Tracy L Kivell's research while affiliated with University of Kent and other places
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Publications (132)
Objectives:
Recent studies have associated subarticular trabecular bone distribution in the extant hominid first metacarpal (Mc1) with observed thumb use, to infer fossil hominin thumb use. Here, we analyze the entire Mc1 to test for interspecific differences in: (1) the absolute volume of trabecular volume fraction, (2) the distribution of the de...
In 1994, Hunt published the 'postural feeding hypothesis'-a seminal paper on the origins of hominin bipedalism-founded on the detailed study of chimpanzee positional behavior and the functional inferences derived from the upper and lower limb morphology of the Australopithecus afarensis A.L. 288-1 partial skeleton. Hunt proposed a model for underst...
Bipedalism, a defining feature of the human lineage, is thought to have evolved as forests retreated in the late Miocene-Pliocene. Chimpanzees living in analogous habitats to early hominins offer a unique opportunity to investigate the ecological drivers of bipedalism that cannot be addressed via the fossil record alone. We investigated positional...
The modern human hand is an intriguing mix of primitive morphology and derived function. Traditionally, its form and function are explained as a functional “trade‐off” between the requirements of locomotion and manipulation, but recently acquired comparative, experimental and fossil evidence suggests that this functional trade‐off is more complex t...
The calcar femorale is an internal bony structure of the proximal femur considered to be functionally related to bipedal locomotion. Among extant primates, the presence of a calcar femorale has been so far documented in extant humans and Pan and, among extinct hominins, in the Late Miocene Orrorin, in a Pliocene Australopithecus, and in a Middle Pl...
The trabecular bone morphology of adult extant primates has been shown to reflect mechanical loading related to locomotion. However, ontogenetic studies of humans and other mammals suggest an adaptive lag between trabecular bone response and current mechanical loading patterns that could result in adult trabecular bone morphology reflecting juvenil...
Changes in hand morphology throughout human evolution have facilitated the use of forceful pad-to-pad precision grips, contributing to the development of fine motor movement and dexterous manipulation typical of modern humans. Today, variation in human hand function may be affected by demographic and/or lifestyle factors, but these remain largely u...
The morphology of the proximal carpals (scaphoid, lunate, triquetrum) are linked to the range of motion (ROM) at the radiocarpal and midcarpal joints. While the relationship between ROM and habitual locomotor mode is well established, it has yet to be investigated whether relative patterns of internal bone architecture reflect the kinematics and ki...
African apes engage in a distinct form of locomotion called knuckle-walking, but there is much ambiguity as to when and how this locomotor behaviour evolved. This study aims to elucidate potential differences in knuckle-walking elbow posture and loading in African apes through the study of trabecular bone. Using a whole-epiphysis approach, we quant...
Soft and hard tissue structures at the ulna-side of the wrist, such as reduced ulna-triquetrum articulation and the Triangular Fibrocartilage Complex (TFCC), are conspicuous synapomorphies of hominoids [Fig.1]. These structures are historically linked to biomechanics characteristic of the hominoids such as high degrees of ulnar deviation and forear...
Homo naledi fossils from the Rising Star cave system provide important insights into the diversity of hand morphology within the genus Homo. Notably, the pollical (thumb) metacarpal (Mc1) displays an unusual suite of characteristics including a median longitudinal crest, a narrow proximal base, and broad flaring intrinsic muscle flanges. The presen...
Although hand grip strength is critical to the daily lives of humans and our arboreal great ape relatives, the human hand has changed in form and function throughout our evolution due to terrestrial bipedalism, tool use, and directional asymmetry (DA) such as handedness. Here we investigate how hand form and function interact in modern humans to ga...
Morphological variation in the hominoid capitate has been linked to differences in habitual locomotor activity due to its importance in movement and load transfer at the midcarpal joint proximally and carpometacarpal joints distally. Although the shape of bones and their articulations are linked to joint mobility, the internal structure of bones ha...
Modelling a muscle key to flexing the thumb, a new study suggests that the powerful opposability that characterises the dextrous human hand evolved in some of our fossil relatives about two million years ago — a time when tool use became more important.
Much research has debated the technological abilities of Neanderthals relative to those of early modern humans, with a particular focus on subtle differences in thumb morphology and how this may reflect differences in manipulative behaviors in these two species. Here, we provide a novel perspective on this debate through a 3D geometric morphometric...
Haeusler et al. (1) suggest that our analysis (2) of the distribution of relative bone volume across the articular surface (figure 5) does not justify different taxonomic allocations or locomotor classifications. We agree with their first suggestion, and we did not use these data to make direct arguments for the taxonomic attribution of either spec...
Objectives
As is the case among many complex motor tasks that require prolonged practice before achieving expertise, aspects of the biomechanics of knapping vary according to the relative experience/skill level of the practitioner. In archaeological experiments focused on the production of Plio‐Pleistocene stone tools, these skill‐mediated biomecha...
The capitate sits centrally in the wrist. Its morphology plays a key role proximally as the ‘ball’ component of the mobile mid-carpal joint, where it articulates with
the loosely tethered proximal row. Distally, it articulates with the tightly bound distal row and is loaded in tension by the attaching extrinsic and intrinsic ligaments.
Morphologica...
The human lineage is marked by a transition in hand use, from locomotion towards increasingly dexterous manipulation, concomitant with bipedalism. The forceful precision grips used by modern humans probably evolved in the context of tool manufacture and use, but when and how many times hominin hands became principally manipulative remains unresolve...
This chapter presents description and analysis of the metacarpals and manual phalanges from Sterkfontein. Although the morphology is generally similar across the sample where there are duplicates of the same element, there are differences in size that are quite remarkable within the context of all South African hominins. Some very large specimens s...
Bipedalism is a defining trait of the hominin lineage, associated with a transition from a more arboreal to a more terrestrial environment. While there is debate about when modern human-like bipedalism first appeared in hominins, all known South African hominins show morphological adaptations to bipedalism, suggesting that this was their predominan...
The analysis of internal trabecular and cortical bone has been an informative tool for drawing inferences about behaviour in extant and fossil primate taxa. Within the hand, metacarpal bone architecture has been shown to correlate well with primate locomotion; however, the extent of morphological differences across taxa is unexpectedly small given...
As a central component of the midcarpal and carpometacarpal joints, the capitate plays a primary role in primate hand biomechanics. Capitate morphology facilitates mobility of the midcarpal joint in suspensory apes, limits extension in knuckle-walking apes, and in humans stabilises the capitometacarpal joint for tool behaviours. Biomechanical loadi...
Supplementary Online Material.
The impact of hand proportions on tool grip abilities in humans, great apes and fossil hominins: a biomechanical analysis using musculoskeletal simulation.
Ameline Bardo, Laurent Vigouroux, Tracy L. Kivell, Emmanuelle Pouydebat.
Journal of Human Evolution 125 (2018) 106-121.
Approximately 11.6-million-year-old fossils reveal an ape with arms suited to hanging in trees but human-like legs, suggesting a form of locomotion that might push back the timeline for when walking on two feet evolved. Ancient ape fossil reveals a previously unknown form of locomotion.
Objectives: The dexterity of fossil hominins is often inferred by assessing the comparative manual anatomy and behaviors of extant hominids, with a focus on the thumb. The aim of this study is to test whether trabecular structure is consistent with what is currently known about habitually loaded thumb postures across extant hominids.
Materials and...
Introduction
Knowledge of internal finger loading during human and non-human primate activities such as tool use or knuckle-walking has become increasingly important to reconstruct the behaviour of fossil hominins based on bone morphology. Musculoskeletal models have proven useful for predicting these internal loads during human activities, but loa...
Although gorillas rarely use tools in the wild, their manipulative skills during plant processing may be similar to those of other tool-using great apes. Virunga mountain gorillas (Gorilla beringei beringei) are known for the complexity in their methods of thistle and nettle plant preparation in the wild. However, there are no comparable data on fo...
Understanding the manual abilities of fossil hominins has been a focus of palaeoanthropological research for decades. Of interest are the morphological characteristics of the thumb due to its fundamental role in manipulation, particularly that of the trapeziometacarpal joint. Considerably less attention has been given to the thumb metacarpophalange...
Trabecular bone remodels during life in response to loading and thus should, at least in part, reflect potential variation in the magnitude, frequency and direction of joint loading across different hominid species. Here we analyse the trabecular structure across all non‐pollical metacarpal distal heads (Mc2‐5) in extant great apes, expanding on pr...
Previously, a micro-finite element (micro-FE)-based inverse remodelling method was presented in the literature that reconstructs the loading history of a bone based on its architecture alone. Despite promising preliminary results, it remains unclear whether this method is sensitive enough to detect differences of bone loading related to pathologies...
Trabecular ontogeny of great ape third metacarpals
Studies of femoral trabecular structure have shown that the orientation and volume of bone are associated with variation in loading and could be informative about individual joint positioning during locomotion. In this study, we analyse for the first time trabecular bone patterns throughout the femoral head using a whole‐epiphysis approach to inves...
Neandertals disappeared from the fossil record around 40,000 bp, after a demographic history of small and isolated groups with high but variable levels of inbreeding, and episodes of interbreeding with other Paleolithic hominins. It is reasonable to expect that high levels of endogamy could be expressed in the skeleton of at least some Neandertal g...
The suite of anatomical features contributing to the unique gripping capabilities of the modern human hand evolved alongside the proliferation of Lower Palaeolithic flaked tool technologies across the Old World. Experimental studies investigating their potential co-evolution suggest that the use of flakes, handaxes, and other stone tools is facilit...
Force and strain data.
This table presents the force and strain data measured from the three specimens in two tendon path conditions, i.e. parallel and semi-physiological conditions. The force experienced by the third metacarpal bone (Fmc) is presented in the proximal(+)/distal(−), dorsal(+)/volar(−), and radial(+)/ulnar(−) directions with respect...
Background
Musculoskeletal and finite element modelling are often used to predict joint loading and bone strength within the human hand, but there is a lack of in vitro evidence of the force and strain experienced by hand bones.
Methods
This study presents a novel experimental setup that allows the positioning of a cadaveric digit in a variety of...
Load cell accuracy.
The load cell was tested before being used in the experiment. The error was less than 0.05 N and the full-scale accuracy error was less than 0.1%.
MATLAB script for processing the force and strain data.
The acquired force and strain signals were calibrated with baseline data collected when the flexor tendons were not loaded. Then, the signals were filtered using a sixth-order Butterworth low-pass filter with the cut-off frequency at 6 Hz.
Analytical and experimental strain values.
The precision of strain gauge measurement was verified via beam models, and the analytical calculations showed the same strain pattern and similar values as the experimental results. Photo credit: Dr Szu-Ching Lu.
Objectives:
Several studies have investigated potential functional signals in the trabecular structure of the primate proximal humerus but with varied success. Here, we apply for the first time a "whole-epiphyses" approach to analysing trabecular bone in the humeral head with the aim of providing a more holistic interpretation of trabecular variat...
Objectives
Trabecular bone structure is known to be influenced by joint loading during life. However, many additional variables have the potential to contribute to trabecular bone structure of an adult individual, including age, sex, body size, genetics, and overall activity level. There is little research into intraspecific variability in trabecul...
Although gorillas rarely use tools in the wild, their manipulative skills during plant processing may be similar to those of other tool-using great apes. Virunga mountain gorillas (Gorilla beringei beringei) are known for the complexity in their methods of thistle and nettle plant preparation in the wild. However, there are no comparable data on fo...
Background
In addition to external bone shape and cortical bone thickness and distribution, the distribution and orientation of internal trabecular bone across individuals and species has yielded important functional information on how bone adapts in response to load. In particular, trabecular bone analysis has played a key role in studies of human...
Fig. S1. Pan BV/TV and DA distributions in the complete sample.
Fig. S4. Homo BV/TV and DA distributions in the complete sample.
Fig. S3. Pongo BV/TV and DA distributions in the complete sample.
Captive specimens: ZSM 1966 0203 (male) and ZSM 1982 0092 (female).
Table S3. Results (p-value) for between taxa differences in the indices.
Captive Pongo are included.
Table S2. Loadings of parameters at each region to PC1 and PC2.
Fig. S2. Gorilla BV/TV and DA distributions in the complete sample.
Table S1. Results (p-value) for between taxa differences in the examined regions.
Captive Pongo are included.
Trabecular parameters for each specimen.
Hand bone morphology is regularly used to link particular hominin species with behaviors relevant to cognitive/technological progress. Debates about the functional significance of differing hominin hand
bone morphologies tend to rely on establishing phylogenetic relationships and/or inferring behavior from epigenetic variation arising from mechanic...
In the Session XVI-7: Is ‘Neanderthal behaviour' a useful concept? Multiscalar approches to the behavioural diversity of middle palaeolithic neanderthal populations
Biomechanical analyses of arboreal locomotion in great apes in their natural environment are scarce and thus attempts to correlate behavioral and habitat differences with variations in morphology are limited. The aim of this study was to investigate the gait characteristics of vertical climbing in mountain gorillas (Gorilla beringei beringei) and c...
Understanding the timing and character of the expansion of Homo sapiens out of Africa is critical for inferring the colonization and admixture processes that underpin global population history. It has been argued that dispersal out of Africa had an early phase, particularly ~130–90 thousand years ago (ka), that reached only the East Mediterranean L...
Understanding the timing and character of the expansion of Homo sapiens out of Africa is critical for inferring the colonization and admixture processes that underpin global population history. It has been argued that dispersal out of Africa had an early phase, particularly ~130–90 thousand years ago (ka), that reached only the East Mediterranean L...
It is widely agreed that biomechanical stresses imposed by stone tool behaviors influenced the evolution of the human hand. Though archaeological evidence suggests that early hominins participated in a variety of tool behaviors, it is unlikely that all behaviors equally influenced modern human hand anatomy. It is more probable that a behavior's lik...
Evolution of the human hand has undergone a transition from use during locomotion to use primarily for manipulation. Previous comparative morphological and biomechanical studies have focused on potential changes in manipulative abilities during human hand evolution, but few have focused on functional signals for arboreal locomotion. Here, we provid...
Aspects of trabecular bone architecture are thought to reflect regional loading of the skeleton, and thus differ between primate taxa with different locomotor and postural modes. However, there are several systemic factors that affect bone structure that could contribute to, or be the primary factor determining, interspecific differences in bone st...
Twenty-nine carpal bones of Homo neanderthalensis have been recovered from the site of El Sidrón (Asturias, Spain) during excavations between 1994 and 2009, alongside ∼2500 other Neandertal skeletal elements dated to ∼49,000 years ago. All bones of the wrist are represented, including adult scaphoids (n = 6), lunates (n = 2), triquetra (n = 4), pis...
Objectives:
Studies on grasping and limb posture during arboreal locomotion in great apes in their natural environment are scarce and thus, attempts to correlate behavioral and habitat differences with variation in morphology are limited. The aim of this study is to compare hand use and forelimb posture during vertical climbing in wild, habituated...
Biomechanical analyses of great ape arboreal locomotion in a natural environment are scarce, thus limiting attempts to correlate behavioural and habitat differences with variation in skeletal morphology. However, little is known about arboreal grips and hand postures, or how these might vary with forelimb posture during vertical climbing on natural...
Objectives:
Internal bone structure, both cortical and trabecular bone, remodels in response to loading and may provide important information regarding behavior. The foot is well suited to analysis of internal bone structure because it experiences the initial substrate reaction forces, due to its proximity to the substrate. Moreover, as humans and...
\textbf{Objectives:}$ Internal bone structure, both cortical and trabecular bone, remodels in response to loading and may provide important information regarding behavior. The foot is well suited to analysis of internal bone structure because it experiences the initial substrate reaction forces, due to its proximity to the substrate. Moreover, as h...
Cranial and mandibular measurements.
DOI:
http://dx.doi.org/10.7554/eLife.24232.046
Canonical variates analysis of carpal morphology.
DOI:
http://dx.doi.org/10.7554/eLife.24232.048
The Rising Star cave system has produced abundant fossil hominin remains within the Dinaledi Chamber, representing a minimum of 15 individuals attributed to Homo naledi. Further exploration led to the discovery of hominin material, now comprising 131 hominin specimens, within a second chamber, the Lesedi Chamber. The Lesedi Chamber is far separated...
Traits of the LES1 cranium in comparison to H. naledi and other hominin species.
DOI:
http://dx.doi.org/10.7554/eLife.24232.045
Postcranial measurements.
DOI:
http://dx.doi.org/10.7554/eLife.24232.047
Taphonomic observations by specimen from the Lesedi Chamber.
DOI:
http://dx.doi.org/10.7554/eLife.24232.049
Bone form reflects both the genetic profile and behavioural history of an individual. As cortical bone is able to remodel in response to mechanical stimuli, interspecific differences in cortical bone thickness may relate to loading during locomotion or manual behaviours during object manipulation. Here, we test the application of a novel method of...
This chapter reviews our current understanding of the extant primate wrist. Following Wood Jones and Napier, the morphology of the primate wrist can be generally described as primitive. It is this primitiveness that allows for the great degree of versatility in wrist function that primates require to navigate their complex locomotor and manipulativ...
This book demonstrates how the primate hand combines both primitive and novel morphology, both general function with specialization, and both a remarkable degree of diversity within some clades and yet general similarity across many others. Across the chapters, different authors have addressed a variety of specific questions and provided their pers...
Objectives:
Humans display an 85-95% cross-cultural right-hand bias in skilled tasks, which is considered a derived behavior because such a high frequency is not reported in wild non-human primates. Handedness is generally considered to be an evolutionary byproduct of selection for manual dexterity and augmented visuo-cognitive capabilities within...