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

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

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

Miniaturization optimized weapon killing power during the social stress of late pre-contact North America (AD 600-1600)

Before Europeans arrived to Eastern North America, prehistoric, indigenous peoples experienced a number of changes that culminated in the development of sedentary, maize agricultural lifeways of varying complexity. Inherent to these lifeways were several triggers of social stress including population nucleation and increase, intergroup conflict (warfare), and increased territoriality. Here, we examine whether this period of social stress co-varied with deadlier weaponry, specifically, the design of the most commonly found prehistoric archery component in late pre-contact North America: triangular stone arrow tips (TSAT). The examination of modern metal or carbon projectiles, arrows, and arrowheads has demonstrated that smaller arrow tips penetrate deeper into a target than do larger ones. We first experimentally confirm that this relationship applies to arrow tips made from stone hafted onto shafts made from wood. We then statistically assess a large sample (n = 742) of late pre-contact TSAT and show that these specimens are extraordinarily small. Thus, by miniaturizing their arrow tips, prehistoric people in Eastern North America optimized their projectile weaponry for maximum penetration and killing power in warfare and hunting. Finally, we verify that these functional advantages were selected across environmental and cultural boundaries. Thus, while we cannot and should not rule out stochastic, production economizing, or non-adaptive cultural processes as an explanation for TSAT, overall our results are consistent with the hypothesis that broad, socially stressful demographic changes in late pre-contact Eastern North America resulted in the miniaturization–and augmented lethality–of stone tools across the region

Multiple novel prostate cancer susceptibility signals identified by fine-mapping of known risk loci among Europeans

Genome-wide association studies (GWAS) have identified numerous common prostate cancer (PrCa) susceptibility loci. We have fine-mapped 64 GWAS regions known at the conclusion of the iCOGS study using large-scale genotyping and imputation in 25 723 PrCa cases and 26 274 controls of European ancestry. We detected evidence for multiple independent signals at 16 regions, 12 of which contained additional newly identified significant associations. A single signal comprising a spectrum of correlated variation was observed at 39 regions; 35 of which are now described by a novel more significantly associated lead SNP, while the originally reported variant remained as the lead SNP only in 4 regions. We also confirmed two association signals in Europeans that had been previously reported only in East-Asian GWAS. Based on statistical evidence and linkage disequilibrium (LD) structure, we have curated and narrowed down the list of the most likely candidate causal variants for each region. Functional annotation using data from ENCODE filtered for PrCa cell lines and eQTL analysis demonstrated significant enrichment for overlap with bio-features within this set. By incorporating the novel risk variants identified here alongside the refined data for existing association signals, we estimate that these loci now explain ∼38.9% of the familial relative risk of PrCa, an 8.9% improvement over the previously reported GWAS tag SNPs. This suggests that a significant fraction of the heritability of PrCa may have been hidden during the discovery phase of GWAS, in particular due to the presence of multiple independent signals within the same regio

Multiorgan MRI findings after hospitalisation with COVID-19 in the UK (C-MORE): a prospective, multicentre, observational cohort study

Introduction: The multiorgan impact of moderate to severe coronavirus infections in the post-acute phase is still poorly understood. We aimed to evaluate the excess burden of multiorgan abnormalities after hospitalisation with COVID-19, evaluate their determinants, and explore associations with patient-related outcome measures. Methods: In a prospective, UK-wide, multicentre MRI follow-up study (C-MORE), adults (aged ≥18 years) discharged from hospital following COVID-19 who were included in Tier 2 of the Post-hospitalisation COVID-19 study (PHOSP-COVID) and contemporary controls with no evidence of previous COVID-19 (SARS-CoV-2 nucleocapsid antibody negative) underwent multiorgan MRI (lungs, heart, brain, liver, and kidneys) with quantitative and qualitative assessment of images and clinical adjudication when relevant. Individuals with end-stage renal failure or contraindications to MRI were excluded. Participants also underwent detailed recording of symptoms, and physiological and biochemical tests. The primary outcome was the excess burden of multiorgan abnormalities (two or more organs) relative to controls, with further adjustments for potential confounders. The C-MORE study is ongoing and is registered with ClinicalTrials.gov, NCT04510025. Findings: Of 2710 participants in Tier 2 of PHOSP-COVID, 531 were recruited across 13 UK-wide C-MORE sites. After exclusions, 259 C-MORE patients (mean age 57 years [SD 12]; 158 [61%] male and 101 [39%] female) who were discharged from hospital with PCR-confirmed or clinically diagnosed COVID-19 between March 1, 2020, and Nov 1, 2021, and 52 non-COVID-19 controls from the community (mean age 49 years [SD 14]; 30 [58%] male and 22 [42%] female) were included in the analysis. Patients were assessed at a median of 5·0 months (IQR 4·2–6·3) after hospital discharge. Compared with non-COVID-19 controls, patients were older, living with more obesity, and had more comorbidities. Multiorgan abnormalities on MRI were more frequent in patients than in controls (157 [61%] of 259 vs 14 [27%] of 52; p<0·0001) and independently associated with COVID-19 status (odds ratio [OR] 2·9 [95% CI 1·5–5·8]; padjusted=0·0023) after adjusting for relevant confounders. Compared with controls, patients were more likely to have MRI evidence of lung abnormalities (p=0·0001; parenchymal abnormalities), brain abnormalities (p<0·0001; more white matter hyperintensities and regional brain volume reduction), and kidney abnormalities (p=0·014; lower medullary T1 and loss of corticomedullary differentiation), whereas cardiac and liver MRI abnormalities were similar between patients and controls. Patients with multiorgan abnormalities were older (difference in mean age 7 years [95% CI 4–10]; mean age of 59·8 years [SD 11·7] with multiorgan abnormalities vs mean age of 52·8 years [11·9] without multiorgan abnormalities; p<0·0001), more likely to have three or more comorbidities (OR 2·47 [1·32–4·82]; padjusted=0·0059), and more likely to have a more severe acute infection (acute CRP >5mg/L, OR 3·55 [1·23–11·88]; padjusted=0·025) than those without multiorgan abnormalities. Presence of lung MRI abnormalities was associated with a two-fold higher risk of chest tightness, and multiorgan MRI abnormalities were associated with severe and very severe persistent physical and mental health impairment (PHOSP-COVID symptom clusters) after hospitalisation. Interpretation: After hospitalisation for COVID-19, people are at risk of multiorgan abnormalities in the medium term. Our findings emphasise the need for proactive multidisciplinary care pathways, with the potential for imaging to guide surveillance frequency and therapeutic stratification

Handaxe Shape Variation in a Relative Context

The nature, extent and causes of shape variation within and between Acheulean handaxe assemblages represent one of the most heavily theorised aspects of Lower Palaeolithic archaeology. To date, however, handaxe shape variation has only ever been studied within an artefact-based comparative context. Here, the 2D and 3D shape of 698 Acheulean handaxes, selected from ten assemblages, is contextualised within a theoretically possible range of forms defined by two intentionally highly diverse modern replica biface sets. Results demonstrate that handaxe artefacts are highly diverse in their 2D plan-view shape, displaying near complete overlap with the shape space of the intentionally diverse replica tools, along with similar levels of variation. The 3D shape of handaxe artefacts, however, displays much stronger form limitations, occupying under 50% of the shape space created by the replica bifaces. Principally, flat and more ‘tabular’ handaxe forms that display low thickness to width ratios were revealed as absent from the archaeological record. It is argued that while there is considerable diversity and variability in the shape of Acheulean handaxe artefacts, their form is nonetheless restricted by strong material volume and ‘refinement’ limits

Form and Function in the Lower Palaeolithic

The causes of morphological variation within Lower Palaeolithic stone tool assemblages have been subject to debate for decades. As a result of numerous explanatory hypotheses, it forms one of the most substantial areas of research within Palaeolithic archaeology. To date, however, very little research has ever been undertaken into the functional causes and consequences of Lower Palaeolithic stone tool form variation. Indeed, despite stone tools being functional objects tasked with the cutting and modification of aspects of the physical environment, previous researchers have preferentially sought to explain their morphology as a result of social, aesthetic, cultural, cognitive, reductive, and raw material influences. Here, this imbalance is addressed and the two principal technological components of the Lower Palaeolithic, ‘basic’ flakes and handaxes, are subject to a number of controlled, statistically robust, and archaeologically inferable experiments investigating relationships between variable tool-forms and functional performance characteristics. Results and subsequent discussion identify a number of important evolutionary, behavioural, and technological implications for Lower Palaeolithic hominins