Becoming Modern Homo sapiens

Human beings are unusual in many ways but perhaps most strikingly in their unique symbolic form of processing information about the world around them. Although based on a long and essential evolutionary history, the modern human cognitive style is not predicted by that history: it is emergent rather than the product of an incremental process of refinement. Homo sapiens is physically very distinctive and is clearly the result of a significant developmental reorganization with ramifications throughout the skeleton and presumably beyond. It is reasonable to suppose that the neural underpinnings of symbolic thought were acquired in this reorganization. However, the fossil and archeological records indicate that the first anatomically recognizable members of the species substantially predated its first members who behaved in a demonstrably symbolic manner. Thus, while the biological potential for symbolic thinking most likely arose in the morphogenetic event that gave rise to H. sapiens as a distinctive anatomical entity, this new capacity was evidently exaptive, in the sense that it had to await its “discovery” and release through a cultural stimulus. Plausibly, this stimulus was the invention of language. One expression of symbolic reasoning is the adoption of technological change in response to environmental challenges, contrasting with earlier responses that typically involved using existing technologies in new ways. As climates changed at the end of the last Ice Age, the new technophile proclivity was expressed in a shift toward agriculture and sedentary lifestyles: a shift that precipitated a fundamentally new (and potentially self-destructive) relationship with nature. Thus, both of what are arguably the two most radical (and certainly the most fateful) evolutionary innovations in the history of life (symbolic thinking and sedentary lifestyles) were both very recent occurrences, well within the (so far rather short) tenure of H. sapiens.     

Art in global context: An evolutionary/functionalist perspective for the 21<Superscript>st</Superscript> century

Regarding the arts as something peopledo — as behaviors, rather than the residue or artifacts of behavior — makes possible a theoretical grounding about their nature and importance, an endeavor that current anthropology of art has largely abandoned. A reconsideration of the suspect and largely discarded terms “functionalism” and “evolutionism” is presented in light of current evolutionary thinking. It is suggested that a contemporary reformulation of these concepts, illustrated by the author's Darwinian or “adaptationist” perspective on art, supports aims and claims of current anthropology of art, and contributes new focus and direction to its endeavors.     

Enzyme-Driven Speciation: Crystallizing Archaea via Lipid Capture

As the origin(s) of life on Earth remains an open question, detailed characteristics about the “last universal ancestor” (LUA) continue to be obscured. Here we provide arguments that strengthen the bacterial-like nature of the LUA. Our view attempts to recreate the evolution of archaeal lipids, the major components of the distinctive membrane that encapsulates these ancient prokaryotes. We show that (S)- 3-O-geranylgeranylglyceryl phosphate synthase (GGGPS), a TIM-barrel protein that performs the committed step in archaeal lipid synthesis, likely evolved from the duplication and fusion of a (βα)₄ half-barrel ancestor. By comparison to the well-characterized HisA and HisF TIM-barrel proteins, we propose a time line for the invention of this diagnostic archaeal biosynthetic pathway. After excluding the possibility of horizontal gene transfer, we conclude that the evolutionary history of GGGPS mirrors the emergence of Archaea from the LUA. We illustrate aspects of this “lipid capture” model that support its likelihood in recreating key evolutionary events and, as our hypothesis is built on a single initiating event, we suggest that the appearance of GGGPS represents an example of enzyme-driven speciation. Reviewing Editor: Dr. Niles Lehman     

On the Structural Regularity in Nucleobases and Amino Acids and Relationship to the Origin and Evolution of the Genetic Code

To explore how chemical structures of both nucleobases and amino acids may have played a role in shaping the genetic code, numbers of sp² hybrid nitrogen atoms in nucleobases were taken as a determinative measure for empirical stereo-electronic property to analyze the genetic code. Results revealed that amino acid hydropathy correlates strongly with the sp² nitrogen atom numbers in nucleobases rather than with the overall electronic property such as redox potentials of the bases, reflecting that stereo-electronic property of bases may play a role. In the rearranged code, five simple but stereo-structurally distinctive amino acids (Gly, Pro, Val, Thr and Ala) and their codon quartets form a crossed intersection “core”. Secondly, a re-categorization of the amino acids according to their β-carbon stereochemistry, verified by charge density (at β-carbon) calculation, results in five groups of stereo-structurally distinctive amino acids, the group leaders of which are Gly, Pro, Val, Thr and Ala, remarkably overlapping the above “core”. These two lines of independent observations provide empirical arguments for a contention that a seemingly “frozen” “core” could have formed at a certain evolutionary stage. The possible existence of this codon “core” is in conformity with a previous evolutionary model whereby stereochemical interactions may have shaped the code. Moreover, the genetic code listed in UCGA succession together with this codon “core” has recently facilitated an identification of the unprecedented icosikaioctagon symmetry and bi-pyramidal nature of the genetic code.     

Depicting the Tree of Life: the Philosophical and Historical Roots of Evolutionary Tree Diagrams

It is a popularly held view that Darwin was the first author to draw a phylogenetic tree diagram. However, as is the case with most popular beliefs, this one also does not hold true. Firstly, Darwin never called his diagram of common descent a tree. Secondly, even before Darwin, tree diagrams were used by a variety of philosophical, religious, and secular scholars to depict phenomena such as “logical relationships,” “affiliations,” “genealogical descent,” “affinity,” and “historical relatedness” between the elements portrayed on the tree. Moreover, historically, tree diagrams themselves can be grouped into a larger class of diagrams that were drawn to depict natural and/or divine order in the world. In this paper, we trace the historical roots and cultural meanings of these tree diagrams. It will be demonstrated that tree diagrams as we know them are the outgrowth of ancient philosophical attempts to find the “true order” of the world, and to map the world “as it is” (ontologically), according to its true essence. This philosophical idea would begin a fascinating journey throughout Western European history. It lies at the foundation of the famous “scala naturae,” as well as religious and secular genealogical thinking, especially in regard to divine, familial (kinship), and linguistic pedigrees that were often depicted by tree images. These scala naturae would fuse with genealogical, pedigree thinking, and the trees that were the result of this blend would, from the nineteenth century onward, also include the element of time. The recognition of time would eventually lead to the recognition of evolution as a fact of nature, and subsequently, tree iconographies would come to represent exclusively the evolutionary descent of species.     

Transforming Our Thinking about Transitional Forms

A common misconception of evolutionary biology is that it involves a search for “missing links” in the history of life. Relying on this misconception, antievolutionists present the supposed absence of transitional forms from the fossil record as evidence against evolution. Students of biology need to understand that evolution is a branching process, paleontologists do not expect to find “missing links,” and evolutionary research uses independent lines of evidence to test hypotheses and make conclusions about the history of life. Teachers can facilitate such learning by incorporating cladistics and tree-thinking into the curriculum and using evograms to focus on important evolutionary transitions.     

Engineering aspects of biological ion channels—from biosensors to computational models for permeation

Molecular sequencing has helped resolve the phylogenetic relationships amongst the diverse groups of algal, fungal-like and protist organisms that constitute the Chromalveolate “superkingdom” clade. It is thought that the whole clade evolved from a photosynthetic ancestor and that there have been at least three independent plastid losses during their evolutionary history. The fungal-like oomycetes and hyphochytrids, together with the marine flagellates Pirsonia and Developayella, form part of the clade defined by Cavalier-Smith and Chao (2006) as the phylum “Pseudofungi”, which is a sister to the photosynthetic chromistan algae (phylum Ochrophyta). Within the oomycetes, a number of predominantly marine holocarpic genera appear to diverge before the main “saprolegnian” and “peronosporalean” lines, into which all oomycetes had been traditionally placed. It is now clear that oomycetes have their evolutionary roots in the sea. The earliest diverging oomycete genera so far documented, Eurychasma and Haptoglossa, are both obligate parasites that show a high degree of complexity and sophistication in their host parasite interactions and infection structures. Key morphological and cytological features of the oomycetes will be reviewed in the context of our revised understanding of their likely phylogeny. Recent genomic studies have revealed a number of intriguing similarities in host–pathogen interactions between the oomycetes with their distant apicocomplexan cousins. Therefore, the earlier view that oomycetes evolved from the largely saprotrophic “saprolegnian line” is not supported and current evidence shows these organisms evolved from simple holocarpic marine parasites. Both the hyphal-like pattern of growth and the acquisition of oogamous sexual reproduction probably developed largely after the migration of these organisms from the sea to land.     

Ecological dominance and the final sprint in hominid evolution

In contrast to many other models of human evolution the “balance of power” theory of Alexander has a clear answer to the question why a runaway selection process for unique social and moral capacities occurred in our ancestry only and not in other species: “ecological dominance” is hypothesized to have diminished the effects of “extrinsic” forces of natural selection such that withinspecies, intergroup competition increased (Alexander, 1989). Alexander seems to be wrong, however, in his claim that already the common HUCHIBO (Humans, Chimps, Bonobo's)-ancestor has crossed the ecological dominance barrier. In this paper an adapted version of Alexander's model is presented and several different ways are proposed to make this adapted version testable. A preliminary survey of the available paleontological and paleoecological data suggests that there is some evidence of a less vulnerable position towards predators in earlyHomo and that there are clear signs related to a crossing of the ecological dominance barrier inHomo sapiens sapiens.     

“Neutral theory” and the dynamics of the evolution of “Modern” human morphology

There is a widespread assumption, even among those who reject the Synthetic Theory of Evolution, that the form of “modern”Homo sapiens is somehow superior to that of archaicHomo sapiens (Tattersall 2000). Those who accept the general outlook of evolutionary biology also tend to assume that “modern” form emerged because it was selected for, which also implies that it was better than that which preceded it. However, after years of using craniofacial measurements to compare human populations, I finally came to realize that, with only a few exceptions, the dimensions measured have no relation to differences in adaptation (Brace 1989, 1996, 2000; Brace et al., 1993). Elsewhere the conclusion has been supported that what is shown by craniometrics is selectively neutral on the average (Relethford 2002). With the documentation that approximately 95% of the genome is not functional, molecular genetics has proved to be useful in documenting the length of time of separation of related human populations by investigating the differences that have accumulated in the neutral parts of the genome. Not surprisingly, the picture revealed by the study of genetic differences is very similar to the one revealed by the study of craniometric differences (Brace et al., 2001). For this reason, the logic behind the “neutral theory” in molecular genetics is very similar to that applied to what happens to morphological characteristics when selection ceases (Brace 1963; Kimura 1968). The difference is that random changes in the neutral part of the genome have no other consequences. However, random changes in the genes that produce specific aspects of morphology will be visible even when selection is no longer controlling the particular trait in question. From an assessment of what random changes in genes controlling morphological traits are likely to do, it follows that the most likely change will probably be a reduction in the trait in question, i.e. the Probable Mutation Effect will produce structural reduction. When survival in the temperate zone during the last glaciation dependend on “obligatory cooking”, one of the unintended consequences was a reduction in the selective pressures maintaining a Middle Pleistocene-sized dentition. The result was a gradual reduction in tooth size and a conversion, of a Neanderthal-sized face into one of “modern” dimensions. The manufacture and use of string for snares and nets similarly reduced the selective pressures maintaining post-cranial levels of robustness and muscularity. The reduction in the latter resulted in the emergence of moderm post-cranial robustness out of what had been a Neanderthal level,in situ wherever the technology can be documented and without any need for invasions and replacements.     

Polymorphism of canonical and noncanonical <Emphasis Type="Italic">gypsy</Emphasis> sequences in different species of <Emphasis Type="Italic">Drosophila </Emphasis><Emphasis Type="Italic">melanogaster</Emphasis> subgroup: possible evolutionary relations

Mobile genetic elements constitute a substantial part of eukaryotic genome and play an important role in its organization and functioning. Co-evolution of retrotransposons and their hosts resulted in the establishment of control systems employing mechanisms of RNA interference that seem to be impossible to evade. However, “active” copies of endogenous retrovirus gypsy escape cellular control in some cases, while its evolutionary elder “inactive” variants do not. To clarify the evolutionary relationship between “active” and “inactive” gypsy we combined two approaches: the analysis of gypsy sequences, isolated from G32 Drosophila melanogaster strain and from different Drosophila species of the melanogaster subgroup, as well as the study of databases, available on the Internet. No signs of “intermediate” (between “active” and “inactive”) gypsy form were found in GenBank, and four full-size G32 gypsy copies demonstrated a convergence that presumably involves gene conversion. No “active” gypsy were revealed among PCR generated gypsy ORF3 sequences from the various Drosophila species indicating that “active” gypsy appeared in some population of D. melanogaster and then started to spread out. Analysis of sequences flanking gypsy variants in G32 revealed their predominantly heterochromatic location. Discrepancy between the structure of actual gypsy sites in G32 and corresponding sequences in database might indicate significant inter-strain heterochromatin diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Palatine fenestrae,the orangutan and hominoid evolution

Although most mammals develop relatively large double anterior palatine fenestrae that patently communicate with the nasal cavity, four extant primates—Homo sapiens, Pongo, Pan andGorilla—do not. While these four have closed-down these foramenal structures,Homo sapiens andPongo are unique in forming a single foramen palatally. Among fossil taxa,Homo, Australopithecus, Sivapithecus (=Ramapithecus) andRudapithecus also develop a single foramen palatally. Dryopithecines, the presumed fossil apes, preserve the two patent fenestrae. In light of dental features that are considered diagnostically “hominid,” which are also found in the orangutan, it is suggested that this “ape,” rather thanPan, is phylogenetically closer toHomo.     

“Force-Talk” in Evolutionary Explanation: Metaphors and Misconceptions

The notion of “pressure” as an evolutionary “force” that “causes” evolution is a pervasive linguistic feature of biology textbooks, journal articles, and student explanatory discourse. We investigated the consequences of using a textbook and curriculum that incorporate so-called force-talk. We examined the frequency with which biology majors spontaneously used notions of evolutionary “pressures” in their explanations, students’ definitions and explanations of what they meant when they used pressures, and the structure of explanatory models that incorporated evolutionary pressures and forces. We found that 12–20 percent of undergraduates spontaneously used “pressures” and/or “forces” as explanatory factors but significantly more often in trait gain scenarios than in trait loss scenarios. The majority of explanations using “force-talk” were characterized by faulty evolutionary reasoning. We discuss the conceptual similarity between faulty notions of evolutionary pressures and linguists’ force-dynamic models of everyday reasoning and ultimately question the appropriateness of force-talk in evolution education.     

Are Cirripedia hopeful monsters? Cytogenetic approach and evidence for a Hox gene cluster in the cirripede crustacean Sacculina carcini

The “hopeful monster” has haunted evolutionary thinking since Richard Goldschmidt coined the phrase in 1933. The phrase is directly related to genetic mechanisms in development and evolution. Cirripedes are peculiar crustaceans in that they all lack abdomens as adults. In a previous study aimed at describing the repertoire of Hox genes of the Cirripedia, we failed to isolate the abdominal-A gene in three species representative of all three cirripede orders. To address the question of whether the cirripede ancestor could have been a “hopeful monster” arising from a rearrangement of the Hox complex, we have performed a cytogenetic analysis of the Hox complex of the cirripede Sacculina carcini. We present here molecular and cytogenetic evidence for the grouping of the Hox genes on a single chromosome. This is the first direct evidence reported for the grouping of Hox genes on the same chromosome in a non-insect arthropod species.     

The evolution of “egalitarian” and “despotic” social systems among macaques

Recent studies of captive macaques have revealed considerable inter-species differences in dominance styles among females. In “egalitarian” species such as stumptail (Macaca arctoides) or tonkean macaques (M. tonkeana), social interactions are more symmetrical and less kin-biased than in “despotic” species such as Japanese (M. fuscata) or rhesus macaques (M. mulatta). Field observations of moor macaques (M. maurus), close relatives of tonkean macaques, suggest that tolerance during feeding characterizes their egalitarian dominance style in the natural habitat. Although it has been proposed that communal defense against other groups may be the main selective force in the evolution of egalitarian dominance style among females, few field data support this prediction. A game theory analysis showed that both an “egalitarian” strategy and a “despotic” strategy are possible evolutionarily stable strategies (ESS) under certain conditions. The difference in dominance styles might reflect the difference in ESS. This means that an egalitarian dominance style can emerge without strong between-group contest competition. A phylogenetic comparison among macaques suggests that despotic dominance styles very likely evolved from egalitarian dominance styles. In the future, primate socioecological studies should pay more attention to the evolutionary history of each species.     

Texas Biology and Biological Anthropology Faculty Express Their Views on Teaching Evolution

Unsurprisingly, survey results indicate that Texas biology and biological anthropology faculty with expertise in an evolutionary area strongly support teaching “just evolution” (100%; N = 54) and not creationism/intelligent design. Importantly, they do not think that religious faith is incompatible with acceptance of evolutionary biology (91%; N = 55), even though 50% (N = 52) describe themselves as “not at all religious.” As school boards nationwide debate science standards, it is important that faculty with relevant expertise have a voice. Biological anthropologists should not be overlooked as a public resource in these debates.     

Improving How Evolution Is Taught: Facilitating a Shift from Memorization to Evolutionary Thinking

In North America, public understanding and acceptance of evolution is alarmingly low. Moreover, acceptance rates are declining, and studies suggest that even students who have taken courses in evolution have the same misunderstandings as the general public. These data signal deficiencies in our educational system and provide a “call to arms” to improve how evolution is taught. Many studies show that student education can be improved by replacing lecture-based pedagogy with active learning approaches—where the role of students changes from passive note taking to active problem solving. Here, we describe changes made to a second-year undergraduate evolution course to facilitate a shift to active learning and improve student understanding of evolution. First, lectures were used only sparingly and were largely replaced by problem-solving activities. Second, standard textbooks were replaced by “popular” books applying evolutionary thinking to topics students encounter on a daily basis. Lastly, predefined laboratory exercises were replaced by student-designed and implemented research projects. These changes led to increased student engagement and enjoyment, improved understanding of evolution and ability to apply evolutionary thinking to biological problems, and increased student recognition that evolutionary thinking is important not only in the classroom but also in their daily lives.     

Microsatellite DNA analysis of rainbow trout (Oncorhynchus mykiss) from western Alberta, Canada: native status and evolutionary distinctiveness of “Athabasca” rainbow trout

Molecular genetic assays can contribute to conservation of aquatic taxa by assessing evolutionary and taxonomic distinctiveness, levels of genetic variation within and between populations, and the degree of introgression with introduced taxa. The Athabasca River drainage of␣western Alberta, Canada is one of only three (and the largest) drainages flowing east of the continental divide that contain native populations of rainbow trout (Salmonidae: Oncorhynchus mykiss). The “Athabasca” rainbow trout has been considered a preglacial relict worthy of special conservation measures. In addition, the native range of Athabasca rainbow trout has seen many instances of introductions of non-native populations since the beginning of the 20th century. We assayed rainbow trout from the Athabasca River drainage, from hatchery populations, and from representative populations in adjacent regions (N = 49 localities) for variation at 10 microsatelite loci to assess the level of evolutionary distinctiveness of Athabasca rainbow trout, and to assess the levels of introgression with non-native hatchery fish. We found that native Athabasca rainbow trout did not form a distinctive genetic assemblage and that the greatest amount of allele frequency variation was attributable to contemporary drainage systems (29.3%) rather than by a Athabasca/non-Athabasca distinction (12.6%). We found that 78% of all fish were confidently assigned to a “wild” rather than a “hatchery” genetic grouping and that most of the inferred introgression with hatchery fish was restricted to a few localities (N = 6). Our results suggest that: (i)␣Athabasca River rainbow trout are likely postglacial immigrants from adjacent populations of the Fraser River, and (ii) that there is no evidence of widespread introgression of hatchery alleles into native Athabasca River drainage rainbow trout.     

Heterogeneity of heterochromatin in six species ofCtenomys (Rodentia: Octodontoidea: Ctenomyidae) from Argentina revealed by a combined analysis of C- and RE-banding

Exceptional chromosomal variability makesCtenomys an excellent model for evolutionary cytogenetic analysis. Six species belonging to three evolutionary lineages were studied by means of restriction endonuclease and C-chromosome banding. The resulting banding patterns were used for comparative analysis of heterochromatin distribution on chromosomes. This combined analysis allowed intra- and inter-specific heterochromatin variability to be detected, groups of species belonging to different lineages to be characterized, and phylogenetic relationships hypothesized from other data to be supported. The “ancestral group”,Ctenomys pundti andC. talarum, share three types of heterochromatin, the most abundant of which was also found in C. aff.C. opimus, suggesting that the latter species also belongs to the “ancestral group”. Additionally, within the subspeciesC. t. talarum, putative chromosomal rearrangements distinguishing two of the three chromosomal races were identified. Two species belong to an “eastern lineage”,C. osvaldoreigi andC. rosendopascuali, and share only one type of heterochromatin homogeneously distributed across their karyotypes.C. latro, the only analyzed species from the “chacoan” lineage, showed three types of heterochromatin, one of them being that which characterizes the “eastern lineage”.C. aff.C. opimus, because of its low heterochromatin content, is the most primitive karyotype of the genus yet described. The heterochromatin variability showed by these species, reflecting the evolutionary divergence toward different heterochromatin types, may have diverged since the origin of the genus. Heterochromatin amplification is proposed as a trend withinCtenomys, occurring independently of chromosomal change in diploid numbers.     

Progressive changes in brain size and musculo-skeletal traits in seven hominoid populations

Neurological complexity has increased over evolutionary time for invertebrates and vertebrates alike, with the hominid brain tripling in size over the last 3 million years. Since magnetic resonance imaging (MRI) studies among humans indicate a significant correlation (meanr>0.40) between individual differences in brain size and general cognitive ability, it is reasonable to hypothesize that increasing brain size confers greater intelligence. However, larger brains have associated costs, taking longer to build and requiring more energy to run. Sufficient advantages must have accrued for them to override these trade-offs. The present paper documents that in hominoids, as brain size increased from 380 to 1364 cm³ over seven hominoid groups (chimpanzees to australopithecines toHomo habilis toHomo erectus to differences amongHomo sapiens), it was accompanied by changes in 74 musculo-skeletal traits (rs=0.90). These occurred on both cranial traits (temporalis fossae, post-orbital constrictions, mandibles, dentition, nuchal muscle attachments) and on post-cranial traits (pelvic widths, femoral heads, tibial plateaus). It is concluded that in the evolutionary competition to find and fill new niches, there was “room at the top” for greater behavioral complexity and larger brain size, leading to cascading effects on other traits.