Grandmother hypothesis and primate life histories

The adaptive significance of midlife menopause in human females has long engaged the attention of evolutionary anthropologists. In spite of extensive debate, the problem has only recently been examined in the context of primate life histories. Here I extend those investigations by comparing life history traits in 16 primate species to test predictions generated from life history theory. In humans, late ages of maturity and higher than expected birth rates are systematically associated with extended postmenopausal longevity. Links among these adjustments on the primate pattern can explain how selection could slow somatic senescence without favoring extension of the fertile span. This conclusion is consistent with the observation that our fertile spans are similar to those of other pongids. The shape of the argument herein demonstrates the utility of life history theory for solving problems of adaptive evolution in female life history traits, with consequences for broader arguments regarding human evolution.     

Hand asymmetry does not relate to key life history traits in post-menopausal contemporary Finnish women

Associations between fluctuating asymmetry (FA, a putative marker of developmental instability, DI) and life history traits have received a great deal of attention in the non-human literature. However, the patterns found are very heterogeneous and generalizations are difficult to make. In humans, only a few studies have related FA to life histories and fitness. In this paper we study such relationships using hand FA and several key life history traits in 209 post-menopausal Finnish women born between 1946 and 1958. Asymmetry measurements were based on scans of the hands and the life histories of these women were collected using questionnaires. No significant associations were detected and trends were opposite to expectations. We did find evidence for directional asymmetry, as traits in the right hand were larger on average. This may be due to handedness, questioning the usefulness of hand FA as a measure of DI. We conclude that future studies in humans should carefully examine the usefulness of traits as measures of DI.     

Molecular changes in fetal Down syndrome brain

Trisomy of human chromosome 21 is a major cause of mental retardation and other phenotypic abnormalities collectively known as Down syndrome. Down syndrome is associated with developmental failure followed by processes of neurodegeneration that are known to supervene later in life. Despite a widespread interest in Down syndrome, the cause of developmental failure is unclear. The brain of a child with Down syndrome develops differently from that of a normal one, although characteristic morphological differences have not been noted in prenatal life. On the other hand, a review of the existing literature indicates that there are a series of biochemical alterations occurring in fetal Down syndrome brain that could serve as substrate for morphological changes. We propose that these biochemical alterations represent and/or precede morphological changes. This review attempts to dissect these molecular changes and to explain how they may lead to mental retardation.     

Reconstructing life history of hominids and humans

Aspects of life history, such as processes and timing of development, age at maturation, and life span are consistently associated with one another across the animal kingdom. Species that develop rapidly tend to mature and reproduce early, have many offspring, and exhibit shorter life spans (r-selection) than those that develop slowly, have extended periods of premature growth, mature later in life, reproduce later and less frequently, have few offspring and/or single births, and exhibit extended life spans (K-selection). In general, primates are among the most K-selected of species. A suite of highly derived life history traits characterizes humans. Among these are physically immature neonates, slowed somatic development both in utero and post-natally, late attainment of reproductive maturity and first birth, and extended post-mature survival. Exactly when, why, and through what types of evolutionary interactions this suite arose is currently the subject of much conjecture and debate. Humankind's biocultural adaptations have helped to structure human life history evolution in unique ways not seen in other animal species. Among all species, life history traits may respond rapidly to alterations in selective pressures through hormonal processes. Selective pressures on life history likely varied widely among hominids and humans over their evolutionary history. This suggests that current patterns of human growth, development, maturation, reproduction, and post-mature survival may be of recent genesis, rather then long-standing adaptations. Thus, life history patterns observed among contemporary human and chimpanzee populations may provide little insight to those that existed earlier in hominid/human evolution.     

Disentangling the Evolution of Early and Late Life History Traits in Humans

Some aspects of human life history are unique among primates. Most notably, humans have a younger weaning age, a later age at first parturition, a shorter female reproductive period, and a longer lifespan than other living hominoid species. Obtaining a better understanding of when and how life history changed during human evolution is important to those studying the evolutionary developmental biology of extinct hominins, as life history traits pace developmental processes. Life history traits are thought to be linked via tradeoffs, such that changes in early life history traits directly affect those that follow later in life, and vice versa. However, it is also worth considering how changes to a single life history trait may indirectly affect other traits by way of modifying selective pressures acting on individuals and groups. For example, because they affect the size and demographic structure of a group, late life history traits (e.g., lifespan) may also affect the evolution of life history traits that occur earlier in life, but by modifying selective pressures acting on juveniles rather than by triggering a physiological tradeoff. This review marks an effort to begin to disentangle the ways in which early and late life history traits may affect each other both directly and indirectly. We concentrate on female life history characteristics. First, we review previous research on the evolution of the postmenopausal lifespan in women. Next we discuss recent findings concerning the relationship between the optimal length of the female reproductive period, mortality, and weaning age that show that selection favors a shorter female reproductive period in the presence of a younger weaning age. We discuss the implications this finding holds for understanding the evolution of life history traits that are of particular interest to developmental biologists.     

Life history focus on a malaria parasite: linked traits and variation among genetic clones

Life history theory has long been a major campaign in evolutionary ecology, but has typically focused only on animals and plants. Life history research on single-celled eukaryotic protists such as malaria parasites (Plasmodium) will offer new insights into the theory’s general utility as well as the parasite’s basic biology. For example, parasitologists have described the Plasmodium life cycle and cell types in exquisite detail, with little discussion of evolutionary issues such as developmental links between traits. We measured 10 life history traits of replicate single-genotype experimental Plasmodium mexicanum infections in its natural lizard host to identify groups of linked traits. These 10 traits formed 4 trait groups: “Rate/Peak” merges measures of growth rate and maximum parasitemia of infections; “Timing” combines time to patency and maximum parasitemia; “Growth Shape” describes the fit to an exponential growth curve; and “Sex Ratio” includes only the gametocyte sex ratio. Parasite genotype (clone) showed no effect on the life history trait groups, with the exception of gametocyte sex ratio. Therefore, variation in most life history traits among infections appears to be driven by environmental (individual host) effects. The findings support the model that life history traits are often linked by developmental constraints. Understanding why life history traits of Plasmodium are linked in this way would offer a new window into the evolution of the parasites, and also should inform public health efforts to control infection prevalence.     

Pooled energy budgets: Resituating human energy ‐allocation trade‐offs

Across taxa, many life‐history traits vary as a function of differences in body size. 1 - 5 Among primates, including humans, allometric relationships explain many trends in metabolic, growth, reproductive, and mortality rates. 6 - 8 But humans also deviate from nonhuman primates with respect to other developmental, reproductive, and parenting characteristics. 9 - 13 Broad relationships between life‐history traits and body size assume that energy expended in activity (foraging effort) is proportional to body size, and that energy available for growth and reproduction are equivalent. Because human subsistence and parenting are based on food sharing, and cooperation in labor and childrearing, the ways by which energy is acquired and allocated to alternate expenditures are expanded. We present a modification of the general allocation model to include a mechanism for these energy transfers. Our goal is to develop a framework that incorporates this mechanism and can explain the human life‐history paradox; that is, slow juvenile growth and rapid reproduction. We suggest that the central characteristics of human subsistence and energy transfer need to be accounted for in order to more fully appreciate human life‐history variability.     

Apes and Tricksters: The Evolution and Diversification of Humans’ Closest Relatives

The evolutionary history of humans comprises an important but small branch on the larger tree of ape evolution. Today’s hominoids—gibbons, orangutans, gorillas, chimpanzees, and humans—are a meager representation of the ape diversity that characterized the Old World from 23–5 million years ago. In this paper, I briefly review this evolutionary history focusing on features important for understanding modern ape and human origins. As the full complexity of ape evolution is beyond this review, I characterize major geographic, temporal, and phylogenetic groups using a few flagship taxa. Improving our knowledge of hominoid evolution both complicates and clarifies studies of human origins. On one hand, features thought to be unique to the human lineage find parallels in some fossil ape species, reducing their usefulness for identifying fossil humans. On the other hand, the Miocene record of fossil apes provides an important source for generating hypotheses about the ancestral human condition; this is particularly true given the dearth of fossils representing our closest living relatives: chimpanzees and gorillas.     

We age because we grow

Why do we age? Since ageing is a near-universal feature of complex organisms, a convincing theory must provide a robust evolutionary explanation for its ubiquity. This theory should be compatible with the physiological evidence that ageing is largely due to deterioration, which is, in principle, reversible through repair. Moreover, this theory should also explain why natural selection has favoured organisms that first improve with age (mortality rates decrease) and then deteriorate with age (mortality rates rise). We present a candidate for such a theory of life history, applied initially to a species with determinate growth. The model features both the quantity and the quality of somatic capital, where it is optimal to initially build up quantity, but to allow quality to deteriorate. The main theoretical result of the paper is that a life history where mortality decreases early in life and then increases late in life is evolutionarily optimal. In order to apply the model to humans, in particular, we include a budget constraint to allow intergenerational transfers. The resultant theory then accounts for all our basic demographic characteristics, including menopause with extended survival after reproduction has ceased.     

Lifetime reproductive effort in humans

Lifetime reproductive effort (LRE) measures the total amount of metabolized energy diverted to reproduction during the lifespan. LRE captures key components of the life history and is particularly useful for describing and comparing the life histories of different organisms. Given a simple energetic production constraint, LRE is predicted to be similar in value for very different life histories. However, humans have some unique ecological characteristics that may alter LRE, such as the long post-reproductive lifespan, lengthy juvenile period and the cooperative nature of human foraging and reproduction. We calculate LRE for natural fertility human populations, compare the findings to other mammals and discuss the implications for human life-history evolution. We find that human life-history traits combine to yield the theoretically predicted value (approx. 1.4). Thus, even with the subsidized energy budget and uniqueness of the adult lifespan, human reproductive strategies converge on the same optimal value of LRE. This suggests that the fundamental demographic variables contained in LRE trade-off against one another in a predictable and highly constrained manner.     

Human encephalization and developmental timing

Human evolution is frequently analyzed in the light of changes in developmental timing. Encephalization in particular has been frequently linked to the slow pace of development in Homo sapiens. The "brain allometry extension" theory postulates that the progressive extension of a conserved primate brain allometry into postnatal life was the basis for brain enlargement in the human lineage. This study shows that published primate and human growth data do not corroborate this model. Instead, the unique encephalization of H. sapiens is alternatively described as the result of evolutionary changes in three aspects of developmental timing. The first is a moderate extension in the duration of brain growth relative to our closest extant relatives, contrary to the view that human brain growth is drastically prolonged into postnatal life. Second, humans evolved a derived brain allometry in comparison with chimpanzees and early hominins. Third, humans (and other anthropoid primates to a lesser degree) display a significant retardation in early postnatal body growth in comparison with other mammals, which directly affects adult encephalization in our species. The rejection of the "brain allometry extension" model may require a reevaluation of the adaptive scenarios proposed to explain how human encephalization evolved.     

Heterogeneity in individual quality in birds: overall patterns and insights from a study on common terns

While life‐history theory predicts a tradeoff between reproduction and survival, positive covariance, indicative of heterogeneity in individual quality, is often reported among individuals from natural populations. We review longitudinal studies of wild bird populations that test the relationship between annual reproductive success and lifespan and find the majority to report a positive correlation, while none reports a negative correlation. Heterogeneity in individual quality in resource acquisition, masking resource‐based tradeoffs, therefore appears to be common in birds. Considering that there is little evidence for heritable variation in fitness, heterogeneity in individual quality among adults may be due to life‐long effects of developmental conditions. In a 20‐year case study on common terns Sterna hirundo, we test for life‐long effects of cohort quality and within‐cohort nest quality, but find no significant effects on long‐term proxies of quality. Since other studies do find strong life‐long effects of developmental conditions, we suggest that the brood reduction strategy adopted by common terns, causing the majority of offspring to die rapidly after hatching, efficiently reduces variation in offspring quality at independence. As such, a brood reduction strategy may contribute to reduced heterogeneity in adult survival in stochastic environments, both suggested to be more common and adaptive in long‐lived species. Further study is required to assess heterogeneity in individual reproduction, especially in relation to environmental stochasticity and species’ life‐history strategies, in order to assess whether the relative strength of selection in early and late life may indeed affect the magnitude of heterogeneity in individual quality over life, and how this is mediated by parent–offspring conflict.     

Mikrozephaliesyndrome und geistige Behinderung

Clarification of the cause of mental retardation, which has a prevalence of 2–3%, is a common reason for genetic consultation. On the basis of the cardinal sign of microcephaly, which also has a prevalence of 2–3%, an overview on different conditions with developmental delay/mental retardation is given according to the mode of inheritance. The current version of the Winter–Baraitser Dysmorphology Database lists 558 conditions with the combination of microcephaly and developmental delay/mental retardation. This makes clear that the following overview gives only a limited look at the comprehensive field of clinical genetics/dysmorphology.     

Grandmothers,hunters and human life history

This paper critiques the competing “Grandmother Hypothesis” and “Embodied Capital Theory” as evolutionary explanations of the peculiarities of human life history traits. Instead, I argue that the correct explanation for human life history probably involves elements of both hypotheses: long male developmental periods and lives probably evolved due to group selection for male hunting via increased female fertility, and female long lives due to the differential contribution women’s complex foraging skills made to their children and grandchildren’s nutritional status within groups provisioned by male hunting.     

Moral Darwinism: Ethical evidence for the descent of man

Could an ethical theory ever play a substantial evidential role in a scientific argument for an empirical hypothesis? InThe Descent of Man, Darwin includes an extended discussion of the nature of human morality, and the ethical theory which he sketches is not simply developed as an interesting ramification of his theory of evolution, but is used as a key part of his evidence for human descent from animal ancestors. Darwin must rebut the argument that, because of our moral nature, humans are essentially different in kind from other animals and so had to have had a different origin. I trace his causal story of how the moral sense could develop out of social instincts by evolutionary mechanisms of group selection, and show that the form of Utilitarianism he proposes involves a radical reduction of the standard of value to the concept of biological fitness. I argue that this causal analysis, although a weakness from a normative standpoint, is a strength when judged for its intended purpose as part of an evidential argument to confirm the hypothesis of human descent.     

The Developmental Basis of Quantitative Craniofacial Variation in Humans and Mice

The human skull is a complex and highly integrated structure that has long held the fascination of anthropologists and evolutionary biologists. Recent studies of the genetics of craniofacial variation reveal a very complex and multifactorial picture. These findings contrast with older ideas that posit much simpler developmental bases for variation in cranial morphology such as the growth of the brain or the growth of the chondrocranium relative to the dermatocranium. Such processes have been shown to have major effects on cranial morphology in mice. It is not known, however, whether they are relevant to explaining normal phenotypic variation in humans. To answer this question, we obtained vectors of shape change from mutant mouse models in which the developmental basis for the craniofacial phenotype is known to varying degrees, and compared these to a homologous dataset constructed from human crania obtained from a single population with a known genealogy. Our results show that the shape vectors associated with perturbations to chondrocranial growth, brain growth, and body size in mice do largely correspond to axes of covariation in humans. This finding supports the view that the developmental basis for craniofacial variation funnels down to a relatively small number of key developmental processes that are similar across mice and humans. Understanding these processes and how they influence craniofacial shape provides fundamental insights into the developmental basis for evolutionary change in the human skull as well as the developmental-genetic basis for normal phenotypic variation in craniofacial form.     

Les origines de l’art et les théories sur l’évolution humaine : le cas français

In recent years, we have witnessed an international debate about the question of the origins of art. On the one hand, some specialists have suggested that art appeared for the first time at the beginning of the Upper Paleolithic associated to the emergence of Homo sapiens sapiens. From this point of view, Paleolithic art as well as other hallmarks of behavioral modernity were exclusive to anatomically modern humans. On the other hand, some scholars have put into question the traditional paradigm concerning the origins of art and have suggested that artistic objects arose over a long period of time among different species, including Neanderthals. In order to contextualize this debate, we analyze in this article the history of the different interpretations and controversies concerning the question of the origins of art. Taking as reference the French case, we examine the connections between the different theories about art's origins suggested by Pleistocene art specialists during the last century and the dominant paradigms in human paleontology during the same period. Informed by one another, the question of the origins of art and that of human evolution seems to be inextricable linked.     

Age-dependency in hunting ability among the Ache of eastern Paraguay

This paper examines changes in hunting ability across the lifespan for the Ache of eastern Paraguay. Hunting ability is decomposed into two components-finding prey and probability of kill upon encounter- and analyzed for important prey species. Results support the argument that skill acquisition is an important aspect of the human foraging niche with hunting outcome variables reaching peaks surprisingly late in life, significantly after peaks in strength. The implications of this study are important for modeling the role of the human foraging niche in the co-evolution of various outstanding human life history characteristics such as large brains, long lifespans, and extended juvenile periods.     

Descent with modification: the unity underlying homology and homoplasy as seen through an analysis of development and evolution

Homology is at the foundation of comparative studies in biology at all levels from genes to phenotypes. Homology is similarity because of common descent and ancestry, homoplasy is similarity arrived at via independent evolution. However, given that there is but one tree of life, all organisms, and therefore all features of organisms, share some degree of relationship and similarity one to another. That sharing may be similarity or even identity of structure and the sharing of a most recent common ancestor--as in the homology of the arms of humans and apes--or it may reflect some (often small) degree of similarity, such as that between the wings of insects and the wings of birds, groups whose shared ancestor lies deep within the evolutionary history of the Metazoa. It may reflect sharing of entire developmental pathways, partial sharing, or divergent pathways. This review compares features classified as homologous with the classes of features normally grouped as homoplastic, the latter being convergence, parallelism, reversals, rudiments, vestiges, and atavisms. On the one hand, developmental mechanisms may be conserved, even when a complete structure does not form (rudiments, vestiges), or when a structure appears only in some individuals (atavisms). On the other hand, different developmental mechanisms can produce similar (homologous) features. Joint examination of nearness of relationship and degree of shared development reveals a continuum within an expanded category of homology, extending from homology --> reversals --> rudiments --> vestiges --> atavisms --> parallelism, with convergence as the only class of homoplasy, an idea that turns out to be surprisingly old. This realignment provides a glimmer of a way to bridge phylogenetic and developmental approaches to homology and homoplasy, a bridge that should provide a key pillar for evolutionary developmental biology (evo-devo). It will not, and in a practical sense cannot, alter how homoplastic features are identified in phylogenetic analyses. But seeing rudiments, reversals, vestiges, atavisms and parallelism as closer to homology than to homoplasy should guide us toward searching for the common elements underlying the formation of the phenotype (what some have called the deep homology of genetic and/or cellular mechanisms), rather than discussing features in terms of shared or independent evolution.     

The Deep Structure of Literary Representations

The cognitive rhetoricians have introduced the idea of cognitive domains into literary theory, but they have not yet developed a model for a comprehensive, species-typical structure of human motives. Evolutionary psychology can provide this model. Elemental human motives and basic emotions provide the deep structure of literary representations, and this deep structure serves to organize the particularities of circumstance and individual identity. Personal power and reproductive success are governing purposes in life and in literary representations. The concept of individual identity is necessary to literary representation, and a theory of literature based in evolutionary psychology has to incorporate models of personality. Literature and its oral antecedents organize experience in personally meaningful ways. They provide models of behavior and help regulate the complex cognitive machinery through which humans negotiate their social and cultural environments.