Manual Loading Distribution During Carrying Behaviors: Implications for the Evolution of the Hominin Hand

The human hand is unparalleled amongst primates in its ability to manipulate objects forcefully and dexterously. Previous research has predominantly sought to explain the evolution of these capabilities through an adaptive relationship between more modern human-like anatomical features in the upper limb and increased stone tool production and use proficiency. To date, however, we know little about the influence that other manipulatively demanding behaviors may have had upon the evolution of the human hand. The present study addresses one aspect of this deficiency by examining the recruitment of the distal phalanges during a range of manual transportation (i.e. carrying) events related to hominin behavioral repertoires during the Plio-Pleistocene. Specifically, forces on the volar pad of each digit are recorded during the transportation of stones and wooden branches that vary in weight and size. Results indicate that in most instances, the index and middle fingers are recruited to a significantly greater extent than the other three digits during carrying events. Relative force differences between digits were, however, dependent upon the size and weight of the object transported. Carrying behaviors therefore appear unlikely to have contributed to the evolution of the robust thumb anatomy observed in the human hand. Rather, results suggest that the manual transportation of objects may plausibly have influenced the evolution of the human gripping capabilities and the 3rd metacarpal styloid process

Investigating interrelationships between Lower Palaeolithic stone tool effectiveness and tool user biometric variation: implications for technological and evolutionary changes

Lower Palaeolithic hominins are thought to have been dependent upon stone tools during the acquisition and processing of food resources. Hence, it is hypothesized that the evolutionary advantages provided by efficient stone tool use may have selected for anatomical changes observed in the hand during this period. Similarly, hominin manipulative capabilities are suggested to have been of consequence to Lower Palaeolithic technological choices and tool-use capabilities. The extent and character of these relationships are not, however, fully understood and it is not known whether these hypothesized co-evolutionary and co-dependent relationships are consistent across varying technological and task-type conditions. Here, six key biometric parameters of the hand are investigated in terms of their statistical relationship with cutting efficiency using both flakes and handaxes over extended periods of use and in multiple types of cutting task. Results indicate that (1) both handaxe and flake cutting efficiency is significantly related with biometric variation of individual tool-users, (2) relationships between biometric parameters and efficiency are consistent across extended durations but vary dependent upon task-type conditions, (3) manipulative strength is the most significant biometric trait in terms of predicting flake efficiency, while (4) hand size is the strongest predictor of handaxe cutting efficiency. These results demonstrate the long-term impact that stone tool use likely had on the evolution of hominin biometric variation during the Lower Palaeolithic, while also highlighting the variable influence of different tool use contexts. Most notably, results indicate that the onset of the Acheulean may have been dependent, a priori, upon hand dimensions that are close to the modern human range, and that prior to the appearance of this anatomy, handaxe use would have been an impractical (i.e. inefficient) tool use behaviour compared to the use of flakes

Early stage blunting causes rapid reductions in stone tool performance

Palaeolithic stone technologies have never been investigated in terms of how sharpness influences their ability to cut. In turn, there is little understanding of how quickly stone cutting edges blunt, how past populations responded to any consequent changes in performance, or how these factors influenced the Palaeolithic archaeological record. Presented here is experimental data quantitatively detailing how variation in edge sharpness influences stone tool cutting performance. Significant increases in force (N) and material displacement (mm) requirements occur rapidly within early stages of blunting, with a single abrasive cutting stroke causing, on average, a 38% increase in the force needed to initiate a cut. In energetic terms, this equates to a 70% increase in work (J). Subsequent to early stages of blunting we identify a substantial drop in the impact of additional edge abrasion. We also demonstrate how edge (included) angle significantly influences cutting force and energy requirements and how it co-varies with sharpness. Amongst other conclusions, we suggest that rapid reductions in performance due to blunting may account for the abundance of lithic artefacts at some archaeological sites, the speed that resharpening behaviours altered tool forms, and the lack of microscopic wear traces on many lithic implements

Torque creation and force variation along the cutting edges of Acheulean handaxes: implications for tip thinning, resharpening and tranchet flake removals

One of the defining characteristics of Acheulean handaxes is the presence of a substantial length of sharp cutting edge, often covering the majority or entirety of their plan-form outline. Recently, factors affecting the efficiency and effectiveness of handaxes for cutting have come under increased scrutiny. Most studies investigate how shape, size, symmetry and other metrics influence cutting performance characteristics. This includes investigations of edge morphology. To date, it is unknown how cutting performance may vary within an individual handaxe dependent on which aspect of its edge is used. Here, we experimentally investigate how loading capabilities (applied forces) vary along the edges of handaxes, from tip to base. Significant differences were identified dependent on the edge-point loaded, with greater forces recorded at the tip of tools relative to more proximally located edges. Notably, at ~20% of a handaxe's length away from the tip, loading levels were reduced by around 24%. Acheulean hominins concerned with maximising cutting stress potential during tool use should, therefore, have preferentially used the tip portion of handaxes when possible. During broader, sweeping cutting motions that use substantial lengths of cutting edge, our data suggest different portions of the edge create variable cutting-stress levels. Such differences likely derive from increases and decreases in torque creation, and the interaction between cutting forces and ergonomic relationships at the hand-tool interface. We discuss how these relationships may have influenced handaxe design during the Acheulean period, including tip focused modifications such as tranchet flake removals, thinning, and increased resharpening

Form and function in the Lower Palaeolithic: history, progress, and continued relevance

Percussively flaked stone artefacts constitute a major source of evidence relating to hominin behavioural strategies and are, essentially, a product or byproduct of a past individual’s decision to create a tool with respect to some broader goal. Moreover, it has long been noted that both differences and recurrent regularities exist within and between Palaeolithic stone artefact forms. Accordingly, archaeologists have frequently drawn links between form and functionality, with functional objectives and performance often being regarded consequential to a stone tool’s morphological properties. Despite these factors, extensive reviews of the related concepts of form and function with respect to the Lower Palaeolithic remain surprisingly sparse. We attempt to redress this issue. First we stress the historical place of form–function concepts, and their role in establishing basic ideas that echo to this day. We then highlight methodological and conceptual progress in determining artefactual function in more recent years. Thereafter, we evaluate four specific issues that are of direct consequence for evaluating the ongoing relevance of form–function concepts, especially with respect to their relevance for understanding human evolution more generally. Our discussion highlights specifically how recent developments have been able to build on a long historical legacy, and demonstrate that direct, indirect, experimental, and evolutionary perspectives intersect in crucial ways, with each providing specific but essential insights for ongoing questions. We conclude by emphasising that our understanding of these issues and their interaction, has been, and will be, essential to accurately interpret the Lower Palaeolithic archaeological record, tool-form related behaviours of Lower Palaeolithic hominins, and their consequences for (and relationship to) wider questions of human evolution

Hand grip diversity and frequency during the use of Lower Palaeolithic stone cutting-tools

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 facilitated by manipulative capabilities consistent with the evolutionary trajectory of the hominin hand during this period. Grip analyses have provided important contributions to this understanding. To date, however, there has been no large-scale investigation of grip diversity during flaked stone-tool use, empirical comparative analyses of grip use frequencies, or examination of ergonomic relationships between grip choice and stone tool type and form. Here, we conduct four experimental studies, using replica Lower Palaeolithic stone tools in a series of actualistic and laboratory-based contexts, to record grip type and frequency of grip use during 1067 stone tool-use events by 123 individuals. Using detailed morphometric data recorded from each tool, we demonstrate how grip choice varies according to the type and form of stone tool used, and how these relationships differ between tool-use contexts. We identify 29 grip types across all tool-use events, with significant differences recorded in their frequency of use dependent on tool type, tool form, and the context of use. Despite the influence of these three factors, there is consistency in the frequent use of a limited number (?4) of grip types within each experiment and the consistent and seemingly forceful recruitment of the thumb and index finger. Accordingly, we argue that there are deep-rooted, ergonomically-related, regularities in how stone tools are gripped during their use, that these regularities may have been present during the use of stone tools by Plio-Pleistocene hominins, and any subsequent selective pressures would likely have been focused on the first and second digit

A citation network analysis of lithic microwear research

The introduction of lithic microwear research into the wider archaeological community by Keeley (1980) was concurrent with the development of the processual paradigm and the adoption of the scientific method. Subsequently, lithic microwear research has benefited from over 35 years of innovation, including the introduction of novel methodological and analytical procedures. The present study employs a citation network to objectively analyse the development of microwear research. Given developments in technology, as well as the institutional isolation of early microwear research, the present analysis considers the citation network that stems from Keeley's seminal 1980 volume. The 363 papers identified as having cited Keeley (1980) in the subsequent 35 years were treated as individual nodes within the citation network. Before analysis, nodes were assigned attributes, including the type of research published and whether they were supportive of three key aspects of Keeley's experimental program: the ability to determine the function of the tool and to ascertain the type of worked material from microwear, as well as the use of high-powered microscopy techniques. Emergent properties of the papers, including closeness centrality, indegree and betweenness centrality, are used to test for significant differences between paper attributes. Similarly a clustering algorithm is used to objectively define distinct clusters of important papers within the discipline. Results indicate that a small number of nodes in the network maintain statistically significant influence on the form of the citation network. These important nodes and the distinct ‘schools of thought’ identified are discussed in the context of Keeley's initial contribution to the sub-field

The unexpected importance of the fifth digit during stone tool production

Unique anatomical features of the human hand facilitate our ability to proficiently and forcefully perform precision grips and in-hand manipulation of objects. Extensive research has been conducted into the role of digits one to three during these manual behaviours, and the origin of the highly derived first digit anatomy that facilitates these capabilities. Stone tool production has long been thought a key influence in this regard. Despite previous research stressing the unique derived morphology of the human fifth digit little work has investigated why humans alone display these features. Here we examine the recruitment frequency, loading magnitude, and loading distribution of all digits on the non-dominant hand of skilled flintknappers during four technologically distinct types of Lower Palaeolithic stone tool production. Our data reveal the fifth digit to be heavily and frequently recruited during all studied behaviours. It occasionally incurred pressures, and was used in frequencies, greater or equal to those of the thumb, and frequently the same or greater than those of the index finger. The fifth digit therefore appears key to >2 million years of stone tool production activities, a behaviour that likely contributed to the derived anatomy observed in the modern human fifth ray

Muscle recruitment and stone tool use ergonomics across three million years of Palaeolithic technological transitions

Ergonomic relationships that minimize muscle activity relative to the creation of cutting stress underpin the design of modern knives, saws, and axes. The Palaeolithic archaeological record, and the > 3 million years of technological behavior that it represents, is predominantly characterized by sharp stone implements used for cutting. To date, we do not know whether Palaeolithic hominins adhered to ergonomic principles when designing stone tools, if lithic technological transitions were linked to ease-of-use advances, or even how muscularly demanding different Palaeolithic tools are on an empirically defined relative basis. Here, we report the results of an experimental program that examines how four key stone tool types, produced between ∼ 3.3 million and ∼ 40 thousand years ago, influence muscle activation in the hominin upper limb. Using standardized laboratory-based tests designed to imitate Pleistocene cutting behaviors, surface electromyography recorded electrical activity (amplitude) in nine muscles across the hand, forearm and shoulder of modern humans during the use of replica Lomekwian, Oldowan, Acheulean and Mousterian stone tools. Results confirm digit flexors and abductors, particularly the first dorsal interosseous and flexor pollicis longus, to be the most heavily recruited muscles during the use of all tool types. Significant differences in muscle activation are, however, identified dependent on the type of stone tool used. Notably, the abductor digiti minimi, flexor pollicis longus, and biceps brachii were highly activated during handaxe use, particularly when compared to the use of Oldowan and Levallois flakes. Results are discussed in light of current understanding on the origin of Lower and Middle Palaeolithic technologies, why specific tool types were produced over others during these periods, and the extent to which early hominins produced ergonomically designed tools

Signals of locomotion and manipulation in the internal trabecular bone structure of extant hominoids and fossil hominins

The enhanced dexterity of the human hand is unique among primates, an ability that is traditionally thought to have evolved in response to tool-related behaviours and a release from the biomechanical constraints of locomotion in our bipedal hominin ancestors. However, recent fossil and archaeological evidence, as well as novel analyses, suggest that dexterity-related morphology and abilities evolved earlier that traditionally thought and that fossil hominins used their hands for locomotion until much later than presumed