Growth changes in internal and craniofacial flexion measurements.

Growth changes in both internal and craniofacial flexion angles are presented for Pan troglodytes, Gorilla gorilla, and modern humans. The internal flexion angle (IFA) was measured from lateral radiographs, and the craniofacial flexion angle (CFA) was calculated from coordinate data. Stage of dental development is used as a baseline for examination of growth changes and nonparametric correlations between flexion angles and dental development stage are tested for significance. In Gorilla, the IFA increases during growth. The IFA is relatively stable in Pan and modern humans. Pan and Gorilla display an increase in the CFA. However, this angle decreases during growth in modern humans. Flexion angles were derived from coordinate data collected for several early hominid crania. Measurements for two robust australopithecine crania indicate strong internal flexion. It has been suggested that cerebellar expansion in this group may relate to derived features of the posterior cranial base. In general, australopithecine crania exhibit craniofacial flexion intermediate between great apes and modern humans. The "archaic" Homo sapiens specimen from Kabwe is most similar to modern humans.     

The Ontogeny of Vocal Sequences: Insights from a Newborn Wild Chimpanzee (Pan troglodytes schweinfurthii)

International Journal of Primatology - Observations of early vocal behaviours in non-human primates (hereafter primates) are important for direct comparisons between human and primate vocal...     

Vocal production mechanisms in a non-human primate: morphological data and a model

Human beings are thought to be unique amongst the primates in their capacity to produce rapid changes in the shape of their vocal tracts during speech production. Acoustically, vocal tracts act as resonance chambers, whose geometry determines the position and bandwidth of the formants. Formants provide the acoustic basis for vowels, which enable speakers to refer to external events and to produce other kinds of meaningful communication. Formant-based referential communication is also present in non-human primates, most prominently in Diana monkey alarm calls. Previous work has suggested that the acoustic structure of these calls is the product of a non-uniform vocal tract capable of some degree of articulation. In this study we test this hypothesis by providing morphological measurements of the vocal tract of three adult Diana monkeys, using both radiography and dissection. We use these data to generate a vocal tract computational model capable of simulating the formant structures produced by wild individuals. The model performed best when it combined a non-uniform vocal tract consisting of three different tubes with a number of articulatory manoeuvres. We discuss the implications of these findings for evolutionary theories of human and non-human vocal production.     

Social learning of a communicative signal in captive chimpanzees

The acquisition of linguistic competency from more experienced social partners is a fundamental aspect of human language. However, there is little evidence that non-human primates learn to use their vocalizations from social partners. Captive chimpanzees (Pan troglodytes) produce idiosyncratic vocal signals that are used intentionally to capture the attention of a human experimenter. Interestingly, not all apes produce these sounds, and it is unclear what factors explain this difference. We tested the hypothesis that these attention-getting (AG) sounds are socially learned via transmission between mothers and their offspring. We assessed 158 chimpanzees to determine if they produced AG sounds. A significant association was found between mother and offspring sound production. This association was attributable to individuals who were raised by their biological mother-as opposed to those raised by humans in a nursery environment. These data support the hypothesis that social learning plays a role in the acquisition and use of communicative vocal signals in chimpanzees.     

A morphometric analysis of craniofacial growth and changes in spatial relations during secondary palatal development in human embryos and fetuses

Staged human embryos and fetuses in the Carnegie Embryological Collection were morphometrically analyzed to show craniofacial dimensions and changes in spatial relations, and to identify patterns that would reflect normal developmental events during palatal formation. Normal embryos aged 7-8 weeks postconception (Streeter-O'Rahilly stages 19-23) and fetuses aged 9-10 weeks postconception, in eight groups with mean crown-rump (CR) lengths of 18-49 mm, were studied with cephalometric methods developed for histologic sections. In the 4-week period studied, facial dimensions increased predominantly in the sagittal plane with extensive changes in length (depth) and height, but limited changes in width. Growth of the mandible was more rapid than the nasomaxillary complex, and the length of Meckel's cartilage exceeded the length of the oronasal cavity at the time of horizontal movement of the shelves during stage 23. Simultaneously with shelf elevation, the upper craniofacial complex lifted, and the tongue and Meckel's cartilage extended forward beneath the primary palate. Analysis of spatial relations in the oronasal cavity showed that the palatomaxillary processes became separated from the tongue--mandibular complex as the head extended, and the tongue became positioned forward with growth of Meckel's cartilage. As the head position extended by 35 degrees, the cranial base angulation was unchanged and the primary palate maintained a 90 degrees position to the posterior cranial base. However, the sagittal position of the maxilla relative to the anterior cranial base increased by 20 degrees between stages 19 and 23. In the late embryonic and early fetal periods, the mean cranial base angulation of approximately 128 degrees and the mean maxillary position angulation of approximately 84 degrees were similar to the angulations previously shown to be present later prenatally and post-natally. The results suggest that human patterns of cranial base angulation and maxillary position to the cranial base develop during the late embryonic period when the chondrocranium and Meckel's cartilage form the primary skeleton.     

The neural origins and implications of imitation, mirror neurons and tool use

Several recent studies report how laboratory-raised, non-human primates exposed to tool use can exhibit intelligent behaviors, such as imitation and reference vocal control, that are never seen in their wild counterparts. Tool-use training appears to forge a novel cortico-cortical connection that underlies this boost in capacity, which normally exists only as latent potential in lower primates. Although tool-use training is patently non-naturalistic, its marked effects on brain organization and behavior could shed light on the evolution of higher intelligence in humans.     

Meningeal arterial patterns in great apes: Implications for hominid vascular evolution

Arterial meningeal patterns were observed for 100 hemispheres from great ape endocasts (Pan paniscus, Pan troglodytes, Gorilla gorilla, and Pongo pygmaeus). Eight patterns emerged based on the relative contributions to the walls and dura mater of the middle part of the braincase of meningeal arteries that stem from two sources. These arteries enter the braincase through either the orbit (delivering blood from the internal carotid artery) or through the base of the middle cranial fossa (via the middle meningeal artery whose blood comes from the external carotid artery). The three genera of apes manifest different frequencies of the eight, patterns, with orangutans highly dependent on orbital meningeal arteries at one extreme, and chimpanzees showing the greatest reliance on the middle meningeal artery at the other. As was the case in an earlier study of rhesus monkeys, there is a trend across the two genera of African apes for increased mean cranial capacity to be associated with increased reliance on the internal carotid artery for supplying the middle portion of the braincase. However, unlike the case for macaques, this trend does not reach statistical significance in African apes. Because it is rare for humans to manifest significant arterial contributions from the orbit to the middle cranial fossa, the comparative data on monkeys, apes, and humans suggest that, during the course of vascular evolution in Homo, the middle meningeal artery eventually took over supply of the entire middle cranial fossa. This hypothesis should be tested in the hominid fossil record. Earlier work on meningeal arterial patterns in apes has traditionally relied on Adachi's system that was determined from humans and focuses on the origin of the middle branch of the middle meningeal artery. As a result, the extensive orbital contributions to the middle portion of the braincase that characterize apes were not recognized and the eight patterns described in this paper were often erroneously assigned to the three patterns that adequately describe only humans. Adachi's system should therefore be abandoned for nonhuman primates and early hominids. A correct understanding of meningeal arterial evolution cannot be achieved until the orbital contributions to the meningeal arteries are recognized and incorporated into an evolutionary study that spans from apes to fossil hominids to living people. © 1993 Wiley-Liss, Inc.     

Human is a unique species among primates in terms of telomere length.

TRF (terminal restriction fragments) length in various tissues of non-human primates such as Macaca mulatta (rhesus monkey), Macaca fuscata (Japanese monkey), Macaca fascicularis (crab-eating monkey), Pan troglodytes (common chimpanzee), and Pongo pygmaeus (orangutan) was at least 23 kb without exception, which was quite different from that of human somatic tissues (smaller than 10 kb). The distribution pattern of telomerase activity among tissues was similar between human and non-human primates, while the activity level showed some differences such as that strong telomerase activity was observed in gastrointestinal and lymphocytic tissues from non-human primates. The human appears to be a unique species among primates in terms of telomere length.     

Bivariate and multivariate growth allometry: statistical and biological considerations

Bivariate and multivariate analyses of 55 cranial dimensions were completed on ontogenetic series of Pygmy chimpanzees ( Pan paniscus ), Common chimpanzees ( Pan troglodytes ), and gorillas ( Gorilla gorilla ). Subanalyses described here were specifically designed to compare and crosscheck quantitative assessments of relative growth as determined using the bivariate and multivariate (principal components analysis—PCA) approaches. Results indicate a strong concordance between the bivariate and multivariate patterns, empirically supporting the claim that PCA provides an effective multivariate approach to analysing growth allometry. Comparison of bivariate and multivariate results also suggests that in multi-group PCA, the first component summarizes shape variation resulting from the sharing and extension of common patterns of growth allometry, while the second and subsequent components summarize shape variation resulting from divergent growth trajectories, reflected in bivariate comparisons as either vertical shifts and/or slope differences. Examination of non-normalized first component loadings, plus a comparison with estimates of logarithmic growth in the cranial dimensions, reveals that the non-normalized loadings are proportional to coefficients of specific growth. This finding further links the bivariate and multivariate approaches, grounding both in Huxley's theoretical notions of multiplicative and relative growth.     

Economic behaviours among non-human primates

Do we have any valid reasons to affirm that non-human primates display economic behaviour in a sufficiently rich and precise sense of the phrase? To address this question, we have to develop a set of criteria to assess the vast array of experimental studies and field observations on individual cognitive and behavioural competences as well as the collective organization of non-human primates. We review a sample of these studies and assess how they answer to the following four main challenges. (i) Do we see any economic organization or institutions emerge among groups of non-human primates? (ii) Are the cognitive abilities, and often biases, that have been evidenced as underlying typical economic decision-making among humans, also present among non-human primates? (iii) Can we draw positive lessons from performance comparisons among primate species, humans and non-humans but also across non-human primate species, as elicited by canonical game-theoretical experimental paradigms, especially as far as economic cooperation and coordination are concerned? And (iv) in which way should we improve models and paradigms to obtain more ecological data and conclusions? Articles discussed in this paper most often bring about positive answers and promising perspectives to support the existence and prevalence of economic behaviours among non-human primates.This article is part of the theme issue ‘Existence and prevalence of economic behaviours among non-human primates’.     

Species association of hepatitis B virus (HBV) in non-human apes; evidence for recombination between gorilla and chimpanzee variants

Hepatitis B virus (HBV) infections are widely distributed in humans, infecting approximately one third of the world's population. HBV variants have also been detected and genetically characterised from Old World apes; Gorilla gorilla (gorilla), Pan troglodytes (chimpanzee), Pongo pygmaeus (orang-utan), Nomascus nastusus and Hylobates pileatus (gibbons) and from the New World monkey, Lagothrix lagotricha (woolly monkey). To investigate species-specificity and potential for cross species transmission of HBV between sympatric species of apes (such as gorillas and chimpanzees in Central Africa) or between humans and chimpanzees or gorillas, variants of HBV infecting captive wild-born non-human primates were genetically characterised. 9 of 62 chimpanzees (11.3%) and two from 11 gorillas (18%) were HBV-infected (15% combined frequency), while other Old world monkey species were negative. Complete genome sequences were obtained from six of the infected chimpanzee and both gorillas; those from P. t .ellioti grouped with previously characterised variants from this subspecies. However, variants recovered from P. t. troglodytes HBV variants also grouped within this clade, indicative of transmission between sub-species, forming a paraphyletic clade. The two gorilla viruses were phylogenetically distinct from chimpanzee and human variants although one showed evidence for a recombination event with a P.t.e.-derived HBV variant in the partial X and core gene region. Both of these observations provide evidence for circulation of HBV between different species and sub-species of non-human primates, a conclusion that differs from the hypothesis if of strict host specificity of HBV genotypes.     

Phylogeny, neoteny and growth of the cranial base in hominoids

This study tests the hypothesis that there is a general pattern in the growth of the cranial base of Homo sapiens that is 'essentially neotenous' [Gould, 1977]. Juvenile and adult crania of Homo sapiens, Gorilla gorilla, Pan troglodytes and Pongo pygmaeus were studied and the cross-sectional growth curves for 10 measurements made on the cranial base (as viewed in norma basilaris) were compared. The results of this study suggest that relatively simple modifications to the timing or pattern of growth are insufficient to explain the observed morphological differences between the cranial base of modern Homo sapiens and the great apes.     

Comparison of cranial ontogenetic trajectories among great apes and humans

Molecular data suggest that humans are more closely related to chimpanzees than either is to the gorillas, yet one finds the closest similarity in craniofacial morphology to be among the great apes to the exclusion of humans. To clarify how and when these differences arise in ontogeny, we studied ontogenetic trajectories for Homo sapiens, Pan paniscus, Pan troglodytes, Gorilla gorilla and Pongo pygmaeus. A total of 96 traditional three-dimensional landmarks and semilandmarks on the face and cranial base were collected on 268 adult and sub-adult crania for a geometric morphometric analysis. The ontogenetic trajectories are compared by various techniques, including a new method, relative warps in size-shape space. We find that adult Homo sapiens specimens are clearly separated from the great apes in shape space and size-shape space. Around birth, Homo sapiens infants are already markedly different from the great apes, which overlap at this age but diverge among themselves postnatally. The results suggest that the small genetic differences between Homo and Pan affect early human ontogeny to induce the distinct adult human craniofacial morphology. Pure heterochrony does not sufficiently explain the human craniofacial morphology nor the differences among the African apes.     

Unconstrained cranial evolution in Neandertals and modern humans compared to common chimpanzees

A variety of lines of evidence support the idea that neutral evolutionary processes (genetic drift, mutation) have been important in generating cranial differences between Neandertals and modern humans. But how do Neandertals and modern humans compare with other species? And how do these comparisons illuminate the evolutionary processes underlying cranial diversification? To address these questions, we used 27 standard cranial measurements collected on 2524 recent modern humans, 20 Neandertals and 237 common chimpanzees to estimate split times between Neandertals and modern humans, and between Pan troglodytes verus and two other subspecies of common chimpanzee. Consistent with a neutral divergence, the Neandertal versus modern human split-time estimates based on cranial measurements are similar to those based on DNA sequences. By contrast, the common chimpanzee cranial estimates are much lower than DNA-sequence estimates. Apparently, cranial evolution has been unconstrained in Neandertals and modern humans compared with common chimpanzees. Based on these and additional analyses, it appears that cranial differentiation in common chimpanzees has been restricted by stabilizing natural selection. Alternatively, this restriction could be due to genetic and/or developmental constraints on the amount of within-group variance (relative to effective population size) available for genetic drift to act on.     

非人灵长类声音通讯的研究进展

声音通讯是非人灵长类研究一个重要的研究领域,有助于了解非人灵长类的社会行为、个体关系、行为进化和社会演化等,甚至对探究人类语言起源和进化等方面也具有十分重要的意义。本文通过对非人灵长类声音通讯的研究内容、影响因素和研究方法等进行了梳理,探讨非人灵长类声音通讯研究的前景和展望,旨在进一步推动国内非人灵长类声音通讯研究的深入,同时为相关研究提供借鉴和参考。     

Chimpanzees Show a Developmental Increase in Susceptibility to Contagious Yawning: A Test of the Effect of Ontogeny and Emotional Closeness on Yawn Contagion

Contagious yawning has been reported for humans, dogs and several non-human primate species, and associated with empathy in humans and other primates. Still, the function, development and underlying mechanisms of contagious yawning remain unclear. Humans and dogs show a developmental increase in susceptibility to yawn contagion, with children showing an increase around the age of four, when also empathy-related behaviours and accurate identification of others’ emotions begin to clearly evince. Explicit tests of yawn contagion in non-human apes have only involved adult individuals and examined the existence of conspecific yawn contagion. Here we report the first study of heterospecific contagious yawning in primates, and the ontogeny of susceptibility thereto in chimpanzees, Pan troglodytes verus. We examined whether emotional closeness, defined as attachment history with the yawning model, affected the strength of contagion, and compared the contagiousness of yawning to nose-wiping. Thirty-three orphaned chimpanzees observed an unfamiliar and familiar human (their surrogate human mother) yawn, gape and nose-wipe. Yawning, but not nose-wiping, was contagious for juvenile chimpanzees, while infants were immune to contagion. Like humans and dogs, chimpanzees are subject to a developmental trend in susceptibility to contagious yawning, and respond to heterospecific yawn stimuli. Emotional closeness with the model did not affect contagion. The familiarity-biased social modulatory effect on yawn contagion previously found among some adult primates, seem to only emerge later in development, or be limited to interactions with conspecifics. The influence of the ‘chameleon effect’, targeted vs. generalised empathy, perspective-taking and visual attention on contagious yawning is discussed.     

PCDH11 is X/Y homologous in Homo sapiens but not in Gorilla gorilla and Pan troglodytes

Protocadherin X (PCDHX) and Protocadherin Y (PCDHY) are cell-surface adhesion molecules expressed predominantly in brain. The human PCDH11X/Y gene pair is located in the non-pseudoautosomal X-Y homologous region (Xq21.3/Yp11.2). The possible existence of PCDH11 gene dosage differences between human and non-human primates is of evolutionary significance with respect to species differences and escape from X inactivation, and has been repeatedly debated. Previous investigations on the X/Y homologous status of PCDH11 and adjacent sequences in non-human primates have highlighted the complexity of the molecular pattern and evolutionary history of this genomic region. This paper provides for the first time direct evidence for the absence of the PCDH11 genefrom the Y chromosome of chimpanzee (Pan troglodytes) as well as gorilla (Gorilla gorilla). By confirmingthe suspected lack of X-Y homologous status for PCDH11 in non-human primates, our results reinforce the hypothesis of a hominid-specific role for this gene in brain development.     

Rank and relationships in the evolution of spoken language

If evolutionary benefits associated with language were predominantly referential, as many theorists assume, then there must have been a preparatory stage in which an 'appetite' for information, not evident in the other primates, developed. To date, no such stage has been demonstrated. The problem dissipates, however, if it is assumed that language emerged from a function more nearly shared with other primates. An obvious candidate is displaying. Here I argue that performative functions associated with oral sound-making provided initial pressures for vocal communication by promoting rank and relationships. These benefits, I suggest, facilitated conflict avoidance and resolution, collaboration, and reciprocal sharing of needed resources. By valuing the performative applications of language, which continue in modern humans, one can more easily derive speech from the social-vocal behaviours of non-human primates, providing greater continuity in accounts of linguistic evolution     

Anthropoid cranial base architecture and scaling relationships

This paper examines how various measures of basicranial length and cranial base angulation affect the relationship between basicranial flexion and relative brain size in anthropoids, including Homo sapiens. Most recent studies support the "spatial packing" hypothesis, that basicranial flexion in haplorhines maximizes braincase volume relative to basicranial length. However, a few studies find the basicranium is less flexed in H. sapiens than expected for other anthropoids, suggesting that other factors contribute to variation in hominin basicranial flexion. The measure of relative brain size used to test the spatial packing hypothesis, the Index of Relative Encephalization (IRE), is calculated with basicranial length (BL) in its denominator, so that shorter BL and larger brain size potentially inflate H. sapiens IREs. To investigate this problem, the lengths of midline cranial floor sections were scaled relative to the cube root of endocranial volume in 157 specimens from 18 anthropoid species. Results indicate that the posterior cranial base and planum sphenoideum are significantly shorter in H. sapiens than in other anthropoids, accounting for higher IREs. Including the cribriform plate in BL, advisable in studies using anthropoids, affects whether H. sapiens differs from other anthropoids for basicranial flexion vs. IRE. However, despite a shorter BL and elevated IRE, H. sapiens does not deviate significantly from the anthropoid relationship between basicranial flexion and relative brain size for two cranial base angles. Because different measures of cranial base angulation change how H. sapiens falls along the anthropoid regression line, it remains equivocal whether the basicranium is less flexed in H. sapiens than in other anthropoids when compared to relative brain size.     

Relative growth of the skull and postcranium in giant transgenic mice

Cross-sectional allometric growth patterns of the cranial and postcranial skeleton were compared between giant transgenic (MT-rGH) mice and their normal littermate controls. Body weights, external body dimensions, and a series of cranial and postcranial linear dimensions of the skeleton were determined for samples of known age. Comparative bivariate and multivariate allometric analyses were completed in order to determine whether (1) the larger transgenic mice differed significantly from the normal controls in aspects of body and skeletal proportions, and (2) any such proportion differences resulted from general allometric effects of overall weight or skeletal size increase. Results demonstrate that the transgenic mice do exhibit significantly different body and skeletal proportions than normal control adults. Allometric comparisons of the skeletal dimensions relative to body weight reveal similar coefficients of growth allometry but several differences in gamma-intercept values in the transgenic vs. control groups. The comparisons among the skeletal dimensions of the skull and postcranium generally reveal the sharing and differential extension of common growth allometries in the two groups. Thus, the elevated levels of growth hormone (GH) and insulin-like growth factor I (IGF-I) in the transgenic mice appear to result in increased overall growth for the various skeletal elements, but in the relative proportions determined by intrinsic growth controls within that system.