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Mechanical properties of foods used in experimental studies of primate masticatory function.

In vivo studies of jaw-muscle behavior have been integral factors in the development of our current understanding of the primate masticatory apparatus. However, even though it has been shown that food textures and mechanical properties influence jaw-muscle activity during mastication, very little effort has been made to quantify the relationship between the elicited masticatory responses of the subject and the mechanical properties of the foods that are eaten. Recent work on human mastication highlights the importance of two mechanical properties-toughness and elastic modulus (i.e., stiffness)-for food breakdown during mastication. Here we provide data on the toughness and elastic modulus of the majority of foods used in experimental studies of the nonhuman primate masticatory apparatus. Food toughness ranges from approximately 56.97 Jm(-2) (apple pulp) to 4355.45 Jm(-2) (prune pit). The elastic modulus of the experimental foods ranges from 0.07 MPa for gummy bears to 346 MPa for popcorn kernels. These data can help researchers studying primate mastication select among several potential foods with broadly similar mechanical properties. Moreover, they provide a framework for understanding how jaw-muscle activity varies with food mechanical properties in these studies.

Animals↗

The neglected sense-olfaction in primate behavior, ecology, and evolution.

This special issue emerged from a symposium held during the 20th Congress of the International Primatological Society in Torino, Italy, in August 2004. The symposium brought together scientists studying several different aspects of olfaction in primates. The topics addressed ranged from the morphology and physiology of the sensory apparatus, the genetics and chemistry of olfactory signals and the use of such signals in primate communication, to a comparative analysis of the role of olfaction in neural evolution. The papers in this issue reflect a surge of interest in diverse aspects of olfaction-an interest that has been stimulated by the more rigorous theoretical approaches and new techniques that have recently become available. This introduction briefly reviews past research on primate olfaction, summarizes the scope of this special issue, and provides a somewhat speculative glimpse of the future.

Animal Communication↗

New perspectives on taste and primate evolution: the dichotomy in gustatory coding for perception of beneficent versus noxious substances as supported by correlations among human thresholds.

In various environments where primates are presently observed, as well as in forests and savannas which have been inhabited by australopithecines and early hominids, there are (or there have been presumably) categories of substances eliciting taste signals associated with stereotyped responses. Such is the case for various soluble sugars of fruits and nectars, attracting consumers, and for several plant compounds in which bitter or strongly astringent properties have a repulsive effect. The occurrence of such classes of tasty substances among natural products appears to be related to the evolutionary trends that shaped primate sensory perception (for detecting either beneficent or potentially noxious substances) in the context of a long history of coevolution between animals and plants. Here, we present original psychophysical data on humans (412 individuals aged 17-59 years) as an analogy with which to test recent evidence from electrophysiology in nonhuman primates (Hellekant et al. [1997] J. Neurophysiol. 77:978-993; Danilova et al. [1998] Ann. N.Y. Acad. Sci. 855:160-164) that taste fibers can be grouped into clusters of "best-responding fibers" with two more specific clusters, one for sugars and one for quinine and tannins. The collinearity found between human taste responses (recognition thresholds) for fructose and sucrose, as well as for quinine and tannins, is presented and discussed as another evidence of the two-direction evolutionary trend determining taste sensitivity. Salt perception appears to be totally independent of these trends. Accordingly, the appreciation of a salty taste seems to be a recent culturally learned response, and not a primary taste perception. The very existence of primary tastes is discussed in the context of evolutionary trends, past and present.

Adolescent↗

The functional adaptations of primate molar teeth.

Measurements were taken on the upper and lower molars of 37 species of primates and one tupaiid to assess the relative importance of shearing, crushing and grinding features. Significant correlations were found between pairs of allometrically standardized dimensions which measure the same molar function (shearing, crushing, or grinding). Correlations between pairs of dimensions which do not measure the same function are not significant. Second molar adaptations for shearing, crushing, and grinding, as well as the length of the second lower molar, and the total surface of the post-canine dentition are negatively allometric with respect to metabolic rate. Species which take different proportions of fruit, leaves, and insects in their diets have different molar structure. Frugivores have small teeth for their adult body size with poorly developed shearing, crushing, and grinding features on their molars. By contrast, leaf-eating species tend to have large teeth for their adult body size with well developed shearing, crushing, and grinding. The second molars of insectivorous species were found to parallel closely those of leaf-eating species. The two groups are clearly distinguishable from the former on the basis of body size alone: the smallest living primate leaf-eater is on order of magnitude larger than the largest living primate insectivore.

Adaptation, Physiological↗

Homologies of the anterior teeth in Indriiadae and a functional basis for dental reduction in primates.

In a recent paper Schwartz ('74) proposes revised homologies of the deciduous and permanent teeth in living lemuriform primates of the family Indriidae. However, new evidence provided by the deciduous dentition of Avahi suggests that the traditional interpretations are correct, specifically: (1) the lateral teeth in the dental scraper of Indriidae are homologous with the incisors of Lemuriadae and Lorisidae, not the canines; (2) the dental formula for the lower deciduous teeth of indriids is 2.1.3; (3) the dental formula for the lower permanent teeth of indriids is 2.0.2.3; and (4) decrease in number of incisors during primate evolution was usually in the sequence I3, then I2, then I1. It appears that dental reduction during primate evolution occurred at the ends of integrated incisor and cheek tooth units to minimize disruption of their functional integrity.

Animals↗

Allometry of primate hair density and the evolution of human hairlessness.

Allometric analyses of hair densities in 23 anthropoid primate taxa reveal that increasingly massive primates have systematically fewer hairs per equal unit of body surface. Considering the absence of effective sweating in monkeys and apes, the negative allometry of relative hair density may represent an architectural adaptation to thermal constraints imposed by the decreasing ratios of surface area to volume in progressively massive primates. Judging by estimates of body volume, denudation of the earliest hominids should have progressed to a considerable extent prior to their shift from a forest to a grassland habitat during the Pliocene. We propose that, lacking a reflective coat of hair, the exploitation of eccrine sweating emerged as the primary mechanism for adaptation to the increased heat leads of man's new environment and permitted further reduction of the remnant coat to its present vestigial condition.

Animals↗

Sexual dimorphism and allometry in primate ossa coxae.

Five measurements were taken on the ossa coxae of 454 adult primates representing Ceboidea, Cercopithecoidea and Hominoidea. Sex differences in these variables and their relationships to overall body size and sexual dimorphism were tested by means of Student's T-test and regression analysis. The study attempts to clarify the nature of primate pelvic sexual dimorphism, including allometric effects, and more specifically, test the assertion made by Mobb and Wood (1977) that sexual dimorphism in body size in not an important determinant in pelvic sex differences. Variables that contribute to the size of the birth canal tend to be larger in females than males in all taxa studied except two. In these, Hylobates and Alouatta, there were no significant differences between the sexes for any of the five variables. In general, sexual dimorphism in variables contributing to the size of the birth canal was correlated (r approximately or equal to 0.8) with sexual dimorphism in body size. Furthermore, the coefficients of allometry underlying pelvic sex differences were shown to be moderately correlated (r approximately or equal to 0.5) with sexual dimorphism in size. The influence of other adaptive factors on primate pelvic sexual dimorphism are also briefly discussed.

Animals↗

Primate cortical bone microstructure: relationship to locomotion.

Mechanical factors are implicated in the control of osteonal (Haversian) remodeling. Twenty primate species, representing a diverse range of locomotor behaviors, have been examined to determine whether the arrangement of secondary osteonal bone at femoral mid-diaphysis reflects behaviorally related biomechanical influences. The proportion of bone cortex occupied by secondary osteons (percent osteonal bone) is related to the biomechanical environment of these femora. Percent osteonal bone delineates groups which correspond to other skeletal and kinesiological categorizations of primate locomotion and distinguishes among arboreal quadrupedal, terrestrial quadrupedal, suspensory, and bipedal primates.

Animals↗

Stresses on the limbs of quadrupedal primates.

Data is presented from eight primates on the ground reaction forces on the limbs during locomotion. These subjects supported from 30 to 45% of their body weight on their forelimbs. Other quadrupedal mammals support 55-60% of their body weight on their forelimbs. The increase of peak vertical force with speed varies greatly between the subjects. The variation in weight supported by the forelimbs and the peak forces on the forelimbs is proposed to correlate with variation in locomotor adaptations. It is suggested that the occurrence of bipedalism in primates represents the extreme expression of the tendency in primates to reduce the compressive forces on their forelimbs.

Animals↗

Dental function and diet in the Carpolestidae (Primates, Plesiadapiformes).

Microwear analysis has long been used to infer dental function in primates; however, this is the first study to combine microwear and morphometric analyses to infer the dental function of the highly derived dentition of an early Tertiary primate. The Carpolestidae, a family of plesiadapiform primates, are characterized in part by a bladelike lower dentition (P4 and trigonid of M1). This study reveals a dual function of this dental complex. During the preparatory cycle, the tall, vertical enamel surfaces and broad basal lobes on P4 and trigonid of M1 probably functioned to wedge foods apart. This cycle is designated slicing-crushing and is uncommon among mammals, having been documented only for two extinct taxa: the multituberculates and the carpolestids. In addition to this special function, P4 and trigonid of M1 were used extensively with the molars in Phase I shearing. Facet analysis has revealed two new Phase I facets located on P3,4, whose formation is a result of normal Phase I movements and the derived morphology of these teeth. Slicing-crushing, used with Phase I shearing, would probably be most useful in processing food items of combined textures, particularly a soft interior covered by either a brittle or ductile coat, which are characteristic of an omnivorous diet composed of invertebrates, nuts, and seeds.

Animals↗

Polymorphism of the vitamin D binding protein (DBP) among primates: an evolutionary analysis.

The distribution of the DBP (vitamin D binding protein) polymorphism is now well characterized among human populations but for primates only limited results are known. The aim of this paper is to describe the electrophoretic polymorphism of this protein among various species. Using three different electrophoretic methods, we are able to detect an unknown polymorphism and to classify the different alleles observed. These results may be used to set an international nomenclature for further comparisons. The different electrophoretic mobilities between Old and New World Monkeys show that: 1) the Cercopithecoïdea are presenting the largest genetic heterogeneity; 2) the DBP among the Galago corresponds to the lowest isoelectric points observed among Primates; 3) during the evolution from nonhuman Primates to Man, the DBP is able to keep its affinity for vitamin D derivatives despite the occurrence of significant molecular modifications; 4) among Anthropoïdea, the electrophoretic patterns of DBP are very close to the human Gc1 proteins. These results show that evolution at the DBP level can be considered as a continuous mechanism of structural modifications. A significant transition occurs during the differentiation between Cercopithecoïdea and Anthropoïdea. It is not too speculative to consider that some electrophoretic forms detected among Gorilla, Pongo, or Pan may be identical to rare variants observed among humans.

Animals↗

Interspecific perspective on mechanical and nonmechanical models of primate circumorbital morphology.

Linear dimensions and angular orientations of the browridge, postorbital bar, and postorbital septum were obtained from a representative series of primates and compared with variables associated with several nonmechanical and biomechanical/mechanical models put forward to explain the form and function of the circumorbital region. Analyses of the results indicate that face size is the primary determinant of variation in primate circumorbital morphology. Anteroposterior browridge thickness is correlated with neural-orbital disjunction among anthropoid primates, but not among prosimians. This difference appears related to differences in the construction of the upper face and anterior cranial fossa between prosimians and anthropoids. Little support is demonstrated for the anterior dental loading model of browridge development. Mediolateral postorbital bar width and (to a lesser degree) browridge height are correlated with neurofacial torsion during mastication and variation in masticatory muscle size. These analyses further suggest that since circumorbital structures (especially the browridges) are located the farthest away from the chewing apparatus, they are least affected by masticatory stresses.

Animals↗

Osteoarthritis, calcium pyrophosphate deposition disease, and osseous infection in Old World primates.

Uncertainties as to the nature and implications of osteoarthritis and calcium pyrophosphate deposition disease (CPPD) in primates were subject to critical review through examination of 153 prosimians and 1,250 Old World non-prosimian primates. Osteoarthritis, calcium pyrophosphate deposition disease, and infectious arthritis/osteomyelitis were relatively rare phenomena, affecting only 1.7% and 2.5%, respectively, of free-ranging prosimians and other Old World non-prosimian primates. Frequency of infection in Indri and Presbytis appears to reflect a unique susceptibility or exposure. Papio may have a unique predisposition to CPPD. The dichotomy (frequency and joint distribution) between free-ranging and artificially restrained animals suggests caution in interpretation of osteoarthritis or CPPD in non-free-ranging animals.

Animals↗

New characters for the functional interpretation of primate scapulae and proximal humeri.

The study of muscle function in nonhuman primates through the technique of electromyography (EMG) has facilitated the identification of specific functional roles for muscles in particular behaviors. This has led to a more complete understanding of the biomechanics of certain regions of the musculoskeletal system, and should facilitate our ability to identify morphological features useful in the functional interpretation of fossil material. The current paper represents one such investigation of a new set of morphometric characters of the scapula and proximal humerus suggested by EMG analyses of shoulder muscle function. A set of new metric variables were examined on the scapulae and proximal humeri of 25 species of extant anthropoid primates, as well as on casts of scapulae and humeri of three fossil primate taxa. The variables are primarily related to the line of action and attachments of the rotator cuff muscles. The position of the scapular spine, the degree of lateral expansion of the subscapular fossa, the size and shape of the subscapularis insertion facet on the lesser tubercle, and the orientation of the infraspinatus insertion facet on the greater tubercle all appear to successfully sort the extant taxa into locomotor groups. Their appearance on the fossil specimens generally supports previous functional interpretations of each taxon's locomotor abilities based on a variety of other characters, suggesting that these traits are equally applicable to fossil material.

Animals↗

Phylogenetic analyses of dimorphism in primates: evidence for stronger selection on canine size than on body size.

Phylogenetic comparative methods were used to analyze the consequences of sexual selection on canine size and canine size dimorphism in primates. Our analyses of previously published body mass and canine size data revealed that the degree of sexual selection is correlated with canine size dimorphism, as well as with canine size in both sexes, in haplorhine but not in strepsirrhine primates. Consistent with these results, male and female canine size was found to be highly correlated in all primates. Since canine dimorphism and canine size in both sexes in haplorhines were found to be not only related to mating system but also to body size and body size dimorphism (characters which are also subject to or the result of sexual selection), it was not apparent whether the degree of canine dimorphism is the result of sexual selection on canine size itself, or whether canine dimorphism is instead a consequence of selection on body size, or vice versa. To distinguish among these possibilities, we conducted matched-pairs analyses on canine size after correcting for the effects of body size. These tests revealed significant effects of sexual selection on relative canine size, indicating that canine size is more important in haplorhine male-male competition than body size. Further analyses showed, however, that it was not possible to detect any evolutionary lag between canine size and body size, or between canine size dimorphism and body size dimorphism. Additional support for the notion of special selection on canine size consisted of allometric relationships in haplorhines between canine size and canine size dimorphism in males, as well as between canine size dimorphism and body size dimorphism. In conclusion, these analyses revealed that the effects of sexual selection on canine size are stronger than those on body size, perhaps indicating that canines are more important than body size in haplorhine male-male competition.

Animals↗

Evolution of somatosensory and motor cortex in primates.

Inferences about how the complex somatosensory systems of anthropoid primates evolved are based on comparative studies of such systems in extant mammals. Experimental studies of members of the major clades of extant mammals suggest that somatosensory cortex of early mammals consisted of only a few areas, including a primary area, S1, bordered by strip-like rostral and caudal somatosensory fields, SR and SC. In addition, the second somatosensory area, S2, and the parietal ventral area, PV, were probably present. S1, S2, and PV were activated independently via parallel projections from the ventroposterior nucleus, VP. Little posterior parietal cortex existed, and it was unlikely that a separate primary motor area, M1, existed until placental mammals evolved. Early primates retained this basic organization and also had a larger posterior parietal region that mediated sensorimotor functions via connections with motor and premotor areas. The frontal cortex included M1, dorsal and ventral premotor areas, supplementary motor area, and cingulate motor fields. Ventroposterior superior and ventroposterior inferior nuclei were distinct from the ventroposterior nucleus in the thalamus. In early anthropoid primates, areas S1, SR, and SC had differentiated into the fields now recognized as areas 3b, 3a, and 1. Areas 3b and 1 contained parallel mirror-image representations of cutaneous receptors and a parallel representation in area 2 was probable. Serial processing became dominant, so that neurons in areas 1, S2, and PV became dependent on area 3b for activation. Posterior parietal cortex expanded into more areas that related to frontal cortex. Less is known about changes that might have occurred with the emergence of apes and humans, but their brains were larger and posed scaling problems most likely solved by increasing the number of cortical areas and reducing the proportion of long connections.

Adaptation, Physiological↗

Convergence of sensory systems in the orbitofrontal cortex in primates and brain design for emotion.

In primates, stimuli to sensory systems influence motivational and emotional behavior via neural relays to the orbitofrontal cortex. This article reviews studies on the effects of stimuli from multiple sensory modalities on the brain of humans and some other higher primates. The primate orbitofrontal cortex contains the secondary taste cortex, in which the reward value of taste is represented. It also contains the secondary and tertiary olfactory cortical areas, in which information about the identity and also about the reward value of odors is represented. A somatosensory input is revealed by neurons that respond to the viscosity of food in the mouth, to the texture (mouth feel) of fat in the mouth, and to the temperature of liquids placed into the mouth. The orbitofrontal cortex also receives information about the sight of objects from the temporal lobe cortical visual areas. Information about each of these modalities is represented separately by different neurons, but in addition, other neurons show convergence between different types of sensory input. This convergence occurs by associative learning between the visual or olfactory input and the taste. In that emotions can be defined as states elicited by reinforcers, the neurons that respond to primary reinforcers (such as taste and touch), as well as learn associations to visual and olfactory stimuli that become secondary reinforcers, provide a basis for understanding the functions of the orbitofrontal cortex in emotion. In complementary neuroimaging studies in humans, it is being found that areas of the orbitofrontal cortex are activated by pleasant touch, by painful touch, by taste, by smell, and by more abstract reinforcers such as winning or losing money. Damage to the orbitofrontal cortex in humans can impair the learning and reversal of stimulus-reinforcement associations and thus the correction of behavioral responses when these are no longer appropriate because previous reinforcement contingencies change. It is striking that humans and other catarrhines, being visual specialists like other anthropoids, interface the visual system to other sensory systems (e.g., taste and smell) in the orbitofrontal cortex.

Afferent Pathways↗

Comparative anatomy of the facial motor nucleus in mammals, with an analysis of neuron numbers in primates.

The facial motor nucleus (VII) contains motoneurons that innervate the facial muscles of expression. In this review, the comparative anatomy of this brainstem nucleus is examined. Several aspects of the anatomical organization of the VII appear to be common across mammals, such as the distribution of neuron types, general topography of muscle representation, and afferent connections from the midbrain and brainstem. Phylogenetic specializations are apparent in the proportion of neurons allocated to the representation of subsets of muscles and the degree of differentiation among subnuclei. These interspecific differences may be related to the elaboration of certain facial muscles in the context of socioecological adaptations such as whisking behavior, sound localization, vocalization, and facial expression. Furthermore, current evidence indicates that direct descending corticomotoneuron projections in the VII are present only in catarrhine primates, suggesting that this connectivity is an important substrate for the evolution of enhanced mobility and flexibility in facial expression. Data are also presented from a stereologic analysis of VII neuron numbers in 18 primate species and a scandentian. Using phylogenetic comparative statistics, it is shown that there is not a correlation between group size and VII neuron number (adjusted for medulla volume) among primates. Great apes and humans, however, display moderately more VII neurons that expected for their medulla size.

Anatomy, Comparative↗