cortex. Our results suggest that changes in GFS associated with encephalization represent an evolutionary response to the specific functional requirements underlying increased brain size in mammals.     

Fallacies of progression in theories of brain-size evolution

The tacit assumption that relative enlargement and differentiation of brains reflect a progressive evolutionary trend toward greater intelligence is a major impediment to the study of brain evolution. Theories that purport to establish a linear scale for this presumed correlation between brain size and intelligence are undermined by the absence of an unbiased allometric baseline for estimating differences in encephalization, by the incompatibility of allometric analyses at different taxonomic levels, by the nonlinearity of the criterion of subtraction used to partition the somatic and cognitive components of encephalization, and by the failure to independently demonstrate any cognitive basis for the regularity of brain/body allometry. Analyzing deviations from brain/body allometric trends in terms of encephalization obfuscates the complementarity between brain and body size and ignores selection on body size, which probably determines most deviations. By failing to analyze the effects of allometry at many levels of structure, comparative anatomists have mistaken methodological artifacts for progressive evolutionary trends. Many structural changes, which are assumed to demonstrate progression of brain structure from primitive to advanced forms, are the results of allometric processes. Increased brain size turns out to have some previously unappreciated functional disadvantages.     

Special Issue ofHuman evolution on intelligence and evolutionary biology

This is a Special Issue on intelligence and evolutionary biology, based on selected lectures at a NATO Advanced Study Institute on this topic. The proceedings of the ASI have been published in a separate volume. The papers presented here have been reviewed and updated to reflect information available in 1988.     

Arboreality,terrestriality and bipedalism

The full publication of Ardipithecus ramidus has particular importance for the origins of hominin bipedality, and strengthens the growing case for an arboreal origin. Palaeontological techniques however inevitably concentrate on details of fragmentary postcranial bones and can benefit from a whole-animal perspective. This can be provided by field studies of locomotor behaviour, which provide a real-world perspective of adaptive context, against which conclusions drawn from palaeontology and comparative osteology may be assessed and honed. Increasingly sophisticated dynamic modelling techniques, validated against experimental data for living animals, offer a different perspective where evolutionary and virtual ablation experiments, impossible for living mammals, may be run in silico, and these can analyse not only the interactions and behaviour of rigid segments but increasingly the effects of compliance, which are of crucial importance in guiding the evolution of an arboreally derived lineage.     

No evidence for the 'expensive-tissue hypothesis' from an intraspecific study in a highly variable species

The 'expensive-tissue hypothesis' states that investment in one metabolically costly tissue necessitates decreased investment in other tissues and has been one of the keystone concepts used in studying the evolution of metabolically expensive tissues. The trade-offs expected under this hypothesis have been investigated in comparative studies in a number of clades, yet support for the hypothesis is mixed. Nevertheless, the expensive-tissue hypothesis has been used to explain everything from the evolution of the human brain to patterns of reproductive investment in bats. The ambiguous support for the hypothesis may be due to interspecific differences in selection, which could lead to spurious results both positive and negative. To control for this, we conduct a study of trade-offs within a single species, Thalassoma bifasciatum, a coral reef fish that exhibits more intraspecific variation in a single tissue (testes) than is seen across many of the clades previously analysed in studies of tissue investment. This constitutes a robust test of the constraints posited under the expensive-tissue hypothesis that is not affected by many of the factors that may confound interspecific studies. However, we find no evidence of trade-offs between investment in testes and investment in liver or brain, which are typically considered to be metabolically expensive. Our results demonstrate that the frequent rejection of the expensive-tissue hypothesis may not be an artefact of interspecific differences in selection and suggests that organisms may be capable of compensating for substantial changes in tissue investment without sacrificing mass in other expensive tissues.     

What can dolphins tell us about primate evolution?

Fifty-five million years ago, a furry, hoofed mammal about the size of a dog ventured into the shallow brackish remnant of the Tethys Sea and set its descendants on a path that would lead to their complete abandonment of the land. These early ancestors of cetaceans (dolphins, porpoises, and whales) thereafter set on an evolutionary course that is arguably the most unusual of any mammal that ever lived. Primates and cetaceans, because of their adaptation to exclusively different physical environments, have had essentially nothing to do with each other throughout their evolution as distinct orders. In fact, the closest phylogenetic relatives of cetaceans are even-toed ungulates.