Olduvai Hominid 7 trapezial metacarpal 1 articular morphology: contrasts with recent humans

A consideration of the metacarpal 1 articulation of the Olduvai Hominid 7 trapezium shows that, although it is generally similar to those of recent humans, it is considerably flatter radioulnarly and dorsopalmarly than those of modern humans and similar in its degree of dorsopalmar flattening to those of Upper Pleistocene late archaic humans. Even though this flattening may have slightly limited flexion-extension at the pollical carpometacarpal joint, it is most likely related to the apparently elevated levels of axial joint reaction force through the thumbs of archaic members of the genus Homo. It also documents the continuation of human carpometacarpal articular evolution through the Pleistocene.     

Oreopithecus bambolii: an unlikely case of hominid-like grip capability in a Miocene ape

Oreopithecus bambolii, an ape from the late Miocene of Italy, is said to possess a hand capable of a precision grip like that of humans. Relative hand length, proportions of the thumb, and morphological features of the thumb and wrist were adduced to support the idea that Oreopithecus had a hand that closely matched the pattern in Australopithecus. A reappraisal of earlier arguments and comparisons of Oreopithecus with humans, apes, and Old World monkeys, reveals that Oreopithecus had an essentially ape-like hand that emphasized ape-like power grasping over human-like precision grasping.     

Shanidar 1: a case of hyperostotic disease (DISH) in the middle Paleolithic.

The Shanidar 1 Neandertal partial skeleton presents osteophytic lesions on its vertebrae and appendicular skeleton which appear independent of the multiple traumatic and degenerative joint disease lesions on the individual. In particular, the large flowing osteophyte on the L3 body, a smaller one on the L5 body, and enthesopathic osteophytes on both calcaneal tuberosities, both patellae and the left ulnar olecranon, support a diagnosis of hyperostotic disease (DISH). The diagnosis is supported by small enthesopathic osteophytes on the preserved femoral greater trochanter and scapular corocoid process. This diagnosis would make it the oldest hominid specimen clearly presenting this systemic condition.     

La Ferrassie 6 and the development of Neandertal pubic morphology

Functional interpretations of the mediolateral elongation and superoinferior flattening of the superior pubic ramus of Neandertals require knowledge of its ontogeny. Metric comparisons between the La Ferrassie 6 Neandertal infant, aged 3-5 years, and a sample of modern infants reveal that the acetabulosymphyseal length of La Ferrassie 6, relative to femoral length and iliac breadth, falls at the limits of the range of variation of the modern infants, while the relative height of its superior pubic ramus is indistinguishable from that of the modern sample. It appears that acetabulosymphyseal elongation of the Neandertal pubis is a feature which is expressed quite early in ontogeny, well before puberty. Superoinferior flattening and ventral margin thinning of the superior ramus probably appear later in Neandertal ontogeny.     

Evolution of viviparity in warm-climate lizards: an experimental test of the maternal manipulation hypothesis

The maternal manipulation hypothesis for the evolution of reptilian viviparity has been claimed to apply to any situation where gravid females are able to maintain body temperatures different from those available in external nests, but empirical data that support this hypothesis are very limited. Here, we tested this hypothesis using gravid females of a warm-climate lizard, Mabuya multifasciata, by subjecting them to five thermal regimes for the whole gestation period. We found gravid females selected lower body temperatures and thermoregulated more precisely than did nongravid females. Offspring produced in different treatments differed in head size, limb length and sprint speed, but not in overall body size or mass. Variation in morphological traits of offspring was induced primarily by extreme temperatures. Sprint speed of offspring was more likely affected by the mean but not by the variance of gestation temperatures. Gravid females maintained more stable body temperatures than did nongravid females not because these temperatures resulted in the optimization of offspring phenotypes but because the range of temperatures optimal for embryonic development was relatively narrow. Our data conform to the main predictions from the maternal manipulation hypothesis that females should adjust thermoregulation during pregnancy to provide optimal thermal conditions for developing embryos and that phenotypic traits forged by maternal thermoregulation should enhance offspring fitness.     

Morphological affinities of the Australopithecus afarensis hand on the basis of manual proportions and relative thumb length

The hands of apes and humans differ considerably with regard to proportions between several bones. Of critical significance is the long thumb relative to other fingers, which is the basis for human-like pad-to-pad precision grip capability, and has been considered by some as evidence of tool-making. The nature and timing of the evolutionary transition from ape-like to human-like manual proportions, however, have remained unclear as a result of the lack of appropriate fossil material. In this article, the manual proportions of Australopithecus afarensis from locality AL 333/333w (Hadar, Ethiopia) are investigated by means of bivariate and multivariate morphometric analyses, in order to test the hypothesis that human-like proportions, including an enhanced thumb/hand relationship, originally evolved as an adaptation to stone tool-making. Although some evidence for human-like manual proportions had been previously proposed for this taxon, conclusive evidence was lacking. Our results indicate that A. afarensis possessed overall manual proportions, including an increased thumb/hand relationship that, contrary to previous reports, is fully human and would have permitted pad-to-pad human-like precision grip capability. We show that these human-like proportions in A. afarensis mainly result from hand shortening, as in modern humans, and that these conclusions are robust enough as to be non-dependent on whether the bones belong to a single individual or not. Since A. afarensis predates the appearance of stone tools in the archeological record, the above-mentioned conclusions permit a confident refutation of the null hypothesis that human-like manual proportions are an adaptation to stone tool-making, and thus alternative explanations must be therefore sought. One hypothesis would consider manipulative behaviors (including tool-use and/or non-lithic tool-making) in early hominines exceeding those reported among extant non-human primates. Alternatively, on the basis of the many adaptations to committed bipedalism in A. afarensis, we propose the hypothesis that once arboreal behaviors became adaptively insignificant and forelimb-dominated locomotor selection pressures were relaxed with the adoption of terrestrial bipedalism, human-like manual proportions could have merely evolved as a result of the complex manipulation selection pressures already present in extant non-human primates.Both hypotheses are not mutually exclusive, and even other factors such as pleiotropy cannot be currently discarded.     

Movement of modulator in maize: a test of an hypothesis

A model of Modulator movement (Greenblatt 1968) from one chromosomal site to another requires that all movements produce potential twin mutations within the affected cell lineage. From this model the prediction would be that (1) untwinned red and untwinned light-variegated sectors within the pericarp of medium-variegated maize must occur in equal frequency even though partners become lost, during ear morphogenesis, to the final pericarp tissue, and that (2) among the backcross progeny of a homozygous P(rr)Mp individual mated with P(wr)/P(wr) pollen, red offspring and light-variegated offspring would occur in equal frequency. Both expectations have been realized and herein reported.     

Incidence and patterning of dental enamel hypoplasia among the Neandertals

Dental enamel hypoplasia (DEH), as an indicator of nonspecific stress during development, provides an assessment of the relative morbidity of past human populations. An investigation of 669 Neandertal dental crowns yielded an overall DEH frequency of 36.0% by tooth (41.9% for permanent teeth; 3.9% for deciduous teeth) and about 75% by individual. These incidences place the Neandertals at the top of recent human ranges of variation in DEH frequencies, indicating high levels of stress during development. The paucity of deciduous tooth DEH and M1 DEH, combined with generally increasing levels of DEH through later calcifying teeth, suggests that the stress was primarily nutritional, beginning at weaning and continuing through adolescence. This supports paleontological and archeological interpretations implying significantly lower effectiveness for Neandertal foraging compared to that of modern humans.     

Development of a dynamic index finger and thumb model to study impairment

Modeling of the human hand provides insight for explaining deficits and planning treatment following injury. Creation of a dynamic model, however, is complicated by the actions of multi-articular tendons and their complex interactions with other soft tissues in the hand. This study explores the creation of a musculoskeletal model, including the thumb and index finger, to explore the effects of muscle activation deficits. The OpenSim model utilizes physiological axes of rotation at all joints, passive joint torques, and appropriate moment arms. The model was validated through comparison with kinematic and kinetic experimental data. Simulated fingertip forces resulting from modeled musculotendon loading largely fell within one standard deviation of experimental ranges for most index finger and thumb muscles, although agreement in the sagittal plane was generally better than for the coronal plane. Input of experimentally obtained electromyography data produced the expected simulated finger and thumb motion. Use of the model to predict the effects of activation deficits on pinch force production revealed that the intrinsic muscles, especially first dorsal interosseous (FDI) and adductor pollicis (ADP), had a substantial impact on the resulting fingertip force. Reducing FDI activation, such as might occur following stroke, altered fingertip force direction by up to 83° for production of a dorsal fingertip force; reducing ADP activation reduced force production in the thumb by up to 62%. This validated model can provide a means for evaluating clinical interventions.     

Early ontogeny of the human femoral bicondylar angle

The presence of a femoral bicondylar angle consistently and significantly greater than 0° has been a hallmark of hominid bipedality, but its pattern of development has not been documented. We have therefore compiled cross-sectional data on the development of the articular bicondylar angle for a clinical sample of modern humans and of the metaphyseal bicondylar angle for two Recent human skeletal samples, one predominantly European in origin and the other Amerindian. All three samples exhibit a pattern of a bicondylar angle of 0° at birth and then a steady average increase in the angle from late in the first year postnatal, through infancy, and into the juvenile years. The two skeletal samples reach low adult values by approximately 4 years postnatal, whereas the clinical sample with a lowered activity level appears to attain consistent adult values slightly later (approximately 6 years postnatal). In addition, two modern human individuals, one nonambulatory and the other minimally ambulatory, show no and little development, respectively, of a bicondylar angle. These data, in conjunction with clinical and experimental observations on the potential and form of angular changes during epiphyseal growth, establish a high degree of potential for plasticity in the development of the human bicondylar angle and the direct association of a bipedal locomotion and (especially) posture with the developmental emergence of a human femoral bicondylar angle. © 1994 Wiley-Liss, Inc.     

Particulate versus integrated evolution of the upper body in late pleistocene humans: A test of two models

Evolutionary biologists are largely polarized in their approaches to integrating microevolutionary and macroevolutionary processes. Neo-Darwinians typically seek to identify population-level selective and genetic processes that culminate in macroevolutionary events. Epigeneticists and structuralists, on the other hand, emphasize developmental constraints on the action of natural selection, and highlight the role of epigenetic shifts in producing evolutionary change in morphology. Accordingly, the ways in which these paradigms view and address morphological contrasts between classes of related organisms differ. These paradigms, although seldomly explicitly stated, emerge in paleoanthropology as well. Considerations of postcranial morphological contrasts between archaic and modern humans typically fall into one of two broad interpretive models. The first derives from the neo-Darwinian perspective and holds that evolution in the postcranial skeleton was largely mosaic (operating in a particulate manner), and that temporal change in specific traits informs us about behavioral shifts or genetic evolution affecting isolated anatomical regions (i.e., adaptive behavioral inferences can be made from comparative studies of individual trait complexes). The alternative model follows from the epigeneticist paradigm and sees change in specific postcranial traits as correlated responses to change in overall body form (involving shifts in regulation of skeletal growth, or selective and developmental responses to broad adaptive shifts). By this view, integration of functional systems both constrains and directs evolution of various traits, and morphological contrasts inform us about overall change in body form related to change in such things as overall growth patterns, climatic adaptation, and technological dependency. These models were tested by confirmatory factor analysis using measures of upper body form and upper limb morphological traits in Eurasian Neandertal and early modern fossils and recent human samples. Results indicate (1) a model of morphological integration fits the data better than a model of no integration, but (2) this integration accounts for less than half of the variance in upper limb traits, suggesting a high degree of tolerance for particulate evolution in the context of an integrated upper body plan. Significant relationships were detected between joint shapes and body size, between humeral shaft shape and body size and chest shape, and between measures of biomechanical efficiency and robusticity. The observed morphological differences between late archaic and early modern humans reflect particulate evolution in the context of constraints imposed by genetic and morphological integration. While particulate approaches to interpreting the fossil record appear to be justified, attention must also be paid to delineating the nature and extent of morphological integration and its role in both constraining and producing observed patterns of variation between groups. Confirmatory factor analysis provides a means of examining trait covariance matrices, and serves as a useful method of identifying patterns of integration in morphology. © 1996 Wiley-Liss, Inc.     

Mechanical transduction in the Golgi tendon organ: a hypothesis.

Morphological evidence, gained from light and electron microscopy, has shown that the unmyelinated terminal branches of the Ib afferent fiber innervating the Golgi tendon organ (GTO) lie within the spaces between braids of collagen. Based on empirical data it is proposed that force applied to a muscle's tendon will straighten these collagen braids and cause compressional deformation of the axon branches trapped between them. The mechanical events, which are presumed to occur within the GTO, appear to explain how it may function as a biological force transducer under static loading conditions. The mechanical principal described for the GTO may be a primitive and wide-spread biological mechanism employed by certain types of sensory receptors that function as position (and force) detectors.     

Adaptive advantages of myrmecochory: the predator-avoidance hypothesis tested over a wide geographic range

The predator‐avoidance hypothesis states that once released from the parent plant, myrmecochorous seeds are rapidly taken by ants to their nests, where they are protected from predators. Previous studies conducted to test this hypothesis have frequently neglected two major aspects necessary for its verification: 1) the influence of processes acting after the seed release and 2) the spatial evenness of such processes. Thus, large‐scale variations in the mechanisms acting beyond seed release, and possibly influencing seed escape from predators, remain poorly documented. Here, we present the results of a post‐dispersal seed‐removal experiment on the myrmecochorous herb Helleborus foetidus, aimed at verifing the predator‐avoidance hypothesis by considering two key post‐release aspects of seed fate: seed destination (dispersed or nondispersed) and seed burial (buried or not buried). Experiments were performed in four different regions in the Iberian Peninsula. After three days of exposure of seeds to the main predator (fieldmice Apodemus sylvaticus), ca 30% of the seeds were removed. Seed destination affected the proportion of seeds escaping predation, but the sign, magnitude and statistical significance of the effect varied among the geographical regions. In the southern region (Cazorla), seeds dispersed in ant nests or intermediate destinations suffered scarcely any predation, but seeds under reproductive‐age plants experienced losses ca 50%. Conversely, in the northern region (Caurel), seeds in nests suffered significantly greater losses than seeds under plants or intermediate destinations, suggesting that nests were especially unsafe destinations. Seed burial had a strong impact on seed escape from predators, and its effect was highly consistent among geographical regions. In view of the consistency of its effect at different spatial scales, seed burial was a more general mechanism for predation avoidance than seed relocation to ant nests, which was habitat‐ and/or ant‐species‐dependent. Our results thus only partially support the predator‐avoidance hypothesis for the evolution of myrmecochory.     

A deterministic model of admixture and genetic introgression: the case of Neanderthal and Cro-Magnon

There is an ongoing debate in the field of human evolution about the possible contribution of Neanderthals to the modern human gene pool. To study how the Neanderthal private alleles may have spread over the genes of Homo sapiens, we propose a deterministic model based on recursive equations and ordinary differential equations. If the Neanderthal population was large compared to the Homo sapiens population at the beginning of the contact period, we show that genetic introgression should have been fast and complete meaning that most of the Neanderthal private alleles should be found in the modern human gene pool in case of ancient admixture. In order to test/reject ancient admixture from genome-wide data, we incorporate the model of genetic introgression into a statistical hypothesis-testing framework. We show that the power to reject ancient admixture increases as the ratio, at the time of putative admixture, of the population size of Homo sapiens over that of Neanderthal decreases. We find that the power to reject ancient admixture might be particularly low if the population size of Homo sapiens was comparable to the Neanderthal population size.     

The matrix-tolerance hypothesis: an empirical test with frogs in the Atlantic Forest

The matrix-tolerance hypothesis suggests that the most abundant species in the inter-habitat matrix would be less vulnerable to their habitat fragmentation. This model was tested with leaf-litter frogs in the Atlantic Forest where the fragmentation process is older and more severe than in the Amazon, where the model was first developed. Frog abundance data from the agricultural matrix, forest fragments and continuous forest localities were used. We found an expected negative correlation between the abundance of frogs in the matrix and their vulnerability to fragmentation, however, results varied with fragment size and species traits. Smaller fragments exhibited stronger matrix-vulnerability correlation than intermediate fragments, while no significant relation was observed for large fragments. Moreover, some species that avoid the matrix were not sensitive to a decrease in the patch size, and the opposite was also true, indicating significant differences with that expected from the model. Most of the species that use the matrix were forest species with aquatic larvae development, but those species do not necessarily respond to fragmentation or fragment size, and thus affect more intensively the strengthen of the expected relationship. Therefore, the main relationship expected by the matrix-tolerance hypothesis was observed in the Atlantic Forest; however we noted that the prediction of this hypothesis can be substantially affected by the size of the fragments, and by species traits. We propose that matrix-tolerance model should be broadened to become a more effective model, including other patch characteristics, particularly fragment size, and individual species traits (e.g., reproductive mode and habitat preference).     

Pathology and the posture of the La Chapelle-aux-Saints Neandertal

The depiction of the Neandertals as incompletely erect was based primarily on Boule's (1911, 1912a, 1913) analysis of the La Chapelle-aux-Saints 1 partial skeleton. The inaccurate aspects of Boule's postural reconstruction were corrected during the 1950s. However, it has come to be believed, following Straus and Cave (1957), that Boule's errors of reconstruction were due to the diseased condition of the La Chapelle-aux-Saints 1 remains, rather than to Boule's misinterpretation of morphology. The abnormalities on the La Chapelle-aux-Saints 1 postcranium include: lower cervical, upper thoracic, and lower thoracic intervertebral degenerative joint disease (DJD), a distal fracture of a mid-thoracic rib, extensive DJD of the left hip, DJD of the right fifth proximal interphalangeal articulation, bilateral humeral head eburnation, and minor exostosis formation on the right humerus, ulna, and radius. These were associated with extensive alveolar inflammation including apical abscesses and antemortem tooth loss, some temporomandibular DJD, bilateral auditory exostoses, and minimal occipital condyle DJD. None of these abnormalities significantly affected Boule's Neandertal postural reconstruction, and a review of his analysis indicates that early twentieth century interpretations of skeletal morphology (primarily of the cranium, cervical vertebrae, lumbar and sacral vertebrae, proximal femora and tibiae, posterior tarsals, and hallucial tarsometatarsal joint), combined with Boule's evolutionary preconceptions, were responsible for his mistaken view of Neandertal posture.