Hallucal grasping in Nycticebus coucang: further implications for the functional significance of a large peroneal process

Euprimate grasping feet are characterized by a suite of morphological traits, including an enlarged peroneal process on the base of the first metatarsal, which serves as the insertion site of the peroneus longus muscle. In prosimians, a large process has typically been associated with a powerful hallucal grasp via the contraction of the peroneus longus to adduct the hallux. Recent electromyography (EMG) studies have documented that peroneus longus does not contribute substantially to hallucal grasping in lemurids (Boyer et al., 2007). However, non-lemurid prosimians have a I-V opposable grasp complex that is morphologically different and phylogenetically more primitive than the I-II adductor grasp complex of the lemurids previously studied. Therefore, it is possible that peroneus longus did function during grasping in early euprimates, but lost this function in large-bodied lemurids. The present study tests the hypothesis that a large peroneal process is related to powerful grasping ability in primates displaying the more primitive I-V grasp complex. We use EMG to evaluate the recruitment of peroneus longus, other crural muscles, and adductor hallucis in static and locomotor grasping activities of the slow loris (Nycticebus coucang). Results show that peroneus longus is active during grasping behaviors that require the subject to actively resist inversion of the foot, and likely contributes to a hallucal grasp in these activities. Peroneus longus activity level does not differ between grasping and power grasping activities, nor does it differ between grasping and non-grasping locomotor modes. Conversely, the digital flexors and hallucal adductor are recruited at higher levels during power grasping and grasping locomotor modes. Consequently, we reject the hypothesis that an enlarged peroneal process represents an adaptation specifically to enhance the power of the I-V grasp, but accept that the muscle likely plays a role in adducting the hallux during grasping behaviors that require stabilization of the ankle, and suggest that further work is necessary to determine if this role is sufficient to drive selection for a large peroneal process.     

Comparative functional morphology of the primate peroneal process

The first metatarsal of living Primates is characterized by a well-developed peroneal process, which appears proportionally larger in prosimians than in anthropoids. A large peroneal process has been hypothesized to: 1) reflect powerful hallucal grasping, 2) act as a buttress to reduce strain from loads acting on the entocuneiform-first metatarsal joint during landing and grasping after a leap, and/or 3) correlate with differences in physiological abduction of the hallux. In this study, we address the latter two hypotheses by comparing the morphology of the peroneal process in 143 specimens representing 37 species of extant prosimians, platyrrhine anthropoids, and tupaiids (tree shrews) that engage in different locomotor behaviors. In particular, we compare taxa that vary in leaping frequency and hallucal abduction. Linear and angular measurements on the first metatarsal were obtained to evaluate differences in relative peroneal process thickness and length, first metatarsal abduction angle, and overall first metatarsal shape. Leaping frequency was significantly correlated only with relative peroneal process thickness within extant lorisoids. Relative process length was positively correlated with the angle of hallucal abduction within prosimians; this angle is significantly greater in prosimians than anthropoids. Multivariate analyses of metatarsal shape effectively separate species along phylogenetic lines, but not by locomotor behaviors. The hypothesis that the peroneal process on the first metatarsal reduces the loads on the entocuneiform-first metatarsal joint during landing after a leap is in part supported by data from extant lorisoids (i.e., slow quadrupedal lorises vs. leaping galagos). A peroneal process of greater length within prosimians may serve to increase the lever arm for the peroneus longus muscle in order to prevent hyper-abduction, followed by inversion in locomotor situations where the animal's weight is born on a highly divergent/abducted hallux.     

Functional morphology of the hallucal metatarsal with implications for inferring grasping ability in extinct primates

Primate evolutionary morphologists have argued that selection for life in a fine branch niche resulted in grasping specializations that are reflected in the hallucal metatarsal (Mt1) morphology of extant “prosimians”, while a transition to use of relatively larger, horizontal substrates explains the apparent loss of such characters in anthropoids. Accordingly, these morphological characters—Mt1 torsion, peroneal process length and thickness, and physiological abduction angle—have been used to reconstruct grasping ability and locomotor mode in the earliest fossil primates. Although these characters are prominently featured in debates on the origin and subsequent radiation of Primates, questions remain about their functional significance. This study examines the relationship between these morphological characters of the Mt1 and a novel metric of pedal grasping ability for a large number of extant taxa in a phylogenetic framework. Results indicate greater Mt1 torsion in taxa that engage in hallucal grasping and in those that utilize relatively small substrates more frequently. This study provides evidence that Carpolestes simpsoni has a torsion value more similar to grasping primates than to any scandentian. The results also show that taxa that habitually grasp vertical substrates are distinguished from other taxa in having relatively longer peroneal processes. Furthermore, a longer peroneal process is also correlated with calcaneal elongation, a metric previously found to reflect leaping proclivity. A more refined understanding of the functional associations between Mt1 morphology and behavior in extant primates enhances the potential for using these morphological characters to comprehend primate (locomotor) evolution. Am J Phys Anthropol 156:327–348, 2015. © 2014 Wiley Periodicals, Inc.     

Telemetered electromyography of peroneus longus in Varecia variegata and Eulemur rubriventer: implications for the functional significance of a large peroneal process

A foot specialized for grasping small branches with a divergent opposable hallux (hallucal grasping) represents a key adaptive complex characterizing almost all arboreal non-human euprimates. Evolution of such grasping extremities probably allowed members of a lineage leading to the common ancestor of modern primates to access resources available in a small-branch niche, including angiosperm products and insects. A better understanding of the mechanisms by which euprimates use their feet to grasp will help clarify the functional significance of morphological differences between the euprimate grasp complex and features representing specialized grasping in other distantly related groups (e.g., marsupials and carnivorans) and in closely related fossil taxa (e.g., plesiadapiforms). In particular, among specialized graspers euprimates are uniquely characterized by a large peroneal process on the base of the first metatarsal, but the functional significance of this trait is poorly understood. We tested the hypothesis that the large size of the peroneal process corresponds to the pull of the attaching peroneus longus muscle recruited to adduct the hallux during grasping. Using telemetered electromyography on three individuals of Varecia variegata and two of Eulemur rubriventer, we found that peroneus longus does not generally exhibit activity consistent with an important function in hallucal grasping. Instead, extrinsic digital flexor muscles and, sometimes, the intrinsic adductor hallucis are active in ways that indicate a function in grasping with the hallux. Peroneus longus helps evert the foot and resists its inversion. We conclude that the large peroneal tuberosity that characterizes the hallucal metatarsal of prosimian euprimates does not correlate to "powerful" grasping with a divergent hallux in general, and cannot specifically be strongly linked to vertical clinging and climbing on small-diameter supports. Thus, the functional significance of this hallmark, euprimate feature remains to be determined.     

Hallucal grasping behavior in Caluromys (Didelphimorphia: Didelphidae): Implications for primate pedal grasping

A key feature in primate evolution is a foot with a divergent opposable hallucal metatarsal bearing a large peroneal process. Extant primates are characterized by a powerful hallucal grasp—an either “euprimate” or “plesiadapoid-euprimate” ancestor acquisition—that facilitates the exploitation of fine branches, an ability that increased the fitness of ancestral euprimates. In this context, the didelphid marsupial Caluromys has been used as the extant analog to this primate morphotype stage due to some morphological, ecological, and behavioral features. However, the extent to which and the positional and support contexts in which Caluromys uses powerful hallucal grasping are not known. This renders analogies to any mode of “euprimate” or “stem primate” grasping poorly substantiated. The present paper quantifies locomotor and postural behavior, support use, and associated frequencies of hallucal grasping in captive Caluromys philander via analysis of video recordings. During locomotion, Caluromys primarily used diagonal sequence walk, clamber, and climb, whereas stand, foot-hang, and bipedal stand were the dominant postures. Small, fine, horizontal, and moderately inclined branches were frequently used. Overall rates of “apparently powerful hallucal grasps” were high, but were exceptionally high during clamber, climb, foot-hang, and bipedal stand. Additionally, an “apparently powerful hallucal grasp” was very common upon fine, small, steep, and vertical branches. The extensive use of such powerful hallucal grasping provided stability and safety that enabled Caluromys to proficiently utilize fine branches of various orientations. The ability to negotiate such unstable supports, further reflected in foot anatomy, provides evidence that the morphobehavioral complex of Caluromys can serve as an extant analog to the plesiadapoid-euprimate ancestor, represented as a terminal branch feeder with effective hallucal grasping.     

New primate first metatarsals from the Paleogene of Egypt and the origin of the anthropoid big toe

The specialized grasping feet of primates, and in particular the nature of the hallucal grasping capabilities of living strepsirrhines and tarsiers (i.e., ‘prosimians’), have played central roles in the study of primate origins. Prior comparative studies of first metatarsal (Mt1) morphology have documented specialized characters in living prosimians that are indicative of a more abducted hallux, which in turn is often inferred to be related to an increased ability for powerful grasping. These include a well-developed peroneal process and a greater angle of the proximal articular surface relative to the long axis of the diaphysis. Although known Mt1s of fossil prosimians share these characters with living non-anthropoid primates, Mt1 morphology in the earliest crown group anthropoids is not well known. Here we describe two Mt1s from the Fayum Depression of Egypt - one from the latest Eocene (from the ∼34 Ma Quarry L-41), and one from the later early Oligocene (from the ∼29-30 Ma Quarry M) - and compare them with a sample of extant and fossil primate Mt1s. Multivariate analyses of Mt1 shape variables indicate that the Fayum specimens are most similar to those of crown group anthropoids, and likely belong to the stem catarrhines Catopithecus and Aegyptopithecus specifically, based on analyses of size. Also, phylogenetic analyses with 16 newly defined Mt1 characters support the hypotheses that “prosimian”-like Mt1 features evolved along the primate stem lineage, while crown anthropoid Mt1 morphology and function is derived among primates, and likely differed from that of basal stem anthropoids. The derived loss of powerful hallucal grasping as reflected in the Mt1 morphology of crown anthropoids may reflect long-term selection for improved navigation of large-diameter, more horizontal branches at the expense of movement in smaller, more variably inclined branches in the arboreal environment.     

Mouse hallucal metatarsal cross‐sectional geometry in a simulated fine branch niche

Mice raised in experimental habitats containing an artificial network of narrow “arboreal” supports frequently use hallucal grasps during locomotion. Therefore, mice in these experiments can be used to model a rudimentary form of arboreal locomotion in an animal without other morphological specializations for using a fine branch niche. This model would prove useful to better understand the origins of arboreal behaviors in mammals like primates. In this study, we examined if locomotion on these substrates influences the mid‐diaphyseal cross‐sectional geometry of mouse metatarsals. Thirty CD‐1/ICR mice were raised in either arboreal (composed of elevated narrow branches of varying orientation) or terrestrial (flat ramps and walkways that are stratified) habitats from weaning (21 days) to adulthood (≥4 months). After experiments, the hallucal metatarsal (Mt1) and third metatarsal (Mt3) for each individual were isolated and micro‐computed tomography (micro‐CT) scans were obtained to calculate mid‐shaft cross‐sectional area and polar section modulus. Arboreal mice had Mt1s that were significantly more robust. Mt3 cross sections were not significantly different between groups. The arboreal group also exhibited a significantly greater Mt1/Mt3 ratio for both robusticity measures. We conclude that the hallucal metatarsal exhibits significant phenotypic plasticity in response to arboreal treatment due to habitual locomotion that uses a rudimentary hallucal grasp. Our results support the hypothesis that early adaptive stages of fine branch arboreality should be accompanied by a slightly more robust hallux associated with the biomechanical demands of this niche. J. Morphol. 276:759–765, 2015. © 2015 Wiley Periodicals, Inc.     

The skeleton of early Eocene Cantius, oldest lemuriform primate

A recently discovered partial skeleton of the adapid Cantius trigonodus from the early Eocene Willwood Formation of the Bighorn Basin, Wyoming, documents substantial new information about the anatomy of the oldest lemuriform primates. It is very similar in all features to its descendant, middle Eocene Notharctus, and both exhibit numerous resemblances to certain extant Malagasy lemurs, particularly Lepilemur, Propithecus, Lemur, and Hapalemur griseus. Like these forms, Cantius had relatively long hind limbs and short forelimbs. Forelimb traits (prominent brachialis flange of the humerus, well-developed olecranon process of the ulna, and strong shafts of the ulna and radius) suggest active use of the forelimbs in progression. Specializations in the hind limb (e.g., expanded articular surface of the femoral head, narrow and elevated patellar trochlea and prominent lateral trochlear ridge, posteriorly oriented femoral and tibial condyles, narrow and elongate talus, and hallucal metatarsal with prominent peroneal tubercle) indicate capabilities for leaping and for powerful grasping with an opposable hallux. Cantius was presumably primitive in having a relatively long ischium and much more distal inferior tibial tuberosity than most extant lemurs--traits suggesting that powerful extension of the thigh and flexion at the knee were important in its locomotion and posture. We interpret Cantius as an active arboreal quadruped with a propensity for leaping. The existence of this skeletal structure in one of the oldest primates of modern aspect suggests that it represents the primitive lemuriform morphology.     

Grasping behavior in tufted capuchin monkeys (Cebus apella): grip types and manual laterality for picking up a small food item

This study investigates prehension in 20 tufted capuchins (Cebus apella) in a reaching task requiring individuals to grasp a small food item fixed to a tray. The aim was twofold: 1) to describe capuchins' grasping techniques in detail, focusing on digit movements and on different areas of contact between the grasping fingers; and 2) to assess the relationship between grip types and manual laterality in this species. Capuchins picked up small food items using a wide variety of grips. In particular, 16 precision grip variants and 4 power grip variants were identified. The most frequently used precision grip involved the distal lateral areas of the thumb and the index finger, while the most preferred kind of power grip involved the thumb and the palm, with the thumb being enclosed by the other fingers. Immature capuchins picked up small food items using power grips more often than precision grips, while adult individuals exhibited no significant preference for either grip type. The analysis performed on the time capuchins took to grasp the food and withdraw it from the tray hole revealed that 1) precision grips were as efficient as power grips; 2) for precision grips, the left hand was faster than the right hand; and 3) for power grips, both hands were equally quick. Hand preference analysis, based on the frequency for the use of either hand for grasping actions, revealed no significant hand bias at group level. Likewise, there was no significant relationship between grip type and hand preference.     

Grasping primate development: Ontogeny of intrinsic hand and foot proportions in capuchin monkeys (Cebus albifrons and Sapajus apella)

Young primates have relatively large hands and feet for their body size, perhaps enhancing grasping ability. We test the hypothesis that selection for improved grasping ability is responsible for these scaling trends by examining the ontogeny of intrinsic hand and foot proportions in capuchin monkeys (Cebus albifrons and Sapajus apella). If selection for improved grasping ability is responsible for the observed patterns of hand and foot growth in primates, we predicted that fingers and toes would be longer early in life and proportionally decline with age. We measured the lengths of manual and pedal metapodials and phalanges in a mixed‐longitudinal radiographic sample. Bone lengths were (a) converted into phalangeal indices (summed non‐distal phalangeal length/metapodial length) to test for age‐related changes in intrinsic proportions and (b) fit to Gompertz models of growth to test for differences in the dynamics of phalangeal versus metapodial growth. Manual and pedal phalangeal indices nearly universally decreased with age in capuchin monkeys. Growth curve analyses revealed that metapodials generally grew at a faster rate, and for a longer duration, than corresponding phalanges. Our findings are consistent with the hypothesis that primates are under selection for increased grasping ability early in life. Relatively long digits may be functionally adaptive for growing capuchins, permitting a more secure grasp on both caregivers and arboreal supports, as well as facilitating early foraging. Additional studies of primates and other mammals, as well as tests of grasping performance, are required to fully evaluate the adaptive significance of primate hand and foot growth.     

Getting a grip on the evolution of grasping in musteloid carnivorans: a three‐dimensional analysis of forelimb shape

The ability to grasp and manipulate is often considered a hallmark of hominins and associated with the evolution of their bipedal locomotion and tool use. Yet, many other mammals use their forelimbs to grasp and manipulate objects. Previous investigations have suggested that grasping may be derived from digging behaviour, arboreal locomotion or hunting behaviour. Here, we test the arboreal origin of grasping and investigate whether an arboreal lifestyle could confer a greater grasping ability in musteloid carnivorans. Moreover, we investigate the morphological adaptations related to grasping and the differences between arboreal species with different grasping abilities. We predict that if grasping is derived from an arboreal lifestyle, then the anatomical specializations of the forelimb for arboreality must be similar to those involved in grasping. We further predict that arboreal species with a well‐developed manipulation ability will have articulations that facilitate radio‐ulnar rotation. We use ancestral character state reconstructions of lifestyle and grasping ability to understand the evolution of both traits. Finally, we use a surface sliding semi‐landmark approach capable of quantifying the articulations in their full complexity. Our results largely confirm our predictions, demonstrating that musteloids with greater grasping skills differ markedly from others in the shape of their forelimb bones. These analyses further suggest that the evolution of an arboreal lifestyle likely preceded the development of enhanced grasping ability.     

Electromyographic activity in the Rhesus monkey forelimb muscles during a goal directed movement and locomotion before, during and after spaceflight

The aim of the present study was to analyse the effects of microgravity on i) the achievement of goal-directed arm movements and ii) the quadrupedal non-human primate locomotion. A reaching movement in weightlessness would require less muscle contraction since there is no need to oppose gravity. Consequently the electromyographic (EMG) activity of the monkey forelimb muscles should be changed during or after spaceflight. EMG activity of the biceps and triceps muscles during goal-directed arm movements were studied in Rhesus monkeys before, during and after 14 days of spaceflight and flight simulation at normal gravity. The EMG activity was also recorded during treadmill locomotion before and after spaceflight. When performing arm motor tasks, the delay values of the EMG bursts were unchanged during the flight. On the contrary, mean EMG was significantly decreased during the flight comparatively to the pre- and post-flight values, which were very similar. Compared with flight animals, the control ground monkey showed no change in the burst durations and mean EMG. After spaceflight, quadrupedal locomotion was modified. The animals had some difficulty in moving, and abnormal steps were numerous. The integrated area of triceps bursts was increased for the stance phase during locomotion. Taken together these data showed that spaceflight induces a dual adaptative process: first, the discharge of the motor pools of the forelimb musculature was modified during exposure to microgravity, and then upon return to Earth, monkeys changed their new motor strategy and re-adapt to normal gravity.     

Articular to diaphyseal proportions of human and great ape metatarsals

This study proposes a new way to use metatarsals to identify locomotor behavior of fossil hominins. Metatarsal head articular dimensions and diaphyseal strength in a sample of chimpanzees, gorillas, orangutans, and humans (n = 76) are used to explore the relationships of these parameters with different locomotor modes. Results show that ratios between metatarsal head articular proportions and diaphyseal strength of the hallucal and fifth metatarsal discriminate among extant great apes and humans based on their different locomotor modes. In particular, the hallucal and fifth metatarsal characteristics of humans are functionally related to the different ranges of motion and load patterns during stance phase in the forefoot of humans in bipedal locomotion. This method may be applicable to isolated fossil hominin metatarsals to provide new information relevant to debates regarding the evolution of human bipedal locomotion. The second to fourth metatarsals are not useful in distinguishing among hominoids. Further studies should concentrate on measuring other important qualitative and quantitative differences in the shape of the metatarsal head of hominoids that are not reflected in simple geometric reconstructions of the articulation, and gathering more forefoot kinematic data on great apes to better understand differences in range of motion and loading patterns of the metatarsals. Am J Phys Anthropol 143:198–207, 2010. © 2010 Wiley‐Liss, Inc.     

Activity and functions of the human gluteal muscles in walking,running, sprinting,and climbing

It has been suggested that the uniquely large gluteus maximus (GMAX) muscles were an important adaptation during hominin evolution based on numerous anatomical differences between humans and extant apes. GMAX electromyographic (EMG) signals have been quantified for numerous individual movements, but not across the range of locomotor gaits and speeds for the same subjects. Thus, comparing relative EMG amplitudes between these activities has not been possible. We assessed the EMG activity of the gluteal muscles during walking, running, sprinting, and climbing. To gain further insight into the function of the gluteal muscles during locomotion, we measured muscle activity during walking and running with external devices that increased or decreased the need to control either forward or backward trunk pitch. We hypothesized that 1) GMAX EMG activity would be greatest during sprinting and climbing and 2) GMAX EMG activity would be modulated in response to altered forward trunk pitch demands during running. We found that GMAX activity in running was greater than walking and similar to climbing. However, the activity during sprinting was much greater than during running. Further, only the inferior portion of the GMAX had a significant change with altered trunk pitch demands, suggesting that the hip extensors have a limited contribution to the control of trunk pitch movements during running. Overall, our data suggest that the large size of the GMAX reflects its multifaceted role during rapid and powerful movements rather than as a specific adaptation for a single submaximal task such as endurance running. Am J Phys Anthropol 153:124–131, 2014. © 2013 Wiley Periodicals, Inc.     

Positional behavior and substrate use in wild adult bearded capuchin monkeys (Sapajus libidinosus)

Natural selection for positional behavior (posture and locomotion) has at least partially driven the evolution of anatomical form and function in the order Primates. Examination of bipedal behaviors associated with daily activity patterns, foraging, and terrestrial habitat use in nonhuman primates, particularly those that adopt bipedal postures and use bipedal locomotion, allows us to refine hypotheses concerning the evolution of bipedalism in humans. This study describes the positional behavior of wild bearded capuchins (Sapajus libidinosus), a species that is known for its use of terrestrial substrates and its habitual use of stones as tools. Here, we test the association of terrestrial substrate use with bipedal posture and locomotion, and the influence of sex (which co‐varies with body mass in adults of this species) on positional behavior and substrate use. Behavior and location of 16 wild adult bearded capuchins from two groups were sampled systematically at 15 s intervals for 2 min periods for 1 year (10,244 samples). Despite their different body masses, adult males (average 3.5 kg) and females (average 2.1 kg) in this study did not differ substantially in their positional behaviors, postures, or use of substrates for particular activities. The monkeys used terrestrial substrates in 27% of samples. Bipedal postures and behaviors, while not a prominent feature of their behavior, occurred in different forms on the two substrates. The monkeys crouched bipedally in trees, but did not use other bipedal postures in trees. While on terrestrial substrates, they also crouched bipedally but occasionally stood upright and moved bipedally with orthograde posture. Bearded capuchin monkeys' behavior supports the suggestion from anatomical analysis that S. libidinosus is morphologically better adapted than its congeners to adopt orthograde postures.     

How Aging Affects Grasping Behavior and Pull Strength in Captive Gray Mouse Lemurs (<Emphasis Type="Italic">Microcebus murinus</Emphasis>)

Prehension is essential for animal survival and fitness. It is involved in locomotion and feeding behavior and subject to physical and physiological constraints. Studies of prehension in primates have explored the importance of food properties and of the environment, but aging has rarely been studied although prehensile capacity may deteriorate with age in humans. To test the hypothesis that aging affects grasping abilities and to reveal possible behavioral adaptations to this, we quantified behavioral grasping strategies and pull strength in 10 young adult (2–3 yr old) and 10 aged (7–8 yr old) gray mouse lemurs (Microcebus murinus). We assessed grasping strategies in an experimental cage by quantifying grip types used to grasp static and mobile foods. We measured strength using a Kistler triaxial force platform. Our results show that 1) mobile and static foods affected individuals of different ages in similar ways; 2) older individuals used more mouth grasps than young ones; 3) aged individuals made twice as many attempts as young ones when grasping mobile food items but this difference was not significant; and 4) there were no differences in hand grip strength between age classes but young individuals showed a higher foot pull strength compared to old ones. These data suggest that the observed differences in behavior may be due to a decrease in foot grip strength, which in turn influences stability on narrow branches, forcing animals to use their hands to maintain stability and preventing them from using their hands for food-related tasks.     

Manual Function in Cebus apella. Digital Mobility,Preshaping, and Endurance in Repetitive Grasping

Manual dexterity varies across species of primates in accord with hand morphology and degree of fine motor control of the digits. Platyrrhine monkeys achieve less direct opposition between thumb and index finger than that of catarrhine primates, and many of them typically whole-hand grip. However, tufted capuchins (Cebus apella), exhibit a degree of independent control of the digits and effective thumb–forefinger opposition. We report how capuchins prehended small objects, with particular attention to the form of sequential fine movements of the fingers, choice of hand, and differences between the two hands in the temporal properties of reaching and grasping. We compare these actions across tasks with differing demands for fine motor control. For tasks that required all the digits to flex in synchrony, capuchins displayed smooth, fast, and efficient reach-to-grasp movements and a higher endurance than for tasks requiring more complex digital coordination. These latter tasks induced a slightly differentiated preshaping of the hand when approaching the objects, indicating preparation for grasping in advance of contact with the object. A right-hand preponderance for complex digital coordination was evident. The monkeys coordinated their fingers rather poorly at the substrate, and they took longer to achieve control of the objects when complex coordination was required than when simultaneous flexion was sufficient. We conclude that precise finger coordination is more effortful and less well coordinated, and the coordination is less lateralized, in capuchins than in catarrhine primates.     

The morphological basis of hallucal orientation in extant birds

The perching foot of living birds is commonly characterized by a reversed or opposable digit I (hallux). Primitively, the hallux of nonavian theropod dinosaurs was unreversed and lay parallel to digits II-IV. Among basal birds, a unique digital innovation evolved in which the hallux opposes digits II-IV. This digital configuration is critical for grasping and perching. I studied skeletons of modern birds with a range of hallucal designs, from unreversed (anteromedially directed) to fully reversed (posteriorly directed). Two primary correlates of hallucal orientation were revealed. First, the fossa into which metatarsal I articulates is oriented slightly more posteriorly on the tarsometatarsus, rotating the digit as a unit. Second, metatarsal I exhibits a distinctive torsion of its distal shaft relative to its proximal articulation with the tarsometatarsus, reorienting the distal condyles and phalanges of digit I. Herein, I present a method that facilitates the re-evaluation of hallucal orientation in fossil avians based on morphology alone. This method also avoids potential misinterpretations of hallucal orientation in fossil birds that could result from preserved appearance alone.     

Temporalis and masseter muscle function during incision in macaques and humans

Most previously published electromyographic (EMG) studies have indicated that the temporalis muscles in humans become almost electrically quiet during incisai biting. These data have led various workers to conclude that these muscles may contribute little to the incisai bite force. The feeding behavior and comparative anatomy of the incisors and temporalis muscles of certain catarrhine primates, however, suggest that the temporalis muscle is an important and powerful contributor to the bite force during incision. One purpose of this study is to analyze the EMG activity of the masseter and temporalis muscles in both humans and macaques with the intention of focusing on the conflict between published EMG data on humans and inferences of muscle function based on the comparative anatomy and behavior of catarrhine primates. The EMG data collected from humans in the present study indicate that, in five of seven subjects, the masseter,anterior temporalis, and posterior temporalis muscles are very active during apple incision (i.e., relative to EMG activity levels during apple and almond mastication). In the other two human subjects the EMG levels of these muscles are lower during incision than during mastication, but in no instance are these muscles ever close to becoming electrically quiet. The EMG data on macaques indicate that, in all six subjects, the masseter, anterior temporalis, and posterior temporalis muscles are very active during incision. These data are in general agreement with inferences on muscle function that have been drawn from the comparative anatomy and behavior of primates, but they do not agree with previous experimental data. The reason for this disagreement is probably due to differences in the experimental procedure. In previous studies subjects simply bit isometrically on their incisors and the resulting EMG pattern was compared to the pattern associated with powerful clenching in centric occlusion. In the present study the subjects incised into actual food objects, and the resulting EMG pattern was compared to the pattern associated with mastication of various foods. It is not surprising that these two procedures result in markedly different EMG patterns, which in turn result in markedly different interpretations of jaw-muscle function. In an attempt to explain the evolution of the postorbital septum in anthropoids, it has been suggested that the anterior temporalis is more active than the masseter during incision (Cachel, 1979). The human and macaque EMG data do not support this hypothesis; during incision, the two muscles show no consistent differences in humans and the masseter appears to be in fact more active than the anterior temporalis in macaques.     

Effect of footwear on intramuscular EMG activity of plantar flexor muscles in walking

One of the purposes of footwear is to assist locomotion, but some footwear types seem to restrict natural foot motion, which may affect the contribution of ankle plantar flexor muscles to propulsion. This study examined the effects of different footwear conditions on the activity of ankle plantar flexors during walking. Ten healthy habitually shod individuals walked overground in shoes, barefoot and in flip-flops while fine-wire electromyography (EMG) activity was recorded from flexor hallucis longus (FHL), soleus (SOL), and medial and lateral gastrocnemius (MG and LG) muscles. EMG signals were peak-normalised and analysed in the stance phase using Statistical Parametric Mapping (SPM). We found highly individual EMG patterns. Although walking with shoes required higher muscle activity for propulsion than walking barefoot or with flip-flops in most participants, this did not result in statistically significant differences in EMG amplitude between footwear conditions in any muscle (p > 0.05). Time to peak activity showed the lowest coefficient of variation in shod walking (3.5, 7.0, 8.0 and 3.4 for FHL, SOL, MG and LG, respectively). Future studies should clarify the sources and consequences of individual EMG responses to different footwear.