Variability selection in hominid evolution

Variability selection (abbreviated as VS) is a process considered to link adaptive change to large degrees of environment variability. Its application to hominid evolution is based, in part, on the pronounced rise in environmental remodeling that took place over the past several million years. The VS hypothesis differs from prior views of hominid evolution, which stress the consistent selective effects associated with specific habitats or directional trends (e.g., woodland, savanna expansion, cooling). According to the VS hypothesis, wide fluctuations over time created a growing disparity in adaptive conditions. Inconsistency in selection eventually caused habitat-specific adaptations to be replaced by structures and behaviors responsive to complex environmental change. Key hominid adaptations, in fact, emerged during times of heightened variability. Early bipedality, encephalized brains, and complex human sociality appear to signify a sequence of VS adaptations—i.e., a ratcheting up of versatility and responsiveness to novel environments experienced over the past 6 million years. The adaptive results of VS cannot be extrapolated from selection within a single environmental shift or relatively stable habitat. If some complex traits indeed require disparities in adaptive setting (and relative fitness) in order to evolve, the VS idea counters the prevailing view that adaptive change necessitates long-term, directional consistency in selection. © 1998 Wiley-Liss, Inc.     

Mutation-biased adaptation in a protein NK model

Evolutionary trends responsible for systematic differences in genome and proteome composition have been attributed to GC:AT mutation bias in the context of neutral evolution or to selection acting on genome composition. A possibility that has been ignored, presumably because it is part of neither the Modern Synthesis nor the Neutral Theory, is that mutation may impose a directional bias on adaptation. This possibility is explored here with simulations of the effect of a GC:AT bias on amino acid composition during adaptive walks on an abstract protein fitness landscape called an "NK" model. The results indicate that adaptation does not preclude mutation-biased evolution. In the complete absence of neutral evolution, a modest GC:AT bias of realistic magnitude can displace the trajectory of adaptation in a mutationally favored direction, to such a degree that amino acid composition is biased substantially and persistently. Thus, mutational explanations for evolved patterns need not presuppose neutral evolution.     

Early paleolithic of China and eastern Asia

In attempting to understand the course of human evolution and the nature of hominid adaptation over the past few million years, it is necessary to consider prevailing evidence from all parts of the world. Eastern Asia provides a range of important questions and challenges with regard to this evolutionary puzzle. Although evidence for earlier ape evolution is present in China (for example, at Lufeng in Yunnan Province), the earliest evidence for hominid presence appears to be in the Early Pleistocene, apparently the result of a migration of hominids to and subsequent adaptation within Eastern Asia. The archeological record provides a closer look at some technological aspects of this adaptation during the Early and Middle Pleistocene, showing both distinctive contrasts and intriguing continuities relative to the rest of the Old World.     

Observed and expected variation in hominid evolution

All of the major groups of fossil hominids (australopithecines, pithecanthropines, Neandertals, and early sapiens) were discovered by 1925, and therefore prior to the formulation of the synthetic theory of evolution that revolutionized the concept of the species in systematics. While these fossil finds were being made the framework for their interpretation included several assumptions: (1) that the number of living hominoid species was great, and that intraspecific variation was slight (authoritative sources recognized as many as 14 separate species of chimpanzees and 15 species of gorillas); (2) that the timescale of human evolution was brief (measured in tens or hundreds of thousands of years). As a result of these premises the consensus that hominid evolution was characterized by a large number of sympatric and synchronic species was virtually inevitable.In contrast, recent molecular studies demonstrate that genetic diversity among recent hominoids is so slight that even humans and chimpanzees differ at only about 1% of the loci that have been sampled so far; evidently, very small genetic differences can produce rather great contrasts in morphology. At the same time, geological break-throughs have increased the timescale for human evolution to several million years.It is concluded that morphological differences among fossil hominids, even if very appreciable and complex, do not necessarily reflect a great degree of either genetic or taxonomic diversity. Potential effects of evolutionary change through time should be incorporated into models of hominid evolution as a means of assessing the minimum number of lineages required to account for observed variations among hominid specimens.     

Rates of evolution: Is there a conflict between neo-darwinian evolutionary theory and the fossil record?

Neo-darwinian and population genetics theory assumes that the necessary and sufficient set of conditions for all genetic, therefore evolutionary, change has been identified. Punctuationalists have assumed the opposite and cite the fossil record as evidence for change too rapid to be explained in neo-darwinian theory. Data is given here to provide estimates of the rate of evolution in hominid fossils, in living populations, and of that rate which would qualify as punctuational in the hominid fossil record. Evolution in living populations is orders of magnitude greater than that found in the fossil record and far greater than necessary to create apparently instantaneous saltations in the fossil record. It is suggested that such saltations may not represent more rapid rates of evolution but, rather, the persistence of evolutionary change in a given direction for a longer than normal period.     

化石人类脑演化研究概况

脑演化是人类演化的一个重要组成部分,其研究可以为人类起源、演化、人群关系及语言、智力等方面提供重要的信息。脑演化的主要证据是通过研究颅内模（endocast）及颅骨的形态得到的。颅内模是从颅骨内表面得到的脑的外部形态。有时颅骨的内腔充满泥沙,并且为钙质所结固,可以自然形成颅内模。也可以人工制作颅内模。颅内模和颅骨的内表面能够提供人类脑髓及神经进化方面的直接证据。对化石人类脑演化的研究主要包括以下几个方面的内容：测量或估计脑量的大小及其和身体大小之间的关系;研究脑量随时间的变化过程;通过对早期人类颅内模表面沟回形态特征的研究,探索脑功能区在早期人类和猿类的区别及在演化上的变化;左右大脑不对称性与一侧优势关系,探讨语言的起源和惯用手的脑功能基础等;脑膜中动脉系统、静脉窦系统及与血液循环相关的排泄孔的变化,探讨大脑各部分比例的变化和功能的日益复杂对供血需求的影响;通过对人类脑演化的研究,探讨人类进化的原因。本文通过对以上几个方面及其中国化石脑演化研究的介绍,对化石人类脑演化的研究概况作综合论述和简单回顾。     

Restrictions to RNA virus adaptation: an experimental approach

Some basic properties of RNA viruses are their high mutation rate, their enormous population sizes and their short generation time. These properties allow RNA virus populations to quickly explore fitness landscapes. A great adaptability has been amply demonstrated in experimental, as well as in natural, populations of RNA viruses. However, at least from a theoretical point of view, a limit to the extent of viral adaptation may exist as a consequence of adaptive trade-offs arising during evolution in changing environmental conditions. Here, I review previously published results searching for such fitness trade-offs. The following scenario has been explored: the cost of host-range expansion, the cost of resistance to antiviral drugs, and the adaptation to different population densities. Despite the environmental conditions tested, results show a common pattern: whenever a virus adapt to a simple environmental situation it pays a cost in terms of adaptation to alternative situations. However, in those cases where the virus has been simultaneously adapted to different environmental conditions, this cost disappears or, at least, is greatly reduced. Finally, and as another factor imposing a limit to their speed of adaptation, I review results showing that clonal interference also plays an important role during viral evolution. This revised version was published online in August 2006 with corrections to the Cover Date.     

The evolution of human fatness and susceptibility to obesity: an ethological approach

Human susceptibility to obesity is an unusual phenomenon amongst animals. An evolutionary analysis, identifying factors favouring the capacity for fat deposition, may aid in the development of preventive public health strategies. This article considers the proximate causes, ontogeny, fitness value and evolutionary history of human fat deposition. Proximate causes include diet composition, physical activity level, feeding behaviour, endocrine and genetic factors, psychological traits, and exposure to broader environmental factors. Fat deposition peaks during late gestation and early infancy, and again during adolescence in females. As in other species, human fat stores not only buffer malnutrition, but also regulate reproduction and immune function, and are subject to sexual selection. Nevertheless, our characteristic ontogenetic pattern of fat deposition, along with relatively high fatness in adulthood, contrasts with the phenotype of other mammals occupying the tropical savannah environment in which hominids evolved. The increased value of energy stores in our species can be attributed to factors increasing either uncertainty in energy availability, or vulnerability to that uncertainty. Early hominid evolution was characterised by adaptation to a more seasonal environment, when selection would have favoured general thriftiness. The evolution of the large expensive brain in the genus Homo then favoured increased energy stores in the reproducing female, and in the offspring in early life. More recently, the introduction of agriculture has had three significant effects: exposure to regular famine; adaptation to a variety of local niches favouring population-specific adaptations; and the development of social hierarchies which predispose to differential exposure to environmental pressures. Thus, humans have persistently encountered greater energy stress than that experienced by their closest living relatives during recent evolution. The capacity to accumulate fat has therefore been a major adaptive feature of our species, but is now increasingly maladaptive in the modern environment where fluctuations in energy supply have been minimised, and productivity is dependent on mechanisation rather than physical effort. Alterations to the obesogenic environment are predicted to play a key role in reducing the prevalence of obesity.     

The first rib of hominoids

Homo sapiens is unique among extant hominoids in displaying a univertebral articular pattern for the first rib; that is, the head of the first rib articulates only with the body of the first thoracic vertebra. All other hominoids, indeed virtually all other mammals, display a bivertebral pattern; that is, the head of the first rib articulates with the bodies of both the seventh cervical and the first thoracic vertebrae, as well as the intervening disk. Two fossil hominid partial first ribs, A.L. 288-lax and A.L. 333-118, show that the univertebral pattern was fully established in the hominid lineage by the appearance of Australopithecus afarensis. Four hypotheses, based in functional anatomy, can be postulated for the evolution of the univertebral pattern: (1), it increases the volume (via increased length) of the neck, which could, in turn, compensate for the functional loss of the laryngeal sac systems in hominid vocalization; (2), it is a consequence of the more barrel-shaped thorax in hominids; (3), it is a consequence of functional modifications in the hominid shoulder girdle; and/or (4), it is a consequence of modifications in hominid first rib motion while breathing in an upright stance. Fossil evidence supports all but the first hypothesis, and most strongly supports the third. However, evidence for the first hypothesis does suggest that the evolution of descent of the upper respiratory system in the hominid lineage may have been permitted by the presence of the univertebral pattern, while the reverse is probably not true. Furthermore, fossil evidence for the third hypothesis shows that, by the appearance of A. afarensis, the hominid upper limb had been freed from locomotor constraints, which concomitantly confirms full adaptation to upright posture. Thus, because of their potential relationship with upright posture, the two remaining hypotheses (i.e., "thoracic shape" and "first rib movement during breathing") also have support from the fossil evidence.     

FROM MORE TO FEWER? TESTING AN ALLEGEDLY PERVASIVE TREND IN THE EVOLUTION OF MORPHOLOGICAL STRUCTURE

While evolutionary trends have long received much attention and have been widely disputed, new methods are now allowing the testing of directional hypotheses with increased rigor. Here, we test a general hypothesis about the way many kinds of discrete characters are thought to evolve, termed oligomerization. This is the tendency for serial structures (such as arthropod body and appendage segments) or armature (such as spines) to evolve primarily through loss and fusion. Focusing on the Crustacea, we use maximum likelihood methods to test for directional evolution in a large sample (> 500) of discrete traits, analyzed against molecular-based phylogenies. We find evidence for a significant trend toward trait loss, in accordance with the reduction principle. However, this trend is far from ubiquitous, with many characters exhibiting a reconstructed bias toward gains. These results suggest that caution must be used before drawing conclusions about which taxa are "primitive" or about the directionality of morphological shifts in the absence of phylogenetic analysis. Nevertheless, oligomerization-as a trend rather than a law-may be an important process that influences evolutionary trajectories from both morphological and functional perspectives.