Search

The plantar aponeurosis in the human foot has been extensively studied and thoroughly described, in part, because of the incidence of plantar fasciitis in humans. It is commonly assumed that the human plantar aponeurosis is a unique adaptation to bipedalism that evolved in concert with the longitudinal arch. However, the comparative anatomy of the plantar aponeurosis is poorly known in most mammals, even among non-human primates, hindering efforts to understand its function. Here, we review previous anatomical descriptions of 40 primate species and use phylogenetic comparative methods to reconstruct the evolution of the plantar aponeurosis and its relationship to the plantaris muscle in primates. Ancestral state reconstructions suggest that the overall organization of the human plantar aponeurosis is shared with chimpanzees and that a similar anatomical configuration evolved independently in different primate clades as an adaptation to terrestrial locomotion. The presence of a plantar aponeurosis with clearly developed lateral and central bands in the African apes suggests that this structure is not prohibitive to suspensory locomotion and that these species possess versatile feet adapted for both terrestrial and arboreal locomotion. This plantar aponeurosis configuration would have been advantageous in enhancing foot stiffness for bipedal locomotion in the earliest hominins, prior to the evolution of a longitudinal arch. Hominins may have subsequently evolved thicker and stiffer plantar aponeuroses alongside the arch to enable a windlass mechanism and elastic energy storage for bipedal walking and running, although this idea requires further testing., (© 2020 The Authors. Journal of Anatomy published by John Wiley & Sons Ltd on behalf of Anatomical Society Anatomical Society.)

Meissner's corpuscles (MCs) are tactile mechanoreceptors found in the glabrous skin of primates, including fingertips. These receptors are characterized by sensitivity to light touch, and therefore might be associated with the evolution of manipulative abilities of the hands in primates. We examined MCs in different primate species, including common marmoset (Callithrix jacchus, n = 5), baboon (Papio anubis, n = 2), rhesus macaque (Macaca mulatta, n = 3), chimpanzee (Pan troglodytes, n = 3), bonobo (Pan paniscus, n = 1) and human (Homo sapiens, n = 8). Fingertips of the first, second and fourth digits were collected from both hands of specimens, dissected and histologically stained using hematoxylin and eosin. The density (MCs per 1 mm(2) ) and the size (cross-sectional diameter of MCs) were quantified. Overall, there were no differences in the densities of MCs or their size among the digits or between the hands for any species examined. However, MCs varied across species. We found a trend for higher densities of MCs in macaques and humans compared with chimpanzees and bonobos; moreover, apes had larger MCs than monkeys. We further examined whether the density or size of MCs varied as a function of body mass, measures of dexterity and dietary frugivory. Among these variables, only body size accounted for a significant amount of variation in the size of MCs., (© 2015 Anatomical Society.)

The pelvis is a critical link in the hindlimb locomotor system and has a central role in resisting loads associated with locomotion, but our understanding of its structural biomechanics is quite limited. Empirical data on how the pelvis responds to the loads it encounters are important for understanding pelvic adaptation to locomotion, and for testing hypotheses regarding how the pelvis is adapted to its mechanical demands. This paper presents in vitro strain gauge data on a sample of monkey and ape cadaveric specimens (Macaca, Papio, Ateles, Hylobates), and assesses strain magnitudes and distributions through the bones of the pelvis: the ilium, ischium and pubis. Pelves were individually mounted in a materials testing system, loads were applied across three hindlimb angular positions, and strains were recorded from 18 locations on the pelvic girdle. Peak principal strains range from 2000 to 3000 με, similar to peak strains recorded from other mammals in vivo. Although previous work has suggested that the bones of the pelvis may act as bent beams, this study suggests that there are likely additional loading regimes superimposed on bending. Specifically, these data suggest that the ilium is loaded in axial compression and torsion, the ischium in torsion, the pubic rami in mediolateral bending, and the pubic symphysis is loaded in a combination of compression and torsion. Compressive strains dominate the pelves of all species representatives. Shear strains change with limb position; hip flexion at 45° induces smaller shear strains than mid-stance (90°) or hip extension (105°). The pelvic girdle is a complex structure that does not lend itself easily to modeling, but finite element analyses may prove useful to generate and refine hypotheses of pelvic biomechanics., (© 2015 Anatomical Society.)

Here we provide the first report about the rates of muscle evolution derived from Bayesian and parsimony cladistic analyses of primate higher-level phylogeny, and compare these rates with published rates of molecular evolution. It is commonly accepted that there is a 'general molecular slow-down of hominoids', but interestingly the rates of muscle evolution in the nodes leading and within the hominoid clade are higher than those in the vast majority of other primate clades. The rate of muscle evolution at the node leading to Homo (1.77) is higher than that at the nodes leading to Pan (0.89) and particularly to Gorilla (0.28). Notably, the rates of muscle evolution at the major euarchontan and primate nodes are different, but within each major primate clade (Strepsirrhini, Platyrrhini, Cercopithecidae and Hominoidea) the rates at the various nodes, and particularly at the nodes leading to the higher groups (i.e. including more than one genera), are strikingly similar. We explore the implications of these new data for the tempo and mode of primate and human evolution., (© 2013 The Authors Journal of Anatomy © 2013 Anatomical Society.)

Shoulder morphology is functionally related to different patterns of locomotion in primates. To investigate this we performed a quantitative analysis of the relationship between cortical bone thickness (Cbt) of the muscle/tendon attachment site on the humerus and physiological cross-sectional area (PCSA) of the shoulder muscle in primates with different locomotory habits. The deltoid, subscapularis, supraspinatus, and infraspinatus were investigated. A chimpanzee, a gibbon, a baboon, two species of macaque, a lutong, a capuchin, and a squirrel monkey were included in the study. The total length of the humerus was measured and the values were converted into three-dimensional reconstructed data on a computer by computed tomography. The Cbt values were obtained from the volumes divided by the areas of the muscle/tendon attachment sites of the humerus by computer analysis. Muscle mass, muscle fascicle length, and muscle pennation angle were measured and PCSA was calculated using these parameters. A relatively high Cbt and small PCSA were characteristic of the gibbon. The gibbon's high Cbt suggests that passive tension in the muscle/tendon attachment site of suspensory primates (brachiators) may be greater than that of quadrupedal primates, whereas the relatively small PCSA indicates an association with a large amount of internal muscle fascia to endure the passive stress of brachiation. Although chimpanzees undertake some suspensory locomotion, the results for this species resemble those of the digitigrade/palmigrade quadrupedal primates rather than those of the suspensory primate. However, the deltoid and subscapularis in chimpanzee differ from those of the other primates and appear to be affected by the peculiar locomotion of knuckle-walking, i.e. the moment arm of forelimb in chimpanzees is relatively longer than that of digitigrade/palmigrade quadrupedal primates. Hence, a large PCSA in the deltoid and subscapularis may contribute to sustaining the body weight during locomotion. On the other hand, a thin cortical bone relative to a large PCSA was a feature of the cercopithecids, indicating that digitigrade/palmigrade quadrupedal locomotion produces less tension at the muscle/tendon attachment sites compared with that produced by brachiators., (© 2011 The Authors. Journal of Anatomy © 2011 Anatomical Society of Great Britain and Ireland.)

One of the most prominent features of the cranial component of the temporomandibular joint (TMJ) is the articular eminence (AE). This bar of bone is the primary surface upon which the condyle translates and rotates during movements of the mandible, and is therefore the primary point at which forces are transmitted from the mandible to the cranium during loading of the masticatory apparatus. The shape of the AE is highly variable across primates, and the raised eminence of humans has often been considered a defining feature of the human TMJ, yet few data exist to address whether this variation is functionally significant. This study used a broad interspecific sample of anthropoid primates to elaborate upon and test the predictions of a previously proposed model of AE function. This model suggests that AE inclination acts to resist non-normal forces at the TMJ, thereby maximizing bite forces (BFs). AE inclination was predicted to covary with two specific features of the masticatory apparatus: height of the TMJ above the occlusal plane; and inclination of the masticatory muscles. A correlate of this model is that taxa utilizing more resistant food objects should also exhibit relatively more inclined AEs. Results of the correlation analyses found that AE inclination is strongly correlated with height of the TMJ above the occlusal plane, but less so with inclination of the masticatory muscles. Furthermore, pairwise comparisons of closely related taxa with documented dietary differences found that the AE is consistently more inclined in taxa that utilize more resistant food items. These data preliminarily suggest that variation in AE morphology across anthropoid primates is functionally related to maximizing BFs, and add to the growing dataset of masticatory morphologies linked to feeding behavior., (© 2011 The Author. Journal of Anatomy © 2011 Anatomical Society of Great Britain and Ireland.)

Apart from molecular data, nearly all the evidence used to study primate relationships comes from hard tissues. Here, we provide details of the first parsimony and Bayesian cladistic analyses of the order Primates based exclusively on muscle data. The most parsimonious tree obtained from the cladistic analysis of 166 characters taken from the head, neck, pectoral and upper limb musculature is fully congruent with the most recent evolutionary molecular tree of Primates. That is, this tree recovers not only the relationships among the major groups of primates, i.e. Strepsirrhini {Tarsiiformes [Platyrrhini (Cercopithecidae, Hominoidea)]}, but it also recovers the relationships within each of these inclusive groups. Of the 301 character state changes occurring in this tree, ca. 30% are non-homoplasic evolutionary transitions; within the 220 changes that are unambiguously optimized in the tree, ca. 15% are reversions. The trees obtained by using characters derived from the muscles of the head and neck are more similar to the most recent evolutionary molecular tree than are the trees obtained by using characters derived from the pectoral and upper limb muscles. It was recently argued that since the Pan/Homo split, chimpanzees accumulated more phenotypic adaptations than humans, but our results indicate that modern humans accumulated more muscle character state changes than chimpanzees, and that both these taxa accumulated more changes than gorillas. This overview of the evolution of the primate head, neck, pectoral and upper limb musculature suggests that the only muscle groups for which modern humans have more muscles than most other extant primates are the muscles of the face, larynx and forearm., (© 2011 The Authors. Journal of Anatomy © 2011 Anatomical Society of Great Britain and Ireland.)

The morphology of postcranial articular surfaces is expected to reflect their weight-bearing properties, as well as the stability and mobility of the articulations to which they contribute. Previous studies have mainly confirmed earlier predictions of isometric scaling between articular surface areas and body mass; the exception to this is 'male-type', convex articular surface areas, which may scale allometrically due to differences in locomotor strategies within the analysed samples. In the present study, we used new surface scanning technology to quantify more accurately articular surface areas and to test those predictions within the talus of hominoid primates, including modern humans. Our results, contrary to predictions, suggest that there are no generalised rules of articular scaling within the talus of hominoids. Instead, we suggest that articular scaling patterns are highly context-specific, depending on the role of each articulation during locomotion, as well as taxon- and sex-specific differences in locomotion and ontogenetic growth trajectories within any given sample. While this may prove problematic for inferring body mass based on articular surface area, it also offers new opportunities of gaining substantial insights into the locomotor patterns of extinct species., (© 2011 The Authors. Journal of Anatomy © 2011 Anatomical Society of Great Britain and Ireland.)

Micro-computed tomographic analyses of trabecular bone architecture have been used to clarify the link between positional behavior and skeletal anatomy in primates. However, there are methodological decisions associated with quantifying and comparing trabecular anatomy across taxa that vary greatly in body size and morphology that can affect characterizations of trabecular architecture, such as choice of the volume of interest (VOI) size and location. The potential effects of these decisions may be amplified in small, irregular-shaped bones of the hands and feet that have more complex external morphology and more heterogeneous trabecular structure compared to, for example, the spherical epiphysis of the femoral head. In this study we investigate the effects of changes in VOI size and location on standard trabecular parameters in two bones of the hand, the capitate and third metacarpal, in a diverse sample of nonhuman primates that vary greatly in morphology, body mass and positional behavior. Results demonstrate that changes in VOI location and, to a lesser extent, changes in VOI size had a dramatic affect on many trabecular parameters, especially trabecular connectivity and structure (rods vs. plates), degree of anisotropy, and the primary orientation of the trabeculae. Although previous research has shown that some trabecular parameters are susceptible to slight variations in methodology (e.g. VOI location, scan resolution), this study provides a quantification of these effects in hand bones of a diverse sample of primates. An a priori understanding of the inherent biases created by the choice of VOI size and particularly location is critical to robust trabecular analysis and functional interpretation, especially in small bones with complex arthroses., (© 2010 The Authors. Journal of Anatomy © 2010 Anatomical Society of Great Britain and Ireland.)

The mammalian facial muscles are a subgroup of hyoid muscles (i.e. muscles innervated by cranial nerve VII). They are usually attached to freely movable skin and are responsible for facial expressions. In this study we provide an account of the origin, homologies and evolution of the primate facial muscles, based on dissections of various primate and non-primate taxa and a review of the literature. We provide data not previously reported, including photographs showing in detail the facial muscles of primates such as gibbons and orangutans. We show that the facial muscles usually present in strepsirhines are basically the same muscles that are present in non-primate mammals such as tree-shrews. The exceptions are that strepsirhines often have a muscle that is usually not differentiated in tree-shrews, the depressor supercilii, and lack two muscles that are usually differentiated in these mammals, the zygomatico-orbicularis and sphincter colli superficialis. Monkeys such as macaques usually lack two muscles that are often present in strepsirhines, the sphincter colli profundus and mandibulo-auricularis, but have some muscles that are usually absent as distinct structures in non-anthropoid primates, e.g. the levator labii superioris alaeque nasi, levator labii superioris, nasalis, depressor septi nasi, depressor anguli oris and depressor labii inferioris. In turn, macaques typically lack a risorius, auricularis anterior and temporoparietalis, which are found in hominoids such as humans, but have muscles that are usually not differentiated in members of some hominoid taxa, e.g. the platysma cervicale (usually not differentiated in orangutans, panins and humans) and auricularis posterior (usually not differentiated in orangutans). Based on our observations, comparisons and review of the literature, we propose a unifying, coherent nomenclature for the facial muscles of the Mammalia as a whole and provide a list of more than 300 synonyms that have been used in the literature to designate the facial muscles of primates and other mammals. A main advantage of this nomenclature is that it combines, and thus creates a bridge between, those names used by human anatomists and the names often employed in the literature dealing with non-human primates and non-primate mammals.

The ontogeny of the nasolacrimal ducts (NLD) and canals (NLC) are investigated in strepsirrhine and haplorhine primates. Developmental series of serially sectioned fetal, perinatal and adult specimens, in combination with juvenile and adult skulls subjected to high-resolution computed tomography, reveal that the vertical NLC and NLD of adult tarsiers and anthropoids are produced by the degeneration of a more horizontal anterior arm of the NLD that is present only transiently in haplorhines, but is maintained throughout life in strepsirrhines. This degeneration manifests as an 'unzipping' of the anterior arm by means of progressive enlargement (in a rostral direction) of a caudally placed opening of the NLD (at the base of the vertical NLC), followed by breakdown of the resulting epithelial groove. The similar mode by which the anterior arm of the membranous NLD degenerates in tarsiers and anthropoids strongly suggests that the conditions in these two taxa are homologous, and provides additional evidence for a monophyletic Haplorhini. The functional relationship between the nasolacrimal duct and the vomeronasal organ is reviewed in light of this evidence, and it is suggested that these changes in the haplorhine NLD were functionally linked to the development of anatomical haplorhinism of the oronasal complex.

As part of the hip joint, the proximal femur is an integral locomotor component. Although a link between locomotion and the morphology of some aspects of the proximal femur has been identified, inclusive shapes of this element have not been compared among behaviourally heterogeneous hominoids. Previous analyses have partitioned complex proximal femoral morphology into discrete features (e.g. head, neck, greater trochanter) to facilitate conventional linear measurements. In this study, three-dimensional geometric morphometrics are used to examine the shape of the proximal femur in hominoids to determine whether femoral shape co-varies with locomotor category. Fourteen landmarks are recorded on adult femora of Homo, Pan, Gorilla, Pongo and Hylobates. Generalized Procrustes analysis (GPA) is used to adjust for position, orientation and scale among landmark configurations. Principal components analysis is used to collapse and compare variation in residuals from GPA, and thin-plate spline analysis is used to visualize shape change among taxa. The results indicate that knucklewalking African apes are similar to one another in femoral shape, whereas the more suspensory Asian apes diverge from the African ape pattern. The shape of the human and orangutan proximal femur converge, a result that is best explained in terms of the distinct requirements for locomotion in each group. These findings suggest that the shape of the proximal femur is brought about primarily by locomotor behaviour.

Primates have shoulders adapted to a wide range of locomotor functions from terrestrial pronograde quadrupedalism to highly arboreal suspensory behaviours. The shape of the scapula tightly follows these functional differences. Previous analyses of primate postcrania, including the scapula, indicate that quadrupedal monkeys are less variable than non-quadrupeds. It was previously suggested that this difference was due to a relationship between the strength of stabilizing selection and the functional demands of the upper limb. Here it is shown that intraspecific scapular shape variance is highly correlated with the degree of committed quadrupedalism. Primates that engage in frequent suspensory behaviours (e.g. apes and ateline monkeys) average twice the amount of shape variance as quadrupeds (e.g. Old World monkeys and Saimiri). Because this difference in intraspecific shape variance is apparent in infants and does not increase or decrease appreciably over ontogeny, it is not likely that differences in postnatal growth, neuromuscular control or environmental factors such as habitat structure/composition are the primary contributors to differences in adult shape variance. Instead variance in embryonic factors that affect the shape/size of the scapula or epigenetic factors associated with muscle attachments are more likely candidates. In particular, the heterogeneous functional demands of the non-quadrupedal shoulder probably reduce the stringency of stabilizing selection, resulting in the persistence into adulthood of increased amounts of embryonically generated scapular shape variance.

A morphological study of the oblique ligament in the thumb is presented. The ligament was consistently described in human specimens and compared with dissections of non-human primates from different species. The oblique ligament was found in some, but not all, specimens in each of the following species examined: chimpanzee, orangutan, gibbon, anubis baboon, hamadryas baboon, squirrel monkey, lemur and marmoset. A revised identity of the oblique ligament is proposed as a reinforced distal border of a fibro-osseous annular pollical flexor sheath and whose function is not independent of the flexor sheath. The constant presence and tendinous trait of the pollical oblique ligament in humans, when compared with non-human primates, supports the notion that the oblique ligament strengthens the pollical flexor sheath in humans for restraint of the flexor pollicis longus tendon during forceful precision pinching. A derivation of the pollical oblique ligament is considered as representing a vestigial radial limb of a flexor pollicis superficialis tendon in the thumb.

A method is described for taking accurate measurements of the bony labyrinth of humans and other primates using high-resolution computed tomography (CT). The measurements comprise 8 dimensions, 14 orientations and 2 indices of the labyrinth, as well as 7 orientations of related structures of the petrous pyramid. Comparison of the measurements taken from CT scans with those taken from subsequently made casts and cryosections demonstrates that the method is sufficiently accurate to permit the morphometric analysis of labyrinthine size and shape. Since the CT method is nondestructive, fast and easy to perform, it is applicable to large samples and to rare or precious anthropological specimens.

Two arteries connect the anterior branch of the middle meningeal artery to the lacrimal artery. One vessel, the sphenoidal artery, passes through the superior orbital fissure. The other vessel, the meningolacrimal artery, passes through the cranio-orbital foramen. An analysis of data derived from embryology, comparative anatomy, and patterns of adult variation indicates that the meningolacrimal artery is homologous with the ramus superior of the stapedial artery, an embryologically and phylogenetically primitive vessel. The sphenoidal artery probably forms late in human ontogeny and represents an evolutionary novelty. Its distribution is probably restricted to hominoids and may be exclusive to humans and orangutans (Pongo pygmaeus). The sphenoidal artery is associated with three other derived features: the sphenoparietal sinus and sulcus and a laterally expanded superior orbital fissure. The high frequency of the sphenoidal artery in Homo and Pongo probably represents a case of parallelism. The earliest paleontological evidence of an enlarged sphenoidal artery in the Hominidae occurs in Indonesian Homo erectus (Sangiran VI).

The morphology of the lateral meniscus has been studied in a series of 316 non-human primates, representative of 43 genera. The lateral meniscus has a crescentic shape in Prosimii, in Platyrrhinii (New World monkeys) and in Pongo pygmaeus. The lateral meniscus is disc-shaped, with a central foramen, in Catarrhinii (Old World monkeys), in Hylobates, in Gorilla and in Pan Troglodytes. In man, the lateral meniscus has a crescentic shape. Discoid lateral menisci are reported as anomalies in man; their origin has given rise to much discussion, but the comparative data favour a phylogenetic origin. The posterior fasciculus (Wrisberg's ligament) of the posterior menisco-femoral ligament is always present and large in all non-human primates; in man, it may be absent and seems to be a regressive structure. On the other hand, the anterior fasciculus (Humphry's ligament) exists only in man and seems to be a progressive structure. The posterior menisco-tibial attachments are weak or non-existent in non-human primates, but they are well-developed in man. The evolutionary development of these characters can be related to human bipedal locomotion. The lateral meniscus in some Prosimii contains one or two intramensical ossicles (lunulae). These structures are absent or very rare in all other primate groups. In man, intrameniscal ossicles are extremely unusual; their origin, phylogenetic or post-traumatic is controversial, but comparative data favour, at least in some cases, the persistence of an ancestral character.

A vomeronasal organ and patent nasopalatine ducts were found in adult specimens of Aotus trivirgatus, Arctocebus calabarensis, Ateles geoffroyi and Saguinus fuscicollis. The vomeronasal complex appeared to be functional except in Ateles. However, the relative size of the organ in the Platyrrhini is much reduced compared to that in prosimians. The possible role of the organ in mediating neuroendocrine processes in primates is discussed.

Evidence is presented to suggest that the eutherian ankle joint has been derived from a meniscus-containing joint such as that found in extant arboreal marsupials. Probable morphological derivatives of this meniscus are identifiable in the Eutheria. The form and function of the joint are described in sub-human Primates and the adaptations which characterize the joint in bipedal man are noted. These morphological findings permit some speculation about the palaeocology of the earliest mammals with particular reference to the emergence of the order Primates.

A description is given of the functional morphology of the intrinsic joints of the foot, with particular reference to the primates, and an attempt is made to define the key human derived features. In the human subtalar joint the compromise axis has become reoriented into a very elevated position and more nearly in line with the functional foot axis than it is in subhuman primates. This provides for torque of the talus, and so the supported body weight, about the axis, during the stance phase of gait. In the subhuman primates the very oblique subtalar axis is important in inverting the foot into a grasping attitude. In subhuman primates the transverse tarsal joint complex plays an important role in supination of the forefoot, which complements inversion at the subtalar joint complex in achieving the ideal grasping position. This movement is only rendered possible by a helical action at the subtalar joint complex. In man the calcaneocuboid joint has been remodelled so as to bring the lamina pedis into a close-packed position during the latter part of the stance phase of gait. The joints between the cuneiforms and the cuboids, and the associated tarsometatarsal joints show significant modifications in man.

One hundred and thirteen forelimbs taken from 62 individuals belonging to 17 primate genera were dissected to reveal the entire course of the superficial venous system. The course of the deep venous system was also documented in at least one forelimb of each primate genus, and the number and location of perforating veins was recorded in 18 human and 45 non-human primate limbs. In Pan, Gorilla and in about 25% of human specimens the lateral superficial vein was confined to the forearm, while in all other primates, and in the majority of humans, this vein extended from the carpus to the clavicular region. Only Pongo and humans exhibited a second main superficial vein on the medial side of the forearm. In all primates the deep veins of the forelimb usually accompanied the arteries. Thus variation in the deep venous system reflected the different arterial patterns exhibited by these primates. The number of perforating veins in the forelimb was related to the length of the limb. Primate genera with longer forelimbs had more perforators, though not as many as would be expected if the number of perforators scaled linearly with limb length.

The sixth layer of the cerebral cortex has been studied by means of the Golgi method in Carnivora (dog and cat), Artiodactyla (cow and sheep), and Primates (human) brains; a basic structural uniformity being observed in all these species. Projection neurons of lamina VIa were large and medium sized pyramidal neurons (including atypical and multiapical), small pyramidal cells, and spinous multipolar neurons with long descending axons. Projection neurons of lamina VIb were medium sized pyramidal neurons and small pyramids, horizontal pyramids, inverted pyramidal cells, spinous multipolar neurons with long descending axons and large fusiform cells. Local circuit neurons of lamina VIa were Martinotti cells, basket neurons, neurogliaform cells, sparsely spinous neurons with whirled axons, spine-free multipolar cells and bipolar neurons. Local circuit neurons of lamina VIb were sparsely spinous and spine-free multipolar cells with short axons and bipolar neurons. Marked differences were observed between gyral, intermediate and fissural regions. Fusiform and bipolar neurons were vertically arranged in the former, but were tangentially orientated in intermediate and fissural regions; inverted pyramidal cells were present in the gyrus but horizontal pyramids were the respective cells in the intermediate and fissural zones. When compared with lissencephalic species, a great horizontal fibrillary system (which is vertically arranged in gyral regions) was observed in convoluted brains. Cells of origin were fusiform neurons, bipolar cells, horizontal and inverted pyramids and pyramidal neurons (the latter by means of horizontal axonal collaterals). The great development of this cortico-cortical association system in gyrencephalic species is considered to be a major step in neocortical evolution.

The examination of the caecum of two groups of cynomolgus and rhesus monkeys, two orang-utans and a chimpanzee, as well as an extensive review of the available literature, confirmed that the length of the caecum, relative to that of the colon, decreased as the position of the species in the primate scale rose. Although absent in prosimians and New World monkeys, there was evidence that the appendix vermiformis began to develop in certain Old World monkeys and became fully developed in the anthropoid apes, showing that, far from being a vestigial organ, it has actually developed progressively in primates.

Electron microscopy of the olfactory epithelium of two prosimian primates, the trees shrew and slow loris, and two simian primates, the macaque and gibbon, has shown that this epithelium consists of three cell types, receptor cells, supporting cells and basal cells, as in other mammals. Receptor cells were ciliated in all the animals investigated except the tree shrew, where, in addition to ciliated receptors, nonciliated receptors bearing only microvilli were occasionally present. Developing receptor cells containing numerous centrioles between nucleus and cell surface were occasionally observed and these cells had poorly developed olfactory knobs and few mitochondria. The olfactory epithelium was similar in morphology in all four species, except that supporting cells showed progressively more numerous, more slender and longer microvilli on their distal surface from tree shrew to slow loris to macaque to gibbon. This may imply a more discriminatory surface in the higher primates in view of the close relationship of these microvilli to the receptor cell surface.

A search for mast cells has been made in the brains of 18 mammalian species in 13 families in the orders Insectivora, Primates, Rodentia and Carnivora. In the larger animals, only the diencephalon and olfactory bulbs were examined. Mast cells were identified by virtue of their heparin-containing granules, which are stained by Alcian blue 8GX and, metachromatically, by toluidine blue 0. Within the cerebral parenchyma, mast cells were confined to the dorsal diencephalon of Erinaceus europaeus (hedgehog), Tupaia glis (tree-shrew) and Nycticebus coucang (slow loris). Some cells were next to capillaries; others were not. Mast cells were sometimes found, though rarely, in the intracerebral perivascular connective tissue leptomeninges and choroid plexuses of some of the other species examined. It is concluded that pericapillary cells (pericytes), which have been called mast cells by some investigators, are not in fact mast cells since there is no evidence for the presence of heparin. The functions of mast cells in the brain are unknown.

The os peroneum is found in only a few primate families and seems to be completely absent in the non-primate pentadactyl mammals, in the Prosimii and in the Platyrrhinii (New World monkeys). In the Cercopithecidae (Catarrhinii, Old World monkeys) and the Hylobatidae, the os peroneum is a coffee bean-shaped constant, large and regular bone. The lateral surface of the bone is convex in all directions and non-articular. The medial surface is covered with hyaline cartilage and articulates by means of a synovial joint with the corresponding facet of the cuboid bone. The histological structure and the mode of ossification of the os peroneum are identical to that of other short bones of the skeleton. The os peroneum of the Cercopithecidae and Hylobatidae is an example of a new skeletal element that has appeared in a tendon subject to unusual mechanical stress. In the case of the peroneus longus tendon the stress is due to repetitive friction because of the functional importance of this muscle in the adduction and pseudo-opposability of the hallux. This osseous element is genetically fixed and hereditarily transmitted. Its mode of appearance is analogous to that accepted for the origin of the patella. In the Pongidae, the os peroneum is absent or rare. In man, this bone is relatively infrequent (approx 20% of mature individuals) and its shape is most irregular. In this case, the os peroneum appears as a regressive form of the typical bone observed in the above families, which is in the process of disappearing. Besides fundamental genetical factors, this regression is probably in relation to the disappearance of the functional importance of the peroneus longus muscle to the loss of the hallux opposability. Thus the mechanical factors cannot be dissociated from the genetic and phylogenetic factors in explaining the appearance and the regression of the os peroneum.

The coraco-acromial ligament in man is a trait shared only with other hominoids (apes) among anthropoid primates. The associated form of the coracoid process and acromion, their lateral projections and the shape of the glenoid cavity likewise distinguish the Hominoidea. These anatomical features add credence to the view that the living hominoids share an independent ancestry within the Anthropoidea associated with the development of several unique locomotor/feeding adaptations.