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The evolution of human speech: the role of enhanced breathing control.

Many cognitive and physical features must have undergone change for the evolution of fully modern human language. One neglected aspect is the evolution of increased breathing control. Evidence presented herein shows that modern humans and Neanderthals have an expanded thoracic vertebral canal compared with australopithecines and Homo ergaster, who had canals of the same relative size as extant nonhuman primates. Based on previously published analyses, these results demonstrate that there was an increase in thoracic innervation during human evolution. Possible explanations for this increase include postural control for bipedalism, increased difficulty of parturition, respiration for endurance running, an aquatic phase, and choking avoidance. These can all be ruled out, either because of their evolutionary timing, or because they are insufficiently demanding neurologically. The remaining possible functional cause is increased control of breathing for speech. The main muscles involved in the fine control of human speech breathing are the intercostals and a set of abdominal muscles which are all thoracically innervated. Modifications to quiet breathing are essential for modern human speech, enabling the production of long phrases on single expirations punctuated with quick inspirations at meaningful linguistic breaks. Other linguistically important features affected by variation in subglottal air pressure include emphasis of particular sound units, and control of pitch and intonation. Subtle, complex muscle movements, integrated with cognitive factors, are involved. The vocalizations of nonhuman primates involve markedly less respiratory control. Without sophisticated breath control, early hominids would only have been capable of short, unmodulated utterances, like those of extant nonhuman primates. Fine respiratory control, a necessary component for fully modern language, evolved sometime between 1.6 Mya and 100,000 ya.

Animals↗

NADP-dependent enzymes. II: Evolution of the mono- and dinucleotide binding domains.

Nicotinamide adenine dinucleotides [NAD and NADP with both referred to as NAD(P)] are among the more diffuse redox cofactors. Despite their stereochemical similarity where the only difference is a phosphomonoester on the ribose near the adenine of NADP, they show different biochemical reactivities with NAD behaving as an oxidant and NADP as a reductant. NAD(P)-dependent enzymes generally share a common open alpha/beta fold with few exceptions only recently structurally characterized. This study of the molecular evolution of the NAD(P) binding domains, possible given the large number of known molecular structures, addresses two main questions: 1) can a common fold exist in different biological systems (divergent evolution) and 2) does a relationship exist among similar biological systems that display different folds (convergent evolution)? Both the structures of mono- and dinucleotide binding domains have been classified by cluster analysis based on the similarity evaluated by their main chain C alpha superposition. Moreover, the cofactor conformations and the stereochemical characteristics of their pockets have also been classified by analogous methods on the basis of the published tertiary structures. Two primary results appear: 1) the classification of the mononucleotide binding domains is different from that of the dinucleotide binding folds and 2) both divergent and convergent evolutionary pathways can be hypothesized, the latter less frequently observed and less pronounced but nevertheless evident. The generally accepted hypothesis that dinucleotide binding domains have evolved by gene duplication of primordial genes coding for the smaller mononucleotide binding domains is acceptable but the two halves of the resulting dinucleotide binding domains are evolutionarily uncorrelated. The NH2-terminal mononucleotide binding domain is less variable than the COOH-terminal half, probably because it involves the binding of the ADP moiety of NAD(P) invariant in all examined systems. There is evidence to postulate that evolutionary pathways for NAD(P)-dependent enzymes are both divergent and convergent. In fact, nearly all combinations of similarity dissimilarity in overall fold, cofactor conformation, and cofactor binding pocket structural characteristics for each enzyme pair examined are possible. The NAD(P)-dependent enzymes apparently provide a canonical example of an evolutionary principle that "anything goes."

Animals↗

Progress in the study of brain evolution: from speculative theories to testable hypotheses.

Darwin's theory of evolution raised the question of how the human brain differs from that of other animals and how it is the same. Early students of brain evolution had constructed rather grand but speculative theories which stated that brains evolved in a linear manner, from fish to man and from simple to complex. These speculations were soundly refuted, however, as contemporary comparative neurobiologists used powerful new techniques and methodologies to discover that complex brains have evolved several times independently among vertebrates (e.g., within teleost fishes and birds) and that brain complexity has actually decreased in the lineages leading to modern salamanders and lungfishes. Moreover, the old idea that brains evolved by the sequential addition of new components has now been replaced by the working hypothesis that brains generally evolve by the divergent modification of preexisting parts. Speculative theories have thus been replaced by testable hypotheses, and current efforts in the field are aimed at making phylogenetic hypotheses even more testable. Particularly promising new directions for comparative neurobiology include (1) the integration of comparative neuroanatomy with comparative embryology and developmental genetics in order to test phylogenetic hypotheses at a mechanistic level, (2) research into how evolutionary changes in the structure of neural circuits are related to evolutionary changes in circuit function and animal behavior, and (3) the analysis of independently evolved similarities to discover general rules about how brains may or may not change during the course of evolution.

Animals↗

The molecular evolution of development.

Morphological differences between species, from simple single-character differences to large-scale variation in body plans, can be traced to changes in the timing and location of developmental events. This has led to a growing interest in understanding the genetic basis behind the evolution of developmental systems. Molecular evolutionary genetics provides one of several approaches to dissecting the evolution of developmental systems, by allowing us to reconstruct the history of developmental genetic pathways, infer the origin and diversification of developmental gene functions, and assess the relative contributions of various evolutionary forces in shaping regulatory gene evolution.

Animal Population Groups↗

Molecular strategies of tooth enamel formation are highly conserved during vertebrate evolution.

The vertebrate body plan is determined by a variety of morphoregulatory genes that are highly conserved throughout evolution. This review presents a phylogenetic analysis of selected molecular and morphological features in vertebrates with particular emphasis upon the phylogeny of tooth morphogenesis and enamel formation. Three lines of evidence support our hypothesis that the agnathans (e.g. hagfishes) are the most primitive extant vertebrates and that enamel gene products are highly conserved during vertebrate evolution. First, an antibody raised against the polypeptide produced by exon 4 of the mouse amelogenin gene recognizes proteins in hagfish, sharks, reptiles and mammals. Second, electron photomicrographic evidence suggests heterochronic shifts in the relative time and rate of enamel formation during vertebrate tooth evolution. Third, mRNA phenotyping suggests significant homology between amelogenin transcripts expressed in species of various vertebrate phyla including agnathans and mammals. These three lines of evidence indicate that amelogenin gene products are expressed in agnathan, reptilian and mammalian teeth.

Ameloblasts↗

Mechanisms of protein evolution and their application to protein engineering.

Protein engineering holds great promise for the development of new biosensors, diagnostics, therapeutics, and agents for bioremediation. Despite some remarkable successes in experimental and computational protein design, engineered proteins rarely achieve the efficiency or specificity of natural enzymes. Current protein design methods utilize evolutionary concepts, including mutation, recombination, and selection, but the inability to fully recapitulate the success of natural evolution suggests that some evolutionary principles have not been fully exploited. One aspect of protein engineering that has received little attention is how to select the most promising proteins to serve as templates, or scaffolds, for engineering. Two evolutionary concepts that could provide a rational basis for template selection are the conservation of catalytic mechanisms and functional promiscuity. Knowledge of the catalytic motifs responsible for conserved aspects of catalysis in mechanistically diverse superfamilies could be used to identify promising templates for protein engineering. Second, protein evolution often proceeds through promiscuous intermediates, suggesting that templates which are naturally promiscuous for a target reaction could enhance protein engineering strategies. This review explores these ideas and alternative hypotheses concerning protein evolution and engineering. Future research will determine if application of these principles will lead to a protein engineering methodology governed by predictable rules for designing efficient, novel catalysts.

Binding Sites↗

The aging brain: the cognitive reserve hypothesis and hominid evolution.

Compared to other primates, humans live a long time and have large brains. Recent theories of the evolution of human life history stages (grandmother hypothesis, intergenerational transfer of information) lend credence to the notion that selection for increased life span and menopause has occurred in hominid evolution, despite the reduction in the force of natural selection operating on older, especially post-reproductive, individuals. Theories that posit the importance (in an inclusive fitness sense) of the survival of older individuals require them to maintain a reasonably high level of cognitive function (e.g., memory, communication). Patterns of brain aging and factors associated with healthy brain aging should be relevant to this issue. Recent neuroimaging research suggests that, in healthy aging, human brain volume (gray and white matter) is well-maintained until at least 60 years of age; cognitive function also shows only nonsignificant declines at this age. The maintenance of brain volume and cognitive performance is consistent with the idea of a significant post- or late-reproductive life history stage. A clinical model, "the cognitive reserve hypothesis," proposes that both increased brain volume and enhanced cognitive ability may contribute to healthy brain aging, reducing the likelihood of developing dementia. Selection for increased brain size and increased cognitive ability in hominid evolution may therefore have been important in selection for increased lifespan in the context of intergenerational social support networks.

Aging↗

On the nature and evolution of the neural bases of human language.

The traditional theory equating the brain bases of language with Broca's and Wernicke's neocortical areas is wrong. Neural circuits linking activity in anatomically segregated populations of neurons in subcortical structures and the neocortex throughout the human brain regulate complex behaviors such as walking, talking, and comprehending the meaning of sentences. When we hear or read a word, neural structures involved in the perception or real-world associations of the word are activated as well as posterior cortical regions adjacent to Wernicke's area. Many areas of the neocortex and subcortical structures support the cortical-striatal-cortical circuits that confer complex syntactic ability, speech production, and a large vocabulary. However, many of these structures also form part of the neural circuits regulating other aspects of behavior. For example, the basal ganglia, which regulate motor control, are also crucial elements in the circuits that confer human linguistic ability and abstract reasoning. The cerebellum, traditionally associated with motor control, is active in motor learning. The basal ganglia are also key elements in reward-based learning. Data from studies of Broca's aphasia, Parkinson's disease, hypoxia, focal brain damage, and a genetically transmitted brain anomaly (the putative "language gene," family KE), and from comparative studies of the brains and behavior of other species, demonstrate that the basal ganglia sequence the discrete elements that constitute a complete motor act, syntactic process, or thought process. Imaging studies of intact human subjects and electrophysiologic and tracer studies of the brains and behavior of other species confirm these findings. As Dobzansky put it, "Nothing in biology makes sense except in the light of evolution" (cited in Mayr, 1982). That applies with as much force to the human brain and the neural bases of language as it does to the human foot or jaw. The converse follows: the mark of evolution on the brains of human beings and other species provides insight into the evolution of the brain bases of human language. The neural substrate that regulated motor control in the common ancestor of apes and humans most likely was modified to enhance cognitive and linguistic ability. Speech communication played a central role in this process. However, the process that ultimately resulted in the human brain may have started when our earliest hominid ancestors began to walk.

Anatomy, Comparative↗

Illness, injury, and disability among Shiwiar forager-horticulturalists: implications of health-risk buffering for the evolution of human life history.

Human life history is distinguished by long lifespan, delayed reproduction, intergenerational asymmetric benefit transfers from adults to juveniles and between adults, and a large brain able to engage in unprecedented levels of learning, reasoning, and insight. The evolution of these traits depends on relatively low human mortality. Understanding why humans have low mortality is therefore critical for understanding the evolution of key human traits. One explanation is that the evolution of food provisioning during periods of health crisis reduced mortality. This hypothesis turns on health risk having posed a significant adaptive problem that could be effectively buffered by healthcare provisioning. Unfortunately, the frequency, duration, and fitness effects of temporary disability are difficult to estimate based on osteological evidence alone, and systematic ethno-biological research on these issues among extant small-scale societies with little access to Western medical care is lacking. Here I present data on 678 injuries and illnesses suffered by 40 Shiwiar forager-horticulturalists, based on physical evidence and informant reports. A subsample of 17 individuals provided data on incidence and duration of disability for 215 pathological incidents. Results indicate that injury and illness occur frequently across the lifespan. Most living individuals have suffered temporarily disabling health crises likely to have been lethal without provisioning. The fitness effects of surviving these episodes are high, suggesting that the Shiwiar population structure and lifeway are dependent on infrequent extended provisioning to temporarily disabled individuals, and that provisioning of aid during healthcare crises effectively lowers mortality in this small-scale society.

Adolescent↗

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.

Animals↗

Modern human origins in Australasia: replacement or evolution?

The controversies surrounding the origins of modern humans have spawned two competing hypotheses, namely Replacement and Multiregional Evolution. The first suggests that modern Homo sapiens evolved first in Africa, as late as 140 ka, and subsequently inhabited the balance of the Old World. Conversely, the second hypothesis posits that modern humans evolved principally from local populations of archaic hominids indigenous to the major regions of the Old World. The hominid mandibular remains (ca. 1 Ma) from Sangiran, central Java, Indonesia, were studied in order to test these hypotheses. Non-metric comparisons were performed between these fossils and aboriginal H. sapiens from Africa and Australia. The Replacement model would be supported by a unique Afro-Australian grouping while Multiregional Evolution would be suggested by a Sangiran-Australasian group which would exclude the modern Africans. These data support the Multiregional Evolution hypothesis in that a plurality (eight) of the seventeen non-metric features link Sangiran to modern Australians, while only three exclusively group the humans from Africa and Australia. These results are suggestive of morphological continuity, which implies the presence of a genetic continuum in Australasia dating back at least one million years.

Animals↗

Correlates of body mass evolution in primates.

Body mass is undoubtedly central to the overall adaptive profile of any organism. Despite this, very little is known of what forces drive evolutionary changes in body mass and, consequently, shape patterns of body mass distribution exhibited by animal radiations. The search for factors that may influence evolutionary processes in general frequently focuses on environmental parameters such as climate change or interspecific competition. With respect to body mass, there is also the suggestion that evolutionary lineages may follow an inherent trend toward increased body mass, known as Cope's rule. The present paper investigates whether overall directional trends of body mass change, or correlations between patterns of body mass evolution and environmental factors have influenced the evolution of body mass in plesiadapiforms and primates. Analyses of the global fossil record of plesiadapiforms and primates suggest that the former did indeed follow an overall trend toward increased body mass compatible with the predictions of Cope's rule. In contrast, neither primates as a whole, nor a number of individual primate radiations (Adapiformes, Omomyiformes, and Anthropoidea), show any indication of overall directional patterns of body mass change. No correlations of primate body mass change with either the latitudinal distribution of fossil species, or with estimates of global temperature trends, were found. There is evidence, however, that direct competition between omomyiforms and adapiforms (the two main primate radiations known from the Paleogene) influenced processes of body mass evolution in omomyiforms.

Analysis of Variance↗

Models of intracellular transport and evolution of the Golgi complex.

We have performed a systematic analysis of models explaining the mechanisms of the intracellular biosecretory transport. The models assessed include not only those based on one mechanism (the dissociation model (and its individual case, the vesicular model), the progression model (and its individual cases, the cisterna maturation/progression and the carrier maturation models), and the lateral diffusion model (and its individual case, the bolus model), but also combined models of transport (the percolating-vesicles model and the synthetic model), including several transport mechanisms. Most of these models are not able to explain recent data on the evolution of genes involved in intracellular transport and Golgi evolution. The carrier maturation model proposing that fusion of the large cargo domain with the distal (closer to the plasmalemma) compartment precedes fission of the domain from the proximal compartment exhibits the best performance in correlation with the available information on evolution of the biosecretory pathway.

Biological Evolution↗

Morphology and histology of chimpanzee primary visual striate cortex indicate that brain reorganization predated brain expansion in early hominid evolution.

Human brain evolution is characterized by an overall increase in brain size, cerebral reorganization, and cerebral lateralization. It is generally understood when brain enlargement occurred during human evolution. However, issues concerning cerebral reorganization and hemispheric lateralization are more difficult to determine from brain endocasts, and they are topics of considerable debate. One region of the cerebral cortex that may represent the earliest evidence for brain reorganization is the primary visual cortex (PVC), or area 17 of Brodmann. In nonhuman primates, this region is larger in volume (demarcated anteriorly by the lunate sulcus), and extends further rostrally than it does in modern humans. In early hominid fossil (Australopithecus) endocasts, this region appears to occupy a smaller area compared to that in nonhuman primates. Some have argued that the brain first underwent size expansion prior to reorganization, while others maintain that reorganization predated brain expansion. To help resolve this question, we provide a description of two male, common chimpanzee (Pan troglodytes) brains, YN77-111 and YN92-115, which clearly display a more posterior lunate sulcal morphology than seen in other chimpanzees. These data show that neurogenetic variability exists in chimpanzees, and that significant differences in organization (e.g., a reduced PVC) can predate brain enlargement. While the human brain has experienced numerous expansion and reorganization events throughout evolution, the data from these two chimpanzees offer significant support for the hypothesis that the neurogenetic basis for brain reorganization was present in our early fossil ancestors (i.e., the australopithecines) prior to brain enlargement.

Animals↗

Evolution of the special senses in primates: past, present, and future.

The present special issue of The Anatomical Record is the result of a symposium entitled Evolution of the Special Senses in Primates. Considered together, the special senses of primates are remarkable because they constitute a singular and definitive suite of mammalian characteristics. Examining their evolution is pivotal for understanding the origin and present-day variation of primate behavior and ecology. Accordingly, the 14 articles assembled here consider the different constraints and opportunities associated with the uptake and use of physical and chemical stimuli. The present issue brings together experts on different primate sensory modalities and stresses events at the sensory periphery, where the organism is exposed to and comes into contact with its environment. Key topics include color vision, the genetics of olfaction, the morphological basis and significance of chemical communication, and the neural organization and scaling of primate sensory systems. The result is a special issue that both reflects our current understanding of primate sensory modalities and challenges certain fundamental assumptions concerning their evolution.

Animals↗

A gene for speed? The evolution and function of alpha-actinin-3.

The alpha-actinins are an ancient family of actin-binding proteins that play structural and regulatory roles in cytoskeletal organisation and muscle contraction. alpha-actinin-3 is the most-highly specialised of the four mammalian alpha-actinins, with its expression restricted largely to fast glycolytic fibres in skeletal muscle. Intriguingly, a significant proportion ( approximately 18%) of the human population is totally deficient in alpha-actinin-3 due to homozygosity for a premature stop codon polymorphism (R577X) in the ACTN3 gene. Recent work in our laboratory has revealed a strong association between R577X genotype and performance in a variety of athletic endeavours. We are currently exploring the function and evolutionary history of the ACTN3 gene and other alpha-actinin family members. The alpha-actinin family provides a fascinating case study in molecular evolution, illustrating phenomena such as functional redundancy in duplicate genes, the evolution of protein function, and the action of natural selection during recent human evolution.

Actinin↗

Floral zygomorphy, the recurring evolution of a successful trait.

The flowers of the primitive angiosperm plants were radially symmetrical (actinomorphic). Flowers with bilateral symmetry (zygomorphic) evolved in several clades independently as an adaptation to specialized methods of pollination and played an important role in the diversification of flowering plants. In the model species Antirrhinum majus (snapdragon), the related genes CYCLOIDEA (CYC) and DICHOTOMA (DICH) are key in the development of this trait. This raises the question of whether they played a role in the evolution of floral bilateral symmetry. To address this, the evolution of CYC in relation to the evolution of zygomorphy is being investigated. Phylogenetic and functional analyses of CYC-like genes are being carried out in groups either closely related to Antirhinum or in families where zygomorphy evolved as an independent event. In addition, the origin of zygomorphy is being studied by comparing the function of CYC-like genes in species with zygomorphic flowers with their function in species with radially symmetrical flowers.

Biological Evolution↗

The structure of photosystem I and evolution of photosynthesis.

Oxygenic photosynthesis is the principal producer of both oxygen and organic matter on earth. The primary step in this process--the conversion of sunlight into chemical energy--is driven by four multi-subunit membrane protein complexes named photosystem I, photosystem II, cytochrome b(6)f complex and F-ATPase. Photosystem I generates the most negative redox potential in nature and thus largely determines the global amount of enthalpy in living systems. The recent structural determination of PSI complexes from cyanobacteria and plants sheds light on the evolutionary forces that shaped oxygenic photosynthesis. The fortuitous formation of our solar system in a space plentiful of elements, our distance from the sun and the long time of uninterrupted evolution enabled the perfection of photosynthesis and the evolution of advanced organisms. The available structural information complements the knowledge gained from genomic and proteomic data to illustrate a more precise scenario for the evolution of life systems on earth.

Biological Evolution↗