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A Dryopithecus skeleton and the origins of great-ape locomotion.

The evolution of skeletal adaptations to orthograde postures, characteristic of extant hominoids, is of great interest as it provides the key to understanding the origins of apes and humans. We report here the recent discovery of an extraordinary partial skeleton of Dryopithecus laietanus from Can Llobateres (Spain). It provides evidence that orthograde postures and locomotion appeared at least 9.5 million years ago. Our results indicate that the body structure of this Miocene ape closely resembles that of extant hominoids and differs from the pronograde pattern of Miocene proconsulids in a set of important morphological characters. Dryopithecus also shows more traits reflecting structural adaptations for suspension than occurs in African apes. A similar positional behaviour is inferred for Sivapithecus indicus, thus strengthening previous hypotheses linking both Miocene forms with Pongo.

Animals↗

The genetical history of humans and the great apes.

When and where did modern humans evolve? How did our ancestors spread over the world? Traditionally, answers to questions such as these have been sought in historical, archaeological, and fossil records. However, increasingly genetic data provide information about the evolution of our species. In this review, we focus on the comparison of the variation in the human gene pool to that of our closest evolutionary relatives, the great apes, because this provides a relevant perspective on human genetical evolution. For instance, comparisons to the great apes show that humans are unique in having little genetic variation as well as little genetic structure in their gene pool. Furthermore, genetic data indicate that humans, but not the great apes, have experienced a period of dramatic growth in their early history.

Animals↗

Evolutionary tree for apes and humans based on cleavage maps of mitochondrial DNA.

The high rate of evolution of mitochondrial DNA makes this molecule suitable for genealogical research on such closely related species as humans and apes. Because previous approaches failed to establish the branching order of the lineages leading to humans, gorillas, and chimpanzees, we compared human mitochondrial DNA to mitochondrial DNA from five species of ape (common chimpanzee, pygmy chimpanzee, gorilla, orangutan, and gibbon). About 50 restriction endonuclease cleavage sites were mapped in each mitochondrial DNA, and the six maps were aligned with respect to 11 invariant positions. Differences among the maps were evident at 121 positions. Both conserved and variable sites are widely dispersed in the mitochondrial genome. Besides site differences, ascribed to point mutations, there is evidence for one rearrangement: the gorilla map is shorter than the other owing to the deletion of 95 base pairs near the origin of replication. The parsimony method of deriving all six maps from a common ancestor produced a genealogical tree in which the common and pygmy chimpanzee maps are the most closely related pair; the closest relative of this pair is the gorilla map; most closely related to this trio is the human map. This tree is only slightly more parsimonious than some alternative trees. Although this study has given a magnified view of the genetic differences among humans and apes, the possibility of a three-way split among the lineages leading to humans, gorillas, and chimpanzees still deserves serious consideration.

Animals↗

Reconstruction of genomic rearrangements in great apes and gibbons by chromosome painting.

The homology between hylobatid chromosomes and other primates has long remained elusive. We used chromosomal in situ suppression hybridization of all human chromosome-specific DNA libraries to "paint" the chromosomes of primates and establish homologies between the human, great ape (chimpanzee, gorilla, and orangutan), and gibbon karyotypes (Hylobates lar species group, 2n = 44). The hybridization patterns unequivocally demonstrate the high degree of chromosomal homology and synteny of great ape and human chromosomes. Relative to human, no translocations were detected in great apes, except for the well-known fusion-origin of human chromosome 2 and a 5;17 translocation in the gorilla. In contrast, numerous translocations were detected that have led to the massive reorganization of the gibbon karyotype: the 22 autosomal human chromosomes have been divided into 51 elements to compose the 21 gibbon autosomes. Molecular cytogenetics promises to finally allow hylobatids to be integrated into the overall picture of chromosomal evolution in the primates.

Animals↗

Emergence of the keratinocyte growth factor multigene family during the great ape radiation.

The structural gene for human keratinocyte growth factor (KGF), a member of the fibroblast growth factor family, consists of three coding exons and two introns typical of other fibroblast growth factor loci. A portion of the KGF gene, located on chromosome 15, is amplified to approximately 16 copies in the human genome, and these highly related copies (which consist of exon 2, exon 3, the intron between them, and a 3' noncoding segment of the KGF transcript) are dispersed to multiple human chromosomes. The KGF-like sequences are transcriptionally active, differentially regulated in various tissues, and composed of three distinct classes of coding sequences that are 5% divergent from each other and from the authentic KGF sequence. Multiple copies of KGF-like genes were also discovered in the genomic DNAs of chimpanzee and gorilla but were not found in lesser apes (gibbon), Old World monkeys (African green monkey and macaques), mice, or chickens. The pattern of evolutionary occurrence suggests that a primordial KGF gene was amplified and chromosomally dispersed subsequent to the divergence of orangutan from African apes but before the trichotomous divergence of human, chimpanzee, and gorilla 5-8 million years ago. The appearance of a transcriptionally active and chromosomally dispersed multigene KGF family may have implications in the evolution of the great apes and humans.

Animals↗

Evidence of hominid-like precision grip capability in the hand of the Miocene ape Oreopithecus.

Functional and allometric analyses of the hand of the late Miocene ape Oreopithecus bambolii (Tuscany, Italy) reveal a series of features that reflect an improved grasping capability including firm pad-to-pad precision gripping that apes are unable to perform. Related features such as hand length, relative thumb length, a deep and large insertion area for the tendon of the long thumb flexor, and the form of the metacarpal 2/capitate articulation are not present in extant or fossil apes. In these features, the Oreopithecus hand closely matches the pattern of early hominids, presumably as a response to similar functional demands.

Animals↗

CXCR4 homologues of gibbon ape, African green monkey, squirrel monkey, and cotton-top marmoset.

CXCR4 gene homologues were isolated from an ape (gibbon), an Old World monkey (African green monkey), and two New World monkeys (squirrel monkey and cotton-top marmoset), and their DNA sequences determined. The squirrel monkey and cotton-top marmoset CXCR4 sequences more closely resemble homologues from apes than Old World monkeys, a pattern not seen for the related chemokine receptor CCR5. The African green monkey CXCR4 gene is similar to its homologue in baboon, a pattern that has also been seen among CCR5 homologues. The gibbon CXCR4 contains the first polymorphisms recognized in ape homologues, the human and chimpanzee CXCR4 proteins being identical, and two of these three differences are also observed in one or more Old World monkey homologues. While 18 positions within CXCR4 are now known to be polymorphic in primates, 7 of these polymorphisms have been observed in multiple examples and 11 have been observed only once.

Amino Acid Sequence↗

Human and ape molecular clocks and constraints on paleontological hypotheses.

Although the relationships of the living hominoid primates (humans and apes) are well known, the relationships of the fossil species, times of divergence of both living and fossil species, and the biogeographic history of hominoids are not well established. Divergence times of living species, estimated from molecular clocks, have the potential to constrain hypotheses of the relationships of fossil species. In this study, new DNA sequences from nine protein-coding nuclear genes in great apes are added to existing datasets to increase the precision of molecular time estimates bearing on the evolutionary history of apes and humans. The divergence of Old World monkeys and hominoids at the Oligocene-Miocene boundary (approximately 23 million years ago) provides the best primate calibration point and yields a time and 95% confidence interval of 5.4 +/- 1.1 million years ago (36 nuclear genes) for the human-chimpanzee divergence. Older splitting events are estimated as 6.4 +/- 1.5 million years ago (gorilla, 31 genes), 11.3 +/- 1.3 million years ago (orangutan, 33 genes), and 14.9 +/- 2.0 million years ago (gibbon, 27 genes). Based on these molecular constraints, we find that several proposed phylogenies of fossil hominoid taxa are unlikely to be correct.

Animals↗

Modern African ape populations as genetic and demographic models of the last common ancestor of humans, chimpanzees, and gorillas.

In order to fully understand human evolutionary history through the use of molecular data, it is essential to include our closest relatives as a comparison. We provide here estimates of nucleotide diversity and effective population size of modern African ape species using data from several independent noncoding nuclear loci, and use these estimates to make predictions about the nature of the ancestral population that eventually gave rise to the living species of African apes, including humans. Chimpanzees, bonobos, and gorillas possess two to three times more nucleotide diversity than modern humans. We hypothesize that the last common ancestor (LCA) of these species had an effective population size more similar to modern apes than modern humans. In addition, estimated dates for the divergence of the Homo, Pan, and Gorilla lineages suggest that the LCA may have had stronger geographic structuring to its mtDNA than its nuclear DNA, perhaps indicative of strong female philopatry or a dispersal system analogous to gorillas, where females disperse only short distances from their natal group. Synthesizing different classes of data, and the inferences drawn from them, allows us to predict some of the genetic and demographic properties of the LCA of humans, chimpanzees, and gorillas.

Africa↗

Evolution of bitter taste receptors in humans and apes.

Bitter taste perception is crucial for the survival of organisms because it enables them to avoid the ingestion of potentially harmful substances. Bitter taste receptors are encoded by a gene family that in humans has been shown to contain 25 putatively functional genes and 8 pseudogenes and in mouse 33 putatively functional genes and 3 pseudogenes. Lineage-specific expansions of bitter taste receptors have taken place in both mouse and human, but very little is known about the evolution of these receptors in primates. We report the analysis of the almost complete repertoires of bitter taste receptor genes in human, great apes, and two Old World monkeys. As a group, these genes seem to be under little selective constraint compared with olfactory receptors and other genes in the studied species. However, in contrast to the olfactory receptor gene repertoire, where humans have a higher proportion of pseudogenes than apes, there is no evidence that the rate of loss of bitter taste receptor genes varies among humans and apes.

Animals↗

A young Alu subfamily amplified independently in human and African great apes lineages.

A variety of Alu subfamilies amplified in primate genomes at different evolutionary time periods. Alu Sb2 belongs to a group of young subfamilies with a characteristic two-nucleotide deletion at positions 65/66. It consists of repeats having a 7-nucleotide duplication of a sequence segment involving positions 246 through 252. The presence of Sb2 inserts was examined in five genomic loci in 120 human DNA samples as well as in DNAs of higher primates. The lack of the insertional polymorphism seen at four human loci and the absence of orthologous inserts in apes indicated that the examined repeats retroposed early in the human lineage, but following the divergence of great apes. On the other hand, similar analysis of the fifth locus (butyrylcholinesterase gene) suggested contemporary retropositional activity of this subfamily. By a semi-quantitative PCR, using a primer pair specific for Sb2 repeats, we estimated their copy number at about 1500 per human haploid genome; the corresponding numbers in chimpanzee and gorilla were two orders of magnitude lower, while in orangutan and gibbon the presence of Sb2 Alu was hardly detectable. Sequence analysis of PCR-amplified Sb2 repeats from human and African great apes is consistent with the model in which the founding of Sb2 subfamily variants occurred independently in chimpanzee, gorilla and human lineages.

Animals↗

Large-scale variation among human and great ape genomes determined by array comparative genomic hybridization.

Large-scale genomic rearrangements are a major force of evolutionary change and the ascertainment of such events between the human and great ape genomes is fundamental to a complete understanding of the genetic history and evolution of our species. Here, we present the results of an evolutionary analysis utilizing array comparative genomic hybridization (array CGH), measuring copy-number gains and losses among these species. Using an array of 2460 human bacterial artificial chromosomes (BACs) (12% of the genome), we identified a total of 63 sites of putative DNA copy-number variation between humans and the great apes (chimpanzee, bonobo, gorilla, and orangutan). Detailed molecular characterization of a subset of these sites confirmed rearrangements ranging from 40 to at least 175 kb in size. Surprisingly, the majority of variant sites differentiating great ape and human genomes were found within interstitial euchromatin. These data suggest that such large-scale events are not restricted solely to subtelomeric or pericentromeric regions, but also occur within genic regions. In addition, 5/9 of the verified variant sites localized to areas of intrachromosomal segmental duplication within the human genome. On the basis of the frequency of duplication in humans, this represents a 14-fold positional bias. In contrast to previous cytogenetic and comparative mapping studies, these results indicate extensive local repatterning of hominoid chromosomes in euchromatic regions through a duplication-driven mechanism of genome evolution.

Animals↗

Morphological analysis of the hindlimb in apes and humans. I. Muscle architecture.

We present quantitative data on the hindlimb musculature of Pan paniscus, Gorilla gorilla gorilla, Gorilla gorilla graueri, Pongo pygmaeus abelii and Hylobates lar and discuss the findings in relation to the locomotor habits of each. Muscle mass and fascicle length data were obtained for all major hindlimb muscles. Physiological cross-sectional area (PCSA) was estimated. Data were normalized assuming geometric similarity to allow for comparison of animals of different size/species. Muscle mass scaled closely to (body mass)(1.0) and fascicle length scaled closely to (body mass)(0.3) in most species. However, human hindlimb muscles were heavy and had short fascicles per unit body mass when compared with non-human apes. Gibbon hindlimb anatomy shared some features with human hindlimbs that were not observed in the non-human great apes: limb circumferences tapered from proximal-to-distal, fascicle lengths were short per unit body mass and tendons were relatively long. Non-human great ape hindlimb muscles were, by contrast, characterized by long fascicles arranged in parallel, with little/no tendon of insertion. Such an arrangement of muscle architecture would be useful for locomotion in a three dimensionally complex arboreal environment.

Animals↗

5'-terminal nucleotide sequences of the Rauscher leukemia virus and gibbon ape leukemia virus genomes exhibit a high degree of correspondence.

The 5'-terminal regions of gibbon ape leukemia virus-Hall's Island and Rauscher murine leukemia virus have been completely sequenced. The chain length for the 5'-terminal region of Rauscher murine leukemia virus is 140 nucleotides, and that for gibbon ape leukemia virus-Hall's Island is 144 nucleotides. An alignment of the sequences maximizing the number of ocrrespondences with the minimum introduction of gaps shows 81% nucleotide matches. From the complementary RNA, secondary structures of this region have been proposed. These data demonstrate the conservation of the 5'-terminal genetic sequences of these viruses and strongly reinforce the concept that viruses of murine origin and viruses of the gibbon ape leukemia virus-Simian sarcoma-associated virus group are closely related.

Animals↗

Cold stress in captive great apes recorded in incremental lines of dental cementum.

Incremental lines in dental cementum of museum specimens of 11 free-ranging great apes were compared to the respective structures in 5 captive specimens of known age-at-death, and with many known life-history parameters. While the dental cementum of the free-ranging apes was regularly structured into alternating dark and light bands, 4 out of 5 captive animals showed marked irregularities in terms of hypomineralized bands which could all be dated to the year 1963. Cementum preservation was insufficient in the fifth specimen and did not permit such a differentiation. All 4 captive apes had been kept in a zoo located in the northern hemisphere, where 1963 was characterized by an extremely cold winter. Since cold stress is a calcium-consuming process, the lack of available calcium in newly forming cementum could be responsible for the observed hypomineralization. The appositional growth characteristics of dental cementum serve as a record for such life-history events.

Adaptation, Physiological↗

Comparative mapping of human claudin-1 (CLDN1) in great apes.

The gene encoding claudin-1 (CLDN1) has been mapped to human chromosome 3 (HSA3; 3q28-->q29) using a radiation hybrid panel. Employing fluorescence in situ hybridization (FISH) we here show that a human P1-derived artificial chromosome (PAC) containing CLDN1 detects the orthologous sites in chromosomes of the great apes, chimpanzee, gorilla, and orangutan. Furthermore, the chromosomal position of CLDN1 was determined in mouse chromosomes by FISH. The position of fluorescent signals is confined to a single chromosomal site in both great apes and mouse and in each case maps to the chromosomal region that has conserved synteny with HSA3 (PTR2q28, GGO2q28, PPY2q38 and MMU16B1). Using a gene-specific probe our results are consistent with reports of the striking similarity of great ape and human genomes as illustrated previously by chromosome painting.

Animals↗

APE-type non-LTR retrotransposons: determinants involved in target site recognition.

Non-long terminal repeat (Non-LTR) retrotransposons represent a diverse and widely distributed group of transposable elements and an almost ubiquitous component of eukaryotic genomes that has a major impact on evolution. Their copy number can range from a few to several million and they often make up a significant fraction of the genomes. The members of the dominating subtype of non-LTR retrotransposons code for an endonuclease with homology to apurinic/apyrimidinic endonucleases (APE), and are thus termed APE-type non-LTR retrotransposons. In the last decade both the number of identified non-LTR retrotransposons and our knowledge of biology and evolution of APE-type non-LTR retrotransposons has increased tremendously.

Retroelements↗

The similarity of chorionic gonadotrophin and its subunits in term placentae from man, apes, old and New World monkeys and a prosimian.

Chorionic gonadotrophin (CG) was estimated, by bioassay and radioimmunoassay (RIA), in placental extracts from 11 ape and monkey species. There was a significant correlation between the results of the two assay systems (r = 0.903, p less than 0.001). The concentration of CG in most primate term placentae was the same as that in the human placenta at term. Extracts from all placentae cross-reacted with antiserum to ovine LH-beta subunit, and those of the chimpanzee and gorilla also had a significant cross-reaction with an antiserum to the carboxyl terminal peptide of the HCG-beta subunit. Primate placentae chromatographed on Sephadex G-200 had components active in the RIA systems for HCG, HCG-alpha HCG-beta subunits. In general, the elution profiles of all ape and monkey placental extracts resemble those made from human term placentae and of purified HCG and its subunits. The shape of the elution patterns from human and non-human material suggests that there was more than one molecular form of CG-alpha subunit activity. A second, more retarded molecular form having beta subunit activity was found in extracts made from human, gorilla, gibbon and rhesus monkey placentae. The similarity between the structure of ape and monkey placental CG with HCG and its subunits implies a function similar to that of HCG in late pregnancy.

Animals↗