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[The Great Ape Project--human rights for the great anthropoid apes].

The Great Ape Project (GAP) is an appeal of 36 scientist from different disciplines aiming at the legal equalisation of the non-human great apes (chimpanzees, gorillas and orang-utans) with man. The appeal is expressed by a number of essays stating zoological, genetical, ethological, anthropological, ethical and psychological knowledge and, based on these arguments, demanding the abolition of the species barrier between human beings and great apes. The central point of the initiative is the "Declaration on Great Apes", claiming the inclusion of great apes in the "community of equals" and thus securing three basic rights for all great apes: 1. The Right of Life; 2. The Protection of Individual Liberty; 3. The Prohibition of Torture. Not only experiments with great apes and their capture from the wilderness will be banned, but it is also intended to enfranchise as many great apes as possible from research laboratories and zoos. As a legal basis for the achievement of basic rights most of the authors plead for the idea of conferring the moral status of "persons" on great apes. Criticism of the GAP is due to its anthropocentrism. Rejection is especially expressed by advocates of pathocentric ethics who argue that the species barrier will not be abolished but only shifted, running then between the great apes and the remaining living beings. However, the GAP resulted in a greater retention in the use of great apes for experiments in several industrial countries. Additionally, the popular literature published by ethologists in the passed decades has supported a more responsible attitude of the public towards primates. Despite of all efforts the survival of the great apes is greatly endangered within their native countries.

Animal Rights↗

Genetic differences between humans and great apes.

The remarkable similarity among the genomes of humans and the African great apes could warrant their classification together as a single genus. However, whereas there are many similarities in the biology, life history, and behavior of humans and great apes, there are also many striking differences that need to be explained. The complete sequencing of the human genome creates an opportunity to ask which genes are involved in those differences. A logical approach would be to use the chimpanzee genome for comparison and the other great ape genomes for confirmation. Until such a great ape genome project can become reality, the next best approach must be educated guesses of where the genetic differences may lie and a careful analysis of differences that we do know about. Our group recently discovered a human-specific inactivating mutation in the CMP-sialic acid hydroxylase gene, which results in the loss of expression of a common mammalian cell-surface sugar throughout all cells in the human body. We are currently investigating the implications of this difference for a variety of issues relevant to humans, ranging from pathogen susceptibility to brain development. Evaluating the uniqueness of this finding has also led us to explore the existing literature on the broader issue of genetic differences between humans and great apes. The aim of this brief review is to consider a listing of currently known genetic differences between humans and great apes and to suggest avenues for future research. The differences reported between human and great ape genomes include cytogenetic differences, differences in the type and number of repetitive genomic DNA and transposable elements, abundance and distribution of endogenous retroviruses, the presence and extent of allelic polymorphisms, specific gene inactivation events, gene sequence differences, gene duplications, single nucleotide polymorphisms, gene expression differences, and messenger RNA splicing variations. Evaluation of the reported findings in all these categories indicates that the CMP-sialic hydroxylase mutation is the only one that has so far been shown to result in a global biochemical and structural difference between humans and great apes. Several of the other known genetic dissimilarities deserve more exploration at the functional level. Among the areas of focus for the future should be genes affecting development, mental maturation, reproductive biology, and other aspects of life history. The approaches taken should include both going from the genome up to the adaptive potential of the organisms and going from novel adaptive regimes down to the relevant repercussions in the genome. Also, as much as we desire a simple genetic explanation for the human phenomenon, it is much more probable that our evolution occurred in multiple genetic steps, many of which must have left detectable footprints in our genomes. Ultimately, we need to know the exact number of genetic steps, the order in which they occurred, and the temporal, spatial, environmental, and cultural contexts that determined their impact on human evolution.

Animals↗

Almost human.

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Animal Experimentation↗

The Yerkes Regional Primate Research Center.

The development and organisation of the centre which now has over 1,500 non-human primates representing more than 30 species, including great apes, is described. Important projects are mainly in the neural and behavioural field, including language acquisition, but also on aspects of experimental pathology.

Academies and Institutes↗

A neuronal morphologic type unique to humans and great apes.

We report the existence and distribution of an unusual type of projection neuron, a large, spindle-shaped cell, in layer Vb of the anterior cingulate cortex of pongids and hominids. These spindle cells were not observed in any other primate species or any other mammalian taxa, and their volume was correlated with brain volume residuals, a measure of encephalization in higher primates. These observations are of particular interest when considering primate neocortical evolution, as they reveal possible adaptive changes and functional modifications over the last 15-20 million years in the anterior cingulate cortex, a region that plays a major role in the regulation of many aspects of autonomic function and of certain cognitive processes. That in humans these unique neurons have been shown previously to be severely affected in the degenerative process of Alzheimer's disease suggests that some of the differential neuronal susceptibility that occurs in the human brain in the course of age-related dementing illnesses may have appeared only recently during primate evolution.

Alzheimer Disease↗

Genome of the apes.

The Human Genome Project has generated both the information and technological infrastructure needed to accelerate genetic comparisons between humans and the African great apes (chimpanzees and gorillas). Sequence and chromosomal organization differences between these highly related genomes will provide clues to the genetic basis for recently evolved, specifically human traits such as bipedal gait and advanced cognitive function. Recent studies comparing the primate genomes have the potential to affect many aspects of human biomedical research and could benefit primate conservation efforts.

Animals↗

Polymorphic aspects of male anthropoid canines.

Interspecific variation in the architecture of male anthropoid maxillary canines is documented. Extant taxa are polymorphic, and most can be sorted into two major groupings based on quantitative measures of shape, distal edge sharpness, and interspecific changes in their linear dimensions (projection, mesiodistal length, and buccolingual breadth) relative to each other and to body mass (scaling). One group includes the great apes and ceboids; the other includes cercopithecoids and hylobatids. Statistically significant differences between these groups were found for canine shape, for trajectories of regressions for canine projection on canine length and canine breadth, and for canine projection and canine breadth relative to body mass. The data indicate that explantations of canine variation in male anthropoids must include a mechanical interpretation of form in addition to assessments of habitus, heritage, and body mass.

Animals↗

The Ai project: historical and ecological contexts.

This paper aims to review a long-term research project exploring the chimpanzee mind within historical and ecological contexts. The Ai project began in 1978 and was directly inspired by preceding ape-language studies conducted in Western countries. However, in contrast with the latter, it has focused on the perceptual and cognitive capabilities of chimpanzees rather than communicative skills between humans and chimpanzees. In the original setting, a single chimpanzee faced a computer-controlled apparatus and performed various kinds of matching-to-sample discrimination tasks. Questions regarding the chimpanzee mind can be traced back to Wolfgang Koehler's work in the early part of the 20th century. Yet, Japan has its unique natural and cultural background: it is home to an indigenous primate species, the Japanese snow monkey. This fact has contributed to the emergence of two previous projects in the wild led by the late Kinji Imanishi and his students. First, the Koshima monkey project began in 1948 and became famous for its discovery of the cultural propagation of sweet-potato washing behavior. Second, pioneering work in Africa, starting in 1958, aimed to study great apes in their natural habitat. Thanks to the influence of these intellectual ancestors, the present author also undertook the field study of chimpanzees in the wild, focusing on tool manufacture and use. This work has demonstrated the importance of social and ecological perspectives even for the study of the mind. Combining experimental approaches with a field setting, the Ai project continues to explore cognition and behavior in chimpanzees, while its focus has shifted from the study of a single subject toward that of the community as a whole.

Adaptation, Psychological↗

The 200-kb segmental duplication on human chromosome 21 originates from a pericentromeric dissemination involving human chromosomes 2, 18 and 13.

Regions close to human centromeres contain DNA fragments spanning hundreds of kilobases that exhibit a high degree of sequence identity (>95%). Here we report the genomic structure and evolution of a family of four paralogous regions related to a 220-kb genomic fragment present on the long arm of human chromosome 21 (21q22.1). Phylogenetic classification of the paralogous sequences obtained from the draft of the Human Genome Project are in agreement with results from comparative fluorescence in situ hybridization on metaphase chromosomes from human and great apes. The original copy present in 21q22.1 in human was duplicated in great apes after the divergence of the orang-utan and inserted in a pericentromeric region, most likely the ancestor of HSA2q, then disseminated by transposition of a larger fragment to other pericentromeric locations: HSA18p11, HSA13q11 and HSA21q11.1. The degree of dissemination varies among species.

Animals↗

Great apes and rhesus monkeys as subjects for psychopharmacological studies of stimulants and depressants.

A group of experiments is described in which chimpanzees and orangutans are utilized as subjects in research projects designed to evaluate the effects of stimulant and depressant drugs on learning and performance. Efficiency of performance on a task which measures spaced responding was impaired when subjects smoked cigarettes containing delta9-tetrahydrocannabinol prior to testing. In a sequential learning task, these subjects also demonstrated reduced performance when stimulatn drugs were orally administered before testing. Depressant drugs did not produce comparable decrements in sequential learning performance. Physical and behavioral tolerance and dependence on ethanol were investigated in rhesus monkey subjects using a variety of experimental procedures, including forced oral acceptance, intragastric intubation, intravenous infusion, and conditioned voluntary oral acceptance.

Animals↗

Comparative mapping of ZFY in the hominoid apes.

Within our project of comparative mapping of candidate genes for sex-determination/testis differentiation, we used a cloned probe from the human ZFY locus for comparative hybridization studies in hominoids. As in the human, the ZFY probe detects X- and Y-specific restriction fragments in the chimpanzee, the gorilla, the orangutan, and the gibbon. Furthermore, the X-specific hybridization site in the great apes resides in Xp21.3, the same locus defining ZFX in the human. The Y-specific locus of ZFY maps closely to the early replicating pseudoautosomal segment in the telomeric or subtelomeric position of the Y chromosomes of the great apes, again as found in the human. Thus, despite cytogenetically visible structural alterations within the euchromatic parts of the Y chromosomes of the human species and the great apes, a segment of the Y chromosome defined by the pseudoautosomal region and ZFY seems to be more strongly conserved than the rest of the Y chromosome.

Animals↗

Cortical orofacial motor representation in Old World monkeys, great apes, and humans. II. Stereologic analysis of chemoarchitecture.

This study presents a comparative stereologic investigation of neurofilament protein- and calcium-binding protein-immunoreactive neurons within the region of orofacial representation of primary motor cortex (Brodmann's area 4) in several catarrhine primate species (Macaca fascicularis, Papio anubis, Pongo pygmaeus, Gorilla gorilla, Pan troglodytes, and Homo sapiens). Results showed that the density of interneurons involved in vertical interlaminar processing (i.e., calbindin- and calretinin-immunoreactive neurons) as well pyramidal neurons that supply heavily-myelinated projections (i.e., neurofilament protein-immunoreactive neurons) are correlated with overall neuronal density, whereas interneurons making transcolumnar connections (i.e., parvalbumin-immunoreactive neurons) do not exhibit such a relationship. These results suggest that differential scaling rules apply to different neuronal subtypes depending on their functional role in cortical circuitry. For example, cortical columns across catarrhine species appear to involve a similar conserved network of intracolumnar inhibitory interconnections, as represented by the distribution of calbindin- and calretinin-immunoreactive neurons. The subpopulation of horizontally-oriented wide-arbor interneurons, on the other hand, increases in density relative to other interneuron subpopulations in large brains. Due to these scaling trends, the region of orofacial representation of primary motor cortex in great apes and humans is characterized by a greater proportion of neurons enriched in neurofilament protein and parvalbumin compared to the Old World monkeys examined. These modifications might contribute to the voluntary dexterous control of orofacial muscles in great ape and human communication.

Animal Communication↗

Relative placement of the mandibular fossa in great apes and humans.

Several researchers have investigated, or commented on, the relative placement of the hominin mandibular fossa with regard to brain expansion and masticatory function. Two confounding factors are identified in this previous work. First, a number of different measurement techniques have been applied, confusing comparisons between studies. Second, the effects of squamous thickening due to temporal bone pneumatization are shown to influence measurements based relative to the ectocranial margin of the skull. To investigate the influence of these factors, a sample of adult human (n=12), chimpanzee (n=12), gorilla (n=15), and orang-utan (n=8) skulls from the Cleveland Museum of Natural History, University of Wisconsin Zoology Museum, and University of Wisconsin Anthropology collections, were CT scanned. Coronal scans were horizontally aligned and measured on a personal computer using ImageJ (NIH). To identify fossa placement, fossa breadth was measured as the projected distance in the coronal plane between the tip of the entoglenoid to lateral margin of the articular surface. A second distance, from the tip of the entoglenoid to a sagittal plane, tangent to the lateralmost margin of the endocranial surface was taken to indicate the extent of medial placement of the fossa. By eliminating the influence of pneumatization, these data unambiguously confirmed the medial placement of the human fossa and show all great apes as having a laterally placed fossa. Similar measurements on three fossil hominins, KNM-BC 1 (Homo sp. indet.), OH 5 and KNM-ER 23000 (Paranthropus boisei) demonstrate that, while all specimens demonstrate a broad fossa, only KNM-BC 1 is characterized by a relatively medial placement while the latter two display lateral placement.

Adult↗

The capacity of animals to acquire language: do species differences have anything to say to us?

Following the Gardners' discovery that an ape named Washoe could learn to produce and combine a number of hand movements similar to those used by deaf human beings, a variety of 'ape-language projects' sprang up. Some projects used different symbol systems, others used different training techniques, and others used different species of apes. While debate still rages regarding the appropriate way to interpret the symbolic productions of apes, three species of great apes (gorilla, orangutan, and chimpanzee) have now been credited with this capacity while no lesser apes or monkeys have been reported, at present, to have acquired such communicative skills. Among all of the claims made for the various animal species, the philosophers have entered the fray attempting to define the essence of what it is about language that makes it 'human'. This paper will compare and contrast the above positions to arrive at behavioural definitions of symbolic usage that can be applied across species. It will then present new data on a fourth ape species Pan paniscus which is proving to be the first non-human species to acquire symbolic skills in a spontaneous manner.

Animals↗

Age at first molar emergence in early Miocene Afropithecus turkanensis and life-history evolution in the Hominoidea.

Among primates, age at first molar emergence is correlated with a variety of life history traits. Age at first molar emergence can therefore be used to broadly infer the life histories of fossil primate species. One method of determining age at first molar emergence is to determine the age at death of fossil individuals that were in the process of erupting their first molars. This was done for an infant partial mandible of Afropithecus turkanensis (KNM-MO 26) from the approximately 17.5 Ma site of Moruorot in Kenya. A range of estimates of age at death was calculated for this individual using the permanent lateral incisor germ preserved in its crypt, by combining the number and periodicity of lateral enamel perikymata with estimates of the duration of cuspal enamel formation and the duration of the postnatal delay in the inception of crown mineralization. Perikymata periodicity was determined using daily cross striations between adjacent Retzius lines in thin sections of two A. turkanensis molars from the nearby site of Kalodirr. Based on the position of the KNM-MO 26 M(1)in relation to the mandibular alveolar margin, it had not yet undergone gingival emergence. The projected time to gingival emergence was estimated based on radiographic studies of M(1)eruption in extant baboons and chimpanzees. The estimates of age at M(1)emergence in KNM-MO 26 range from 28.2 to 43.5 months, using minimum and average values from extant great apes and humans for the estimated growth parameters. Even the absolute minimum value is well outside the ranges of extant large Old World monkeys for which there are data (12.5 to <25 months), but is within the range of chimpanzees (25.7 to 48.0 months). It is inferred, therefore, that A. turkanensis had a life history profile broadly like that of Pan. This is additional evidence to that provided by Sivapithecus parvada (Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptations, 1997, 173) that the prolonged life histories characteristic of extant apes were achieved early in the evolutionary history of the group. However, it is unclear at present whether life-history prolongation in apes represents the primitive catarrhine pace of life history extended through phyletic increase in body mass, or whether it is derived with respect to a primitive, size-adjusted life history that was broadly intermediate between those of extant hominoids and cercopithecoids. Life history evolution in primates as a whole may have occurred largely through a series of grade-shifts, with the establishment of fundamental life-history profiles early in the histories of major higher taxa. These may have included shifts that were largely body mass dependent, as well as those that occurred in the absence of significant changes in body mass.

Adaptation, Physiological↗