Comparative postcranial body shape and locomotion in Chlorocebus aethiops and Cercopithecus mitis.

Body weight and length, chest girth, and seven postcranial limb segment lengths are compared between two guenon species, Chlorocebus (Cercopithecus) aethiops (vervets) and Cercopithecus mitis (blue monkeys), exhibiting different habitual locomotor preferences. The subjects, all adults, were wild caught for a non-related research project (Turner et al. [1986] Genetic and morphological studies on two species of Kenyan monkeys, C. aethiops and C. mitis. In: Else JG, Lee PC, editors. Primate evolution, proceedings of the Xth International Congress of Primatology, Cambridge. London). The morphological results are interpreted within the context of previously published observations of primate locomotion and social organization. The sample is unique in that the body weight of each individual is known, allowing the effects of body-size scaling to be assessed in interspecific and intersexual comparisons. C. mitis has a significantly (P < 0.05) greater body weight and trunk length than C. aethiops. A shorter trunk may function to reduce spinal flexibility for ground-running in the latter. Proximal limb segments (arm and thigh) are significantly greater in C. mitis, reflecting known adaptations to committed arboreal quadrupedal locomotion. By contrast, relative distal limb segments (forearm, crus, and foot) are significantly longer in C. aethiops, concordant with a locomotor repertoire that includes substantial terrestrial quadrupedalism, in addition to arboreal agility, and also the requisite transition between ground and canopy. Although normally associated with arboreal monkeys, greater relative tail length occurs in the more terrestrial vervets. However, because vervets exploit both arboreal and terrestrial habitats, a longer tail may compensate for diminished balance during arboreal quadrupedalism resulting from the greater "brachial" and "crural" indices that enhance their ground quadrupedalism. Most interspecific differences in body proportions are explicable by differences in locomotor modalities. Some results, however, contradict commonly held "tenets" that relate body size and morphology exclusively to locomotion. Generally associated with terrestriality, sexual dimorphism (male/female) is greater in the more arboreal blue monkeys. A more intense, seasonal mating competition may account for this incongruity.

Ontogeny of histochemical fiber types and muscle function in the masseter muscle of miniature swine.

In this study of masticatory maturation, the ontogeny of the histochemical fiber type composition of musculus masseter is examined in the omnivorous miniature swine (Sus scrofa). Fiber type characteristics are interpreted by comparison with electromyography (EMG) recorded during feeding behavior. Similar to locomotion studies, the results suggest a correspondence between the composition and arrangement of motor units and their recruitment pattern. Serial sections of masseter muscles from 10 minipigs, ranging from 2 weeks to slightly over 1 year of age, were stained for myosin adenosine triphosphatase (mATPase) activity to distinguish slow-twitch from fast-twitch fibers, and for nicotinamide adenosine dehydrogenase-tetrazolium reductase to assess the aerobic capacity of the same fibers. Although maintaining a uniformly high aerobic capacity throughout ontogeny and in adult animals, a transition is observed in the relative proportions of fast- and slow-twitch fibers. The primarily fast-twitch neonatal pig masseter eventually comprises approximately 25-30% slow-twitch fibers in adults, with a higher predominance of slow fibers in the deep (vs. superficial) and anterior (vs. posterior) regions of the muscle. Furthermore, while individual fibers of adult masseters generally stain for either alkaline- or acid-stable mATPase activity, a substantial proportion of cells in developing animals exhibits the presence of both isozymes. EMG results indicate functional heterogeneity within the masseter of adult pigs. During chewing, when pig chow is replaced by cracked corn, EMG activity in the deep portion of the muscle either decreases or increases slightly. In the superficial portion, however, muscle amplitudes become dramatically higher for corn, surpassing levels generated for chewing the less obdurate chow. These results are consistent with a behavioral transition from neonatal suckling to sustained mastication of foods of more complex textures eaten by adult pigs. The relationship between these fiber type and EMG results for pig masseter corresponds to those pertaining to motor unit recruitment in the extensor muscles of locomotion. Implications of this work for the evolutionary morphology of mastication also are discussed.

Growth, development, and sexual dimorphism in vervet monkeys (Cercopithecus aethiops) at four sites in Kenya.

Body weight and ten body segment measurements were collected from 367 wild-trapped vervet monkeys (Cercopithecus aethiops) in central and southern Kenya. The animals represent between 70 and 95% of the animals in each of 30 troops at four geographical locations separated by 80 to 380 km. The capture sites differed in altitude, mean annual rainfall and temperature. Two questions are addressed: (1) what are the differences in male and female growth patterns, and (2) what is the relationship between size, climate, and availability of food? Each animal was assigned to an age class based on dental examination. Means for all variables do not diverge for males and females from birth to age class 4 (15-18 months). After this, male and female growth rates diverge. This sexual dimorphism in growth pattern may reflect timing of entry into the reproductive community. A nested analysis of variance (ANOVA) was employed to compare sites, groups within sites and individuals within groups. Statistically significant differences between sites in body weight and body segment measurements are found for adult females. Except for tail length, these differences do not follow Bergmann's or Allen's Rules correlating size differences and temperature, but rather may reflect proximity to cultivated areas or tourist lodges with greater access to human food.

Interlimb coordination, gait, and neural control of quadrupedalism in chimpanzees.

Interlimb coordination is directly relevant to the understanding of the neural control of locomotion, but few studies addressing this topic for nonhuman primates are available, and no data exist for any hominoid other than humans. As a follow-up to Jungers and Anapol's ([1985] Am. J. Phys. Anthropol. 67:89-97) analysis on a lemur and talapoin monkey, we describe here the patterns of interlimb coordination in two chimpanzees as revealed by electromyography. Like the lemur and talapoin monkey, ipsilateral limb coupling in chimpanzees is characterized by variability about preferred modes within individual gaits. During symmetrical gaits, limb coupling patterns in the chimpanzee are also influenced by kinematic differences in hindlimb placement ("overstriding"). These observations reflect the neurological constraints placed on locomotion but also emphasize the overall flexibility of locomotor neural mechanisms. Interlimb coordination patterns are also species-specific, exhibiting significant differences among primate taxa and between primates and cats. Interspecific differences may be suggestive of phylogenetic divergence in the basic mechanisms for neural control of locomotion, but do not preclude morphological explanations for observed differences in interlimb coordination across species.

Fiber architecture of the extensors of the hindlimb in semiterrestrial and arboreal guenons.

Fiber architecture of the extensor musculature of the knee and ankle is examined in two African gueon species--the semiterrestrial Cercopithecus aethiops, and the arboreal C. ascanius. Using histologic and microscopic techniques to measure lengths of sarcomeres, the original lengths of muscle fasciculi and angles of pinnation in quadriceps femoris and triceps surae are reconstructed from direct measurements on cadavers. Calculations of reduced physiological cross-sectional area, mass/predicted effective tetanic tension, maximum excursion, and tendon length/fasciculus+tendon lengths are correlated to preferred locomotor modalities in the wild. For both species, greater morphological differences occur among the bellies of quadriceps femoris--rectus femoris, vastus intermedius, v. lateralis, and v. medialis--than among the bellies of triceps surae--gastrocnemius lateralis, g. medialis, plantaris, and soleus. With regard to quadriceps femoris, few differences occur between species. Interspecific differences in the triceps surae indicate (1) redirection of muscle force to accommodate arboreality in which the substrate is less than body width; (2) muscles more suited for velocity in the semiterrestrial vervets; and (3) muscles used more isotonically in vervets and more isometrically in red-tailed monkeys. The inherent flexibility of muscles may be preadaptive to a primary species shift in locomotor modality until the bony morphology is able to adapt through natural selection.

Morphological adaptation to diet in platyrrhine primates.

Morphological features of the jaws and teeth are examined in eight species of platyrrhine monkeys that coexist in the Suriname rainforest. Z-scores calculated from geometric predictions for several features of the feeding apparatus thought to have some functional significance (e.g., tooth dimensions, jaw robusticity, leverage of primary jaw elevators) are compared to a profile of the naturalistic dietary behavior of these species (i.e., proportions of fruit mesocarp, seeds, leaves, and fauna eaten). Several features are found exclusively in those platyrrhines whose dietary preferences are the most limited. Such specializations appear to be associated with a particular protein source exploited by a species to supplement a largely frugivorous diet. Ateles paniscus, which feeds primarily on the mesocarp of ripe fruit, has an adaptive morphology that emphasizes broad incisors. Chiropotes satanas (and to a slightly lesser extent, Pithecia pithecia) is a frugivore/seed predator with large upper and lower canines and a robust mandible. The frugivore/folivore Alouatta seniculus has a relatively large total molar area and effective mandibular condyle height. In all four of these strictly vegetarian species, the leverage of the masseter muscle is greater than that of temporalis. Of the omnivorous species, Cebus apella and C. nigrivittatus exploit both fauna and seeds for protein and exhibit an array of many of the above features, such as large teeth and thick mandibles. Saimiri sciureus, not particularly known for seed predation, departs from Cebus in having less robust canines and a more gracile mandible. All three cebid omnivores have a temporalis with greater leverage than the masseter, indicating a requirement for resisting anteriorly directed forces, for example, using the jaws for vigorous foraging. The lack of any enlarged features, other than incisors, in the omnivorous Saguinus midas may be attributable to the functional constraints of small body size. Because the small size of the gape limits the size of the food parcel ingested, a requirement to enlarge other dentomandibular structures for trituration is alleviated.

Motor-unit territories in the masseter muscle of infant pigs.

In adult miniature pigs (Sus scrofa) fibres of the masseter contract differentially, helping to produce the successive movements of the mandible during the chewing cycle. In infant pigs, however, most fibres of the masseter contract simultaneously. One hypothesis to explain the ontogenetic change in contraction pattern is that the infant masseter is neurologically immature, with large overlapping motor units incapable of producing differential contractions. This hypothesis was tested by mapping the territories of motor units in the masseters of piglets. Filaments from the masseteric nerve were stimulated repetitively; muscle fibres belonging to the stimulated motor units were identified by their failure to react for glycogen. Just as in older pigs, motor-unit territories were found to be very restricted, occupying only a small portion of total muscle volume. Thus, neural organization does not appear to be immature in piglets. An alternative hypothesis, that the ontogenetic change in activity pattern results from growth changes in the anatomy of the masseter, may be a more likely explanation.

Length-tension relationships of masseter and digastric muscles of miniature swine during ontogeny.

At incremental whole muscle lengths, active isometric and passive elastic forces were recorded from the masseter and digastric muscles of anaesthetized miniature pigs (Hanford) weighing 2.0-20.0 kg. Wet muscle mass and maximum tetanic tension values for masseter exceed those for digastric and increase more rapidly with body mass (age). At any body mass, masseter exceeds digastric in the ratio of optimum length (that length at which maximum tetanic tension is produced) to in situ muscle length (that length which corresponds to the jaw in a closed position) and the proportion of passive tension comprising total (passive plus active) tension. Passive elastic tension begins to rise in masseter at lengths as short as 87% of optimum (in younger pigs). In digastric, passive tension is absent until the muscle is stretched to a length slightly longer than optimum in younger pigs but occurs at shorter lengths in older pigs. Contractile properties explain functional differences between masseter and digastric more clearly than they explain ontogenetic changes in either muscle. The behavioural transition from infant suckling to adult mastication of solid food is best characterized by a disproportionate increase in mass (and force) of the masseter, relative to digastric, and increased reliance upon active (rather than passive) tension.

Neural organization of the masseter muscle in the pig.

The neural organization of the pig masseter, an architecturally and functionally compartmentalized muscle, was investigated by using dissection, glycogen depletion, evoked electromyography, and counts of axon numbers at various levels along the masseteric nerve. The masseteric nerve enters the muscle as two rostral branches, which also supply the zygomatico-mandibularis, and a more caudal main branch, which soon divides into four terminal nerves with variable distributions. Stimulation of filaments containing roughly 50 extrafusal motor axons resulted in glycogen depletion of 5-20% of the muscle fibers in very small subvolumes of the masseter; the affected subvolumes were delimited by perimysium. Electromyography after stimulation of various branches of the nerve confirmed the distributions deduced from anatomy and further indicated that axons do not branch between the rostral and main nerve branches but may occasionally do so among the more distal terminal branches of the main branch. The proximal trunk of the masseteric nerve contains about 3,500 myelinated fibers with a bimodal size distribution. Approximately 1,000 of the larger fibers were estimated to be extrafusal motor axons. Along the proximal trunk of the nerve, fibers were constantly rearranged; coupled with the observation that the locations of motor unit territories were usually not related to the position of the stimulated axons within the nerve, this suggests that the nerve trunk is not strictly ordered somatotopically.

Fossil platyrrhine forelimb bones from the early miocene of Argentina.

Primate scapula and ulna fragments of uncertain taxonomic affinity (MACN-SC 101) have been recovered from the Pinturas deposits at Arroyo Feo, Santa Cruz, Argentina in association with Santacrucian (Early Miocene) land mammals. Least-squares regression of body weight on surface area and on height of the glenoid fossa of the scapula indicates an estimated mean weight of 3.6 kg for this individual. On the basis of qualitative and several metric features, the fossil scapula and ulna most closely resemble living platyrrhine monkeys. In estimated body weight and relative height of the coronoid process, the fossil is similar to arboreal quadrupeds, such as Cebus apella and Chiropotes. However, spinoglenoid, axilloglenoid, and axillospinal angles, length of lever arm, and length and breadth of the sigmoid notch imply behavioral similarity with larger species that also use their forelimbs extensively in climbing, such as Alouatta and Lagothrix. MACN-SC 101 may represent the incipient divergence of a generalized platyrrhine arboreal quadruped toward a more suspensory form.

Morphological and videofluorographic study of the hyoid apparatus and its function in the rabbit (Oryctolagus cuniculus).

The anatomy of the hyoid apparatus and positional changes of the hyoid bone during mastication and deglutition are described in the New Zealand White rabbit (Oryctolagus cuniculus). A testable model is constructed to predict the range of movement during function of the hyoid, a bone entirely suspended by soft tissue. Frame-by-frame analysis of a videofluorographic tape confirms the accuracy of the prediction through observation of hyoid bone excursion during oral behavior. During chewing, translation of the hyoid bone is diminutive and irregular, lacking a clearly discernible path of excursion. However, some movements of the hyoid occur with regularity. During fast opening, anterodorsal movement of the hyoid is interrupted with an abrupt posteroventral depression when the bolus is moved posteriorly toward the cheek teeth by the tongue. This clockwise rotation (when viewed from the right side) of the hyoid accompanies jaw opening and is reversed (posteroventral movement) for the jaw closing sequence. Lateral movements of the hyoid may be slightly coupled to mandibular rotation in the horizontal plane. The findings suggest that the hyoid bone maintains a relatively static position during the dynamics of chewing. The primary function would be to provide a stable base for the movements of the tongue. Another possible function would be to control the position of the larynx within the pharyngeal cavity. Some characteristic features of the rabbit hyoid apparatus may be consequential to relatively erect posture and a saltatory mode of locomotion.

Telemetered electromyography of the fast and slow extensors of the leg of the brown lemur (Lemur fulvus).

Quantitative telemetered electromyography from the four heads of the quadriceps femoris (vastus medialis, rectus femoris, vastus intermedius and vastus lateralis) during normal postures and locomotion (quadrupedal resting, walk/run, gallop and leaping) is presented for the brown lemur, Lemur fulvus. The vastus intermedius is the sole contributor to muscle recruitment during quadrupedal resting postures. It exhibits consistently high levels of electrical activity during all locomotory behaviour. The vastus medialis is recruited least among these muscles during walk/run and tends to be 'saved' for galloping and leaping. The rectus femoris and vastus lateralis are recruited at similarly high levels during all phasic activities. The rectus femoris appears to be used eccentrically, storing 'elastic strain energy' during all phasic activities. The vastus lateralis combines exceptionally high potential effective force with relatively high potential velocity and resistance to fatigue and probably develops the majority of force in all phasic activities. These results support previously documented structural and histochemical data that imply a functional division of labour among these muscle synergists.

The force-velocity relation of the rabbit digastric muscle.

In 30 animals, the digastric was made to pull actively against a slide loaded by a servo-controlled linear motor. Force and velocity were recorded at the end of active shortening to the in-situ (jaw-closed) muscle length. Passive and active force-length relations were also determined in 17 of the rabbits. The empirical force-velocity data were fitted to a hyperbolic equation. The average speed of muscle shortening at zero load was 14.67 cm/s. Mean maximum isometric force at in-situ length (P0) was 1267 g, and the mean ratio a/P0 was 0.18. The average time-to-peak twitch tension was 31.8 ms under isometric conditions. In-situ muscle-belly length was about 3 per cent less than optimum length for isometric force. Maximum muscle force was positively correlated with animal size, but maximum velocity showed no relation to force or length. The estimated maximum speed of sarcomere shortening was 26 micron/s, which is slightly slower than in fast limb muscles of the cat, and may indicate the presence of both histochemical type I and II fibres. The isometric force after shortening had ceased was less than P0, and was correlated with the velocity during shortening. This depression of isometric force may result from an alteration of the excitation-coupling system during activation. These observations suggest a role for the digastric in the rapid acceleration and deceleration of the mandible near the jaw-closed position during opening and closing.

Architectural and histochemical diversity within the quadriceps femoris of the brown lemur (Lemur fulvus).

Physiologically related features of muscle morphology are considered with regard to functional adaptation for locomotor and postural behavior in the brown lemur (Lemur fulvus). Reduced physiological cross-sectional area, estimated maximum excursion of the tendon of insertion, length of tendon per muscle fasciculus, and areal fiber type composition were examined in the quadriceps femoris in order to assess the extent of a "division of labor" among four apparent synergists. Each of these four muscles in this prosimian primate displays a distinguishing constellation of morphological features that implies functional specialization during posture and normal locomotion (walk/run, galloping, leaping). Vastus medialis is best suited for rapid whole muscle recruitment and may be reserved for relatively vigorous activities such as galloping and leaping (e.g., small cross-sectional area per mass, long excursion, predominance of fast-low oxidative fibers, relatively little tendon per fasciculus). In theory, rectus femoris could be employed isometrically in order to store elastic strain energy during all phasic activities (e.g., large cross-sectional area per mass, short excursion, predominance of fast-high oxidative fibers, large amount of tendon per fasciculus). Vastus intermedius exhibits an overall morphology indicative of a typical postural muscle (e.g., substantial cross-sectional area, short excursion, predominance of slow-high oxidative fibers, large amount of tendon per fasciculus). The construction of vastus lateralis reflects an adaptation for high force, relatively high velocity, and resistance to fatigue (e.g., large cross-sectional area, long excursion, most heterogeneous distribution of fiber types, large amount of tendon per fasciculus); this muscle is probably the primary contributor to a wide range of locomotor behaviors in lemurs. Marked dramatic architectural disparity among the four bellies, coupled with relative overall fiber type heterogeneity, suggests the potential for exceptional flexibility in muscle recruitment within this mass. One interpretation of this relatively complex neuromuscular organization in the brown lemur is that it represents an adaptation for the exploitation of a three-dimensional arboreal environment by rapid quadrupedalism and leaping among irregular and spatially disordered substrates.

Limulus striated muscle provides an unusual model for muscle contraction.

Although nearly two decades have passed since de Villafranca (1961) described A-band shortening, controversy persists. Here we will review the data which has been amassed since de Villafranca 's description. We will conclude that A-bands and thick filaments shorten during sarcomere shortening in Limulus striated muscle. Further we will suggest that two machines operate in this muscle: a tension generating sliding filament system and a tension generating thick filament shortening system. Also we will suggest a mechanism of force generation of the filament shortening system and provide evidence for a cycling bridge mechanism for this muscle.

Scapula of Apidium phiomense: a small anthropoid from the Oligocene of Egypt.

A scapular fragment from the Upper Fossil Wood Zone, Quarry I, Fayum Depression, Egypt, is assigned to the taxon Apidium phiomense. This animal may have approximated the size of Galago crassicaudatus or Nycticebus coucang, as predicted by regressing body weight on glenoid surface area and fossa length for an assortment of living primates. Morphometric values for spinoglenoid, axilloglenoid, and axillospinal angles indicate locomotor affinities of A. phiomense with colobine monkeys. Other nonmetric traits align the fossil with Saimiri sciureus, probably its closest behavioral analogue. A. phiomense is thus characterized as an arboreal quadruped with a locomotor repertoire that includes a considerable amount of leaping and the use of forelimbs for clinging and/or suspending during landing.