Genetic and morphological records of the Hominoidea and hominid origins: a synthesis

Molecular genetics has had a major impact on phylogenetics, although many hominoid paleontologists and morphologists ignore or remain unaware of genetic data. However, substantial genetic evidence shows chimpanzees and humans as closest relatives. Living hominoids share many postcranial similarities, many of which are retained from the extant hominoid common ancestor. Miocene hominoid fossils consisted until recently mostly of teeth and jaw fragments which are relatively uninformative phylogenetically. As their postcrania have become better sampled, surprisingly few of these taxa share significant similarities with living apes, suggesting that few if any are related to specific extant lineages. Given the genetically inferred relationships of hominoids and the morphology of the earliest hominids, the common ancestor of humans and chimpanzees was probably chimp-like, a knuckle-walker with small thin-enameled cheek teeth. If correct, this scenario implies that known Miocene hominoids, most of which are postcranially archaic and have large, thickly enameled cheek teeth, throw little if any direct light on hominid origins.     

A gloverin-like antibacterial protein is synthesized in Helicoverpa armigera following bacterial challenge

The elucidation of patterns of cranial skeletal maturation and growth in fossil hominids is possible not only through dental studies but also by mapping different aspects of ossification in both extant African apes and humans. However, knowledge of normal skeletal development in large samples of extant great apes is flimsy. To remedy this situation, this paper offers an extensive survey and thorough discussion of the ossification of the posterior border of the sphenoid greater wing. Indeed, this area provides much information about basicranial skeletal maturation. We investigate three variants: the absence of the foramen spinosum and the position of both the foramen spinosum and the foramen ovale in relation to the sphenosquamosal suture. Providing original data about humans and 1,425 extant great ape skulls and using a sample of 64 fossil hominids, this study aimed to test whether different ossification patterns occurred during the course of human evolution. The incidence of three derived morphologies located on the posterior border of the sphenoid greater wing increases during human evolution at different geological periods. The evolutionary polarity of these three derived morphologies is assessed by outgroup comparison and ontogenetic methods. During human evolution, there is a clear trend for the foramen spinosum to be present and wholly located on the posterior area of the sphenoid greater wing. Moreover, in all the great ape species and in Australopithecus afarensis, the sphenosquamosal suture may split the foramen ovale. Inversely, the foramen ovale always lies wholly within the sphenoid greater wing in Australopithecus africanus, robust australopithecines, early Homo, H. erectus (and/or H. ergaster), and Homo sapiens. From ontogenetic studies in humans, we conclude that, during human evolution, the ossification of the posterior area of the sphenoid greater wing progressively surrounded the middle meningeal artery (passing through the foramen spinosum) and the small meningeal artery (passing through the foramen ovale).     

Associated malformations in infants and fetuses with upper or lower neural tube defects

A number of studies on sexual differences in the ilium have been reported. However, most of these studies have focused on the adult ilium. With regard to sexual differences in the fetal ilium, few studies have been carried out. Especially, there have few studies regarding sexual differences in the fetal ilium using dry bones. In the present study, sexual differences in the morphological characteristics of the greater sciatic notch were investigated using dry fetal iliac bones. We examined 212 fetuses (106 males and 106 females) measuring 20.0 cm or more in fetal length with free of gross malformations or deformities which were collected at Saga Medical School. The iliac bones were excised from these fetuses and dried as materials for study. Fetal length was measured in the fully extended position and was classified into six groups each for males and females in 5-cm increments, giving a total of twelve groups. The inner lateral surface of the greater sciatic notch was magnified 25x using projector, traced, and the tracings were inputted to a computer (7600/120: Apple Co. Ltd) using a flatbed scanner. The width and height of the greater sciatic notch were measured, as well as the area of the greater sciatic notch. No sexual differences were noted regarding the increase in the width of the greater sciatic notch in fetuses up to 39.9 cm in fetal length. In fetuses measuring between 40.0 anf 44.9 cm, a significant sexual difference was observed, with the increase in greater sciatic notch width markedly greater in females. With respect to the increase in the height of the greater sciatic notch, no significant differences were noted. No significant differences were observed in the total area of the greater sciatic notch in fetuses up to 39.9 cm in fetal length. On the other hand, a significant sexual difference was observed in fetuses measuring 40.0 cm or more, with the total area markedly increased in females. In addition, no significant differences were found in the area between the two sides of the greater sciatic notch in fetuses up to 39.9 cm in fetal length, although significant differences were observed in fetuses measuring 40.0 cm or more. In female fetuses 40.0 cm or more in fetal length, the greater sciatic notch was found to be wider than in males, and the peak of the greater sciatic notch was displaced in the anteroinferior direction, farther from the auricular surface of the ilium and nearer the pubic bone. Thus these female fetuses show morphological characteristics of the adult female pelvic bone. Based on the results obtained, it is concluded that sexual differences can be identified in fetuses from the 8th month of pregnancy (fetal length 40.0 cm or more).     

A mutation in human CMP-sialic acid hydroxylase occurred after the Homo-Pan divergence

Sialic acids are important cell-surface molecules of animals in the deuterostome lineage. Although humans do not express easily detectable amounts of N-glycolylneuraminic acid (Neu5Gc, a hydroxylated form of the common sialic acid N-acetylneuraminic acid, Neu5Ac), it is a major component in great ape tissues, except in the brain. This difference correlates with lack of the hydroxylase activity that converts CMP-Neu5Ac to CMP-Neu5Gc. Here we report cloning of human and chimpanzee hydroxylase cDNAs. Although this chimpanzee cDNA is similar to the murine homologue, the human cDNA contains a 92-bp deletion resulting in a frameshift mutation. The isolated human gene also shows evidence for this deletion. Genomic PCR analysis indicates that this deletion does not occur in any of the African great apes. The gene is localized to 6p22-p23 in both humans and great apes, which does not correspond to known chromosomal rearrangements that occurred during hominoid evolution. Thus, the lineage leading to modern humans suffered a mutation sometime after the common ancestor with the chimpanzee and bonobo, potentially affecting recognition by a variety of endogenous and exogenous sialic acid-binding lectins. Also, the expression of Neu5Gc previously reported in human fetuses and tumors as well as the traces detected in some normal adult humans must be mediated by an alternate pathway.     

South Turkwel: a new pliocene hominid site in Kenya

New fossils discovered south of the Turkwel River in northern Kenya include an associated metacarpal, capitate, hamate, lunate, pedal phalanx, mandibular fragment, and teeth. These fossils probably date to around 3.5 m.y.a. Faunal information suggests that the environment at South Turkwel was predominantly bushland. The mandibular and dental remains are fragmentary, but the postcranial fossils are informative. Comparisons with Australopithecus, modern human, chimpanzee and gorilla hand bones suggest that the Turkwel hominid was most like Australopithecus afarensis and A. africanus. Carpometacarpal articulations are intermediate between those of modern humans and African apes, suggesting enhanced gripping capabilities compared with extant apes. The hamulus was strikingly large, similar in proportion only to Neandertals and some gorillas, suggesting the presence of powerful forearms and hands. There are no indicators of adaptations to knuckle-walking or suspensory locomotion in the hand, and the pedal phalanx suggests that this hominid was habitually bipedal.     

Ontogeny of the knee joint in humans, great apes and fossil hominids: pelvi-femoral relationships during postnatal growth in humans

Results of a study of the femoral bicondylar angle in adult and juvenile humans and great apes are presented. These results raise the question of whether or not the measurement reference of this angle is valid. This is because humans and great apes have a very different growth process of the distal epiphyseal suture of the femur during the period between birth and adulthood. The approximately 3 million years old juvenile femoral diaphyses attributed to Australopithecus afarensis (AL 333-110 and AL 333-111) were also studied. These specimens show an insertion of the diaphysis into the epiphysis of the simplified type typical of modern humans. This region is more convoluted in nonhuman anthropoids. Pelvifemoral interrelations are investigated through both longitudinal and cross-sectional radiographic studies of 23 human children. Growth changes in bicondylar and collo-diaphyseal angles, total femoral and femoral neck lengths, and interacetabular distance are correlated with age and to each other. These results are used to demonstrate the distinctive features of the Australopithecus afarensis fossil, AL 288-1.     

Does early care affect joint attention in great apes (Pan troglodytes, Pan paniscus, Pongo abelii, Pongo pygmaeus, Gorilla gorilla)?

The ability to share attention with another is the foundation on which other theory of mind skills are formed. The quality of care received during infancy has been correlated with increased joint attention in humans. The purpose of this study was to assess the effects of care style (responsive or basic) and caregiver type (ape or human) during the first 6 months on joint attention in 4 great ape species (Pan troglodytes, Gorilla gorilla, Pongo spp., and Pan pansicus). Great apes engaged in joint attention with conspecifics and humans regardless of the style of early care they experienced from either a great ape mother or human caregiver. This finding suggests that joint attention is a robust ability in great apes that is resilient against at least some differences in early care. Future studies using additional measures of early care quality are recommended. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Making inferences about the location of hidden food: Social dog, causal ape.

Domestic dogs (Canis familiaris) and great apes from the genus Pan were tested on a series of object choice tasks. In each task, the location of hidden food was indicated for subjects by some kind of communicative, behavioral, or physical cue. On the basis of differences in the ecologies of these 2 genera, as well as on previous research, the authors hypothesized that dogs should be especially skillful in using human communicative cues such as the pointing gesture, whereas apes should be especially skillful in using physical, causal cues such as food in a cup making noise when it is shaken. The overall pattern of performance by the 2 genera strongly supported this social-dog, causal-ape hypothesis. This result is discussed in terms of apes' adaptations for complex, extractive foraging and dogs' adaptations, during the domestication process, for cooperative communication with humans. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Prenatal gonadal hormones and sex differences in human behavior.

Summarizes the literature on perinatal hormone influences in infrahuman mammals, reviews in detail studies on perinatal hormone influences on human behavior, and summarizes methodological procedures that have been used to compensate for problems inherent in research on clinical populations. Several studies provide evidence suggesting that some sex differences in human behavior may be related to prenatal hormone levels. In light of methodological advances and the growing number of studies in this area, it may soon be possible to define human hormone–behavior relations in sufficient detail to allow strong links to research on perinatal hormone effects in other species. Given recent identification of neural sex differences that may underlie behavioral sex differences in infrahuman mammals, such links could lead to greater understanding of the neural basis of sexually dimorphic behaviors in humans. (4 p ref) (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Chloroplast and nuclear gene sequences indicate late Pennsylvanian time for the last common ancestor of extant seed plants

We have estimated the time for the last common ancestor of extant seed plants by using molecular clocks constructed from the sequences of the chloroplastic gene coding for the large subunit of ribulose-1,5-bisphosphate carboxylase/oxygenase (rbcL) and the nuclear gene coding for the small subunit of rRNA (Rrn18). Phylogenetic analyses of nucleotide sequences indicated that the earliest divergence of extant seed plants is likely represented by a split between conifer-cycad and angiosperm lineages. Relative-rate tests were used to assess homogeneity of substitution rates among lineages, and annual angiosperms were found to evolve at a faster rate than other taxa for rbcL and, thus, these sequences were excluded from construction of molecular clocks. Five distinct molecular clocks were calibrated using substitution rates for the two genes and four divergence times based on fossil and published molecular clock estimates. The five estimated times for the last common ancestor of extant seed plants were in agreement with one another, with an average of 285 million years and a range of 275-290 million years. This implies a substantially more recent ancestor of all extant seed plants than suggested by some theories of plant evolution.     

Gravity and solidity in four great ape species (Gorilla gorilla, Pongo pygmaeus, Pan troglodytes, Pan paniscus): Vertical and horizontal variations of the table task.

Three experiments modeled after infant studies were run on four great ape species (Gorilla gorilla, Pongo pygmaeus, Pan troglodytes, Pan paniscus) to investigate their reasoning about solidity and gravity constraints. The aims were: (a) to find out if great apes are subject to gravity biased search or display sensitivity for object solidity, (b) to check for species differences, and (c) to assess if a gravity hypothesis or more parsimonious explanations best account for failures observed. Results indicate that great apes, unlike monkeys, show no reliable gravity bias, that ape species slightly differ in terms of their performance, and that the errors made are best explained by a gravity account. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Sex differences and androgen-dependent regulation of aromatase (CYP19) mRNA expression in the developing and adult rat brain

Sex differences, androgen dependence and asymmetries of aromatase activity have been reported during ontogeny of the rat. It remains to be elucidated, however, whether the changes in aromatase activity are reflected by similar changes in specific mRNA levels. In addition, very little is known regarding mechanism(s) underlying such differential regulation of aromatase expression. To address these questions, we have employed the in situ hybridization (ISH) technique to examine specific mRNA levels in the brain of both male and female rats at selected stages of development. In prenatal stages of development, at gestational day (GD) 18 and 20, aromatase mRNA was detected in several hypothalamic and limbic brain regions. Semiquantitative analysis of aromatase mRNA did not reveal statistically significant sex differences in any of these regions (except in one experiment at GD20, when a sex difference was found in the medial preoptic nucleus). In contrast, clear sex differences were determined at postnatal day (PN) 2; male animals contained significantly more aromatase mRNA in the bed nucleus of the stria terminalis (BST) and the sexually dimorphic nucleus of the preoptic area (SDN) compared to female rats. Four days later in development, at PN6, sex differences of aromatase mRNA signals were observed in the BST, but were no longer detectable in the SDN. At PN15 and in adult animals, no sex differences could be determined. The effect of flutamide treatment (50 mg/kg/day) was investigated in GD20 fetuses as well as in adult rats. No statistically significant changes in aromatase mRNA expression were found in either case. In summary, our results suggest that differential regulation of aromatase mRNA expression during the critical period of sexual differentiation might, in part, account for the establishment of some of the many sexually dimorphic parameters of the rat brain. The role of androgens in the regulation of the sex-specific and developmental expression of aromatase mRNA in the rat brain remains to be clarified.     

Mental evolution and development: Evidence for secondary representation in children, great apes, and other animals.

Recent interest in the development and evolution of theory of mind has provided a wealth of information about representational skills in both children and animals. According to J. Perner (1991), children begin to entertain secondary representations in the 2nd year of life. This advance manifests in their passing hidden displacement tasks, engaging in pretense and means-ends reasoning, interpreting external representations, displaying mirror self-recognition and empathic behavior, and showing an early understanding of "mind" and imitation. New data show a cluster of mental accomplishments in great apes that is very similar to that observed in 2-year-old humans. It is suggested that it is most parsimonious to assume that this cognitive profile is of homologous origin and that great apes possess secondary representational capacity. Evidence from animals other than apes is scant. This analysis leads to a number of predictions for future research. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Bipedal posture and hand preference in humans and other primates.

Hand preference for quadrupedal and bipedal reaching in humans and rhesus macaques (Macaca mulatta) was examined, and the data were compared with postural reaching data that have been reported for 8 other primate species. Population-level biases were found toward use of the right hand for quadrupedal and bipedal reaching in humans and use of the left hand for quadrupedal reaching in rhesus macaques. Rhesus macaques showed a significant shift toward greater use of the right hand for bipedal vs. quadrupedal reaching. Comparisons with other species showed significant variance in the direction and strength of hand preference across reaching postures. The study noted right-hand biases for bipedal reaching in humans, great apes, and tufted capuchins and shifts toward greater use of the right hand for bipedal vs. quadrupedal reaching in great apes, tufted capuchins, and rhesus macaques. These results suggest that posture alters both the direction and strength of primate hand preference and that bipedalism may have facilitated species-typical right-handedness in humans. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Earliest known Old World monkey skull

Similarities of the skull are commonly used to support hypotheses of ancestor-descendant relationships between fossil and living ape genera, especially between the late Miocene apes Sivapithecus and Dryopithecus from Eurasia and the living orang-utan (Pongo) from Borneo and Sumatra. Yet determining whether craniofacial traits shared by extant and Miocene apes are primitive or derived is severely hampered by the rarity of well-preserved fossil crania, particularly of early members of their closest outgroup, the Old World monkeys (Cercopithecoidea). The discovery of a complete and undistorted skull of Victoriapithecus at middle Miocene deposits from Maboko Island, Kenya, provides evidence of intact cranial-vault and basicranial morphology, brain size and craniofacial hafting for a primate from between 32 and 7 million years ago. Victoriapithecus represents a branch of Old World monkey that is intermediate between extant cercopithecids (Colobinae and Cercopithecinae) and the common ancestor they shared with apes (Hominoidea). The skull preserves traits widely thought to be derived for extant and fossil members of a proposed Sivapithecus/Pongo clade, but which now appear to be primitive features of ancestral Old World higher primates in general.     

Restoration of the type and palate of Ankarapithecus meteai: taxonomic and phylogenetic implications

A small number of large hominoid specimens are known from the late Miocene of Turkey. (This paper focuses on the two specimens known until 1996. New fossils described by Alpagut et al. ([1996] Nature 382:349-351) are briefly discussed in this paper as well.) Among these, a fragmentary mandible is the type specimen of Ankarapithecus meteai. Another specimen, a partial face, is from the same taxon. Based on the morphology of this face, Ankarapithecus meteai was synonymized with Sivapithecus (Andrews and Tekkaya [1980], Paleontology 23:85-95). The facial specimen was recently restored by the authors, exposing much anatomy that had previously been obscured. The new anatomical details reveal many important differences from Sivapithecus and justify a revision of the nomen Ankarapithecus. Ankarapithecus was a large hominid (great apes and humans) in the clade that also includes Sivapithecus and Pongo. Sivapithecus and Pongo share derived characters not found in Ankarapithecus, which is thus the sister clade to the Sivapithecus-Pongo clade. While the results of this analysis support the sister relationship of Sivapithecus and Pongo, there is some uncertainty regarding paleobiogeographic and taxonomic relationships to the large hominids from the Siwaliks. An Ankarapithecus-like taxon may have been ancestral to Sivapithecus, or an early Siwalik (Chinji formation) taxon, which predates both Sivapithecus sensu stricto and Ankarapithecus, may be ancestral to both.     

The evolution of the frontal lobes: a volumetric analysis based on three-dimensional reconstructions of magnetic resonance scans of human and ape brains

Scenarios regarding the evolution of cognitive function in hominids depend largely on our understanding of the organization of the frontal lobes in extant humans and apes. The frontal lobe is involved in functions such as creative thinking, planning of future actions, decision making, artistic expression, aspects of emotional behavior, as well as working memory, language and motor control. It is often claimed that the frontal lobe is disproportionately larger in humans than in other species, but conflicting reports exist on this issue. The brain of the apes in particular remains largely unknown. In this report we measure the volume of the frontal lobe as a whole and of its main sectors (including cortex and immediately underlying white matter) in living humans, and in post-mortem brains of the chimpanzee, gorilla, orang-utan, gibbon and the macaque using three-dimensional reconstructions of magnetic resonance (MR) scans of the brain. On the basis of these data we suggest that although the absolute volume of the brain and the frontal lobe is largest in humans, the relative size of the frontal lobe is similar across hominoids, and that humans do not have a larger frontal lobe than expected from a primate brain of the human size. We also report that the relative size of the sectors of the frontal lobe (dorsal, mesial, orbital) is similar across the primate species studied. Our conclusions are preliminary, because the size of our sample, although larger than in previous studies, still remains small. With this caveat we conclude that the overall volume of the frontal lobe in hominids enlarged in absolute size along with the rest of the brain, but did not become relatively larger after the split of the human line from the ancestral African hominoid stock. Aspects other than relative volume of the frontal lobe have to be responsible for the cognitive specializations of the hominids.     

Comparative and familial analysis of handedness in great apes.

Historically, population-level handedness has been considered a hallmark of human evolution. Whether nonhuman primates exhibit population-level handedness remains a topic of considerable debate. This paper summarizes published data on handedness in great apes. Comparative analysis indicated that chimpanzees and bonobos show population-level right handedness, whereas gorillas and orangutans do not. All ape species showed evidence of population-level handedness when considering specific tasks. Familial analyses in chimpanzees indicated that offspring and maternal (but not paternal) handedness was significantly positively correlated, but this finding was contingent upon the classification criteria used to evaluate hand preference. Overall, the proportion of right handedness is lower in great apes compared with humans, and various methodological and theoretical explanations for this discrepancy are discussed. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Sex Differences and Individual Differences in Cognitive Performance and Their Relationship to Endogenous Gonadal Hormones and Gonadotropins.

Sexually dimorphic cognitive performance in men (n=42) and women (n=42) was related to testosterone, estradiol, progesterone, luteinizing hormone, follicle-stimulating hormone, and sex hormone binding globulin, measured in 10-ml blood samples collected between 0900 and 1030 and, among women, during the follicular phase of the menstrual cycle. Significant sex differences favored men on spatial tasks (Mental Rotation and Judgment of Line Orientation) and on an inhibition task and favored women on a verbal task (category fluency). However, there were no significant relationships between any of the hormones and cognitive performance, suggesting that there are few, if any, consistent, substantial relationships between endogenous, nonfluctuating levels of gonadal hormones or gonadotropins and these cognitive abilities in men or women. (PsycINFO Database Record (c) 2010 APA, all rights reserved)