Inferring cultural reproduction from lithic data: A critical review.

The cultural reproduction of lithic technology, long an implicit assumption of archaeological theories, has garnered increasing attention over the past decades. Major debates ranging from the origins of the human culture capacity to the interpretation of spatiotemporal patterning now make explicit reference to social learning mechanisms and cultural evolutionary dynamics. This burgeoning literature has produced important insights and methodological innovations. However, this rapid growth has sometimes led to confusion and controversy due to an under-examination of underlying theoretical and methodological assumptions. The time is thus ripe for a critical assessment of progress in the study of the cultural reproduction of lithic technology. Here we review recent work addressing the evolutionary origins of human culture and the meaning of artifact variation at both intrasite and intersite levels. We propose that further progress will require a more extended and context-specific evolutionary approach to address the complexity of real-world cultural reproduction.

Knowledge vs. know-how? Dissecting the foundations of stone knapping skill.

Stone tools provide some of the best remaining evidence of behavioral change over long periods, but their cognitive and evolutionary implications remain poorly understood. Here, we contribute to a growing body of experimental research on the cognitive and perceptual-motor foundations of stone toolmaking skills by using a flake prediction paradigm to assess the relative importance of technological understanding vs. accurate action execution in Late Acheulean-style handaxe production. This experiment took place as part of a larger, longitudinal study of knapping skill acquisition, allowing us to assemble a large sample of predictions across learning stages and in a comparative sample of experts. By combining group and individual-level statistical analyses with predictive modeling, we show that understanding and predicting specific flaking outcomes in this technology is both more difficult and less important than expected from previous work. Instead, our findings reveal the critical importance of perceptual motor skills needed to manage speed-accuracy trade-offs and reliably detach the large, invasive flakes that enable bifacial edging and thinning. With practice, novices increased striking accuracy, flaking success rates, and (to an extent) handaxe quality by targeting small flakes with acute platform angles. However, only experts were able to combine percussive force and accuracy to produce results comparable with actual Late Acheulean handaxes. The relatively intense demands for accurate action execution documented in our study indicate that biomechanical properties of the upper limb, cortical and cerebellar systems for sensorimotor control, and the cognitive, communicative, and affective traits supporting deliberate practice would all have been likely targets of selection acting on Late Acheulean toolmaking aptitude.

Evolutionary neuroscience of cumulative culture.

Culture suffuses all aspects of human life. It shapes our minds and bodies and has provided a cumulative inheritance of knowledge, skills, institutions, and artifacts that allows us to truly stand on the shoulders of giants. No other species approaches the extent, diversity, and complexity of human culture, but we remain unsure how this came to be. The very uniqueness of human culture is both a puzzle and a problem. It is puzzling as to why more species have not adopted this manifestly beneficial strategy and problematic because the comparative methods of evolutionary biology are ill suited to explain unique events. Here, we develop a more particularistic and mechanistic evolutionary neuroscience approach to cumulative culture, taking into account experimental, developmental, comparative, and archaeological evidence. This approach reconciles currently competing accounts of the origins of human culture and develops the concept of a uniquely human technological niche rooted in a shared primate heritage of visuomotor coordination and dexterous manipulation.

Understanding stone tool-making skill acquisition: Experimental methods and evolutionary implications.

Despite its theoretical importance, the process of stone tool-making skill acquisition remains understudied and poorly understood. The challenges and costs of skill learning constitute an oft-neglected factor in the evaluation of alternative adaptive strategies and a potential source of bias in cultural transmission. Similarly, theory and data indicate that the most salient neural and cognitive demands of stone tool-making should occur during learning rather than expert performance. Unfortunately, the behavioral complexity and extensive learning requirements that make stone knapping skill acquisition an interesting object of study are the very features that make it so challenging to investigate experimentally. Here we present results from a multidisciplinary study of Late Acheulean handaxe-making skill acquisition involving twenty-six naïve participants and up to 90 hours training over several months, accompanied by a battery of psychometric, behavioral, and neuroimaging assessments. In this initial report, we derive a robust quantitative skill metric for the experimental handaxes using machine learning algorithms, reconstruct a group-level learning curve, and explore sources of individual variation in learning outcomes. Results identify particular cognitive targets of selection on the efficiency or reliability of tool-making skill acquisition, quantify learning costs, highlight the likely importance of social support, motivation, persistence, and self-control in knapping skill acquisition, and illustrate methods for reliably reconstructing ancient learning processes from archaeological evidence.

Virtual dissection and comparative connectivity of the superior longitudinal fasciculus in chimpanzees and humans.

Many of the behavioral capacities that distinguish humans from other primates rely on fronto-parietal circuits. The superior longitudinal fasciculus (SLF) is the primary white matter tract connecting lateral frontal with lateral parietal regions; it is distinct from the arcuate fasciculus, which interconnects the frontal and temporal lobes. Here we report a direct, quantitative comparison of SLF connectivity using virtual in vivo dissection of the SLF in chimpanzees and humans. SLF I, the superior-most branch of the SLF, showed similar patterns of connectivity between humans and chimpanzees, and was proportionally volumetrically larger in chimpanzees. SLF II, the middle branch, and SLF III, the inferior-most branch, showed species differences in frontal connectivity. In humans, SLF II showed greater connectivity with dorsolateral prefrontal cortex, whereas in chimps SLF II showed greater connectivity with the inferior frontal gyrus. SLF III was right-lateralized and proportionally volumetrically larger in humans, and human SLF III showed relatively reduced connectivity with dorsal premotor cortex and greater extension into the anterior inferior frontal gyrus, especially in the right hemisphere. These results have implications for the evolution of fronto-parietal functions including spatial attention to observed actions, social learning, and tool use, and are in line with previous research suggesting a unique role for the right anterior inferior frontal gyrus in the evolution of human fronto-parietal network architecture.

Differences in neural activation for object-directed grasping in chimpanzees and humans.

The human faculty for object-mediated action, including tool use and imitation, exceeds that of even our closest primate relatives and is a key foundation of human cognitive and cultural uniqueness. In humans and macaques, observing object-directed grasping actions activates a network of frontal, parietal, and occipitotemporal brain regions, but differences in human and macaque activation suggest that this system has been a focus of selection in the primate lineage. To study the evolution of this system, we performed functional neuroimaging in humans' closest living relatives, chimpanzees. We compare activations during performance of an object-directed manual grasping action, observation of the same action, and observation of a mimed version of the action that consisted of only movements without results. Performance and observation of the same action activated a distributed frontoparietal network similar to that reported in macaques and humans. Like humans and unlike macaques, these regions were also activated by observing movements without results. However, in a direct chimpanzee/human comparison, we also identified unique aspects of human neural responses to observed grasping. Chimpanzee activation showed a prefrontal bias, including significantly more activity in ventrolateral prefrontal cortex, whereas human activation was more evenly distributed across more posterior regions, including significantly more activation in ventral premotor cortex, inferior parietal cortex, and inferotemporal cortex. This indicates a more "bottom-up" representation of observed action in the human brain and suggests that the evolution of tool use, social learning, and cumulative culture may have involved modifications of frontoparietal interactions.

Neuroplasticity enables bio-cultural feedback in Paleolithic stone-tool making.

Stone-tool making is an ancient human skill thought to have played a key role in the bio-cultural co-evolutionary feedback that produced modern brains, culture, and cognition. To test the proposed evolutionary mechanisms underpinning this hypothesis we studied stone-tool making skill learning in modern participants and examined interactions between individual neurostructural differences, plastic accommodation, and culturally transmitted behavior. We found that prior experience with other culturally transmitted craft skills increased both initial stone tool-making performance and subsequent neuroplastic training effects in a frontoparietal white matter pathway associated with action control. These effects were mediated by the effect of experience on pre-training variation in a frontotemporal pathway supporting action semantic representation. Our results show that the acquisition of one technical skill can produce structural brain changes conducive to the discovery and acquisition of additional skills, providing empirical evidence for bio-cultural feedback loops long hypothesized to link learning and adaptive change.

On the psychological origins of tool use.

The ubiquity of tool use in human life has generated multiple lines of scientific and philosophical investigation to understand the development and expression of humans' engagement with tools and its relation to other dimensions of human experience. However, existing literature on tool use faces several epistemological challenges in which the same set of questions generate many different answers. At least four critical questions can be identified, which are intimately intertwined-(1) What constitutes tool use? (2) What psychological processes underlie tool use in humans and nonhuman animals? (3) Which of these psychological processes are exclusive to tool use? (4) Which psychological processes involved in tool use are exclusive to Homo sapiens? To help advance a multidisciplinary scientific understanding of tool use, six author groups representing different academic disciplines (e.g., anthropology, psychology, neuroscience) and different theoretical perspectives respond to each of these questions, and then point to the direction of future work on tool use. We find that while there are marked differences among the responses of the respective author groups to each question, there is a surprising degree of agreement about many essential concepts and questions. We believe that this interdisciplinary and intertheoretical discussion will foster a more comprehensive understanding of tool use than any one of these perspectives (or any one of these author groups) would (or could) on their own.

Technology, expertise and social cognition in human evolution.

Paleolithic stone tools provide concrete evidence of major developments in human behavioural and cognitive evolution. Of particular interest are evolving cognitive mechanisms implied by the cultural transmission of increasingly complex prehistoric technologies, hypothetically including motor resonance, causal reasoning and mentalizing. To test the relevance of these mechanisms to specific Paleolithic technologies, we conducted a functional magnetic resonance imaging study of Naïve, Trained and Expert subjects observing two toolmaking methods of differing complexity and antiquity: the simple 'Oldowan' method documented by the earliest tools 2.5 million years ago; and the more complex 'Acheulean' method used to produce refined tools 0.5 million years ago. Subjects observed 20-s video clips of an expert demonstrator, followed by behavioural tasks designed to maintain attention. Results show that observational understanding of Acheulean toolmaking involves increased demands for the recognition of abstract technological intentions. Across subject groups, Acheulean compared with Oldowan toolmaking was associated with activation of left anterior intraparietal and inferior frontal sulci, indicating the relevance of resonance mechanisms. Between groups, Naïve subjects relied on bottom-up kinematic simulation in the premotor cortex to reconstruct unfamiliar intentions, and Experts employed a combination of familiarity-based sensorimotor matching in the posterior parietal cortex and top-down mentalizing involving the medial prefrontal cortex. While no specific differences between toolmaking technologies were found for Trained subjects, both produced frontal activation relative to Control, suggesting focused engagement with toolmaking stimuli. These findings support motor resonance hypotheses for the evolutionary origins of human social cognition and cumulative culture, directly linking these hypotheses with archaeologically observable behaviours in prehistory.

The cognitive science of technology.

Technology is central to human life but hard to define and study. This review synthesizes advances in fields from anthropology to evolutionary biology and neuroscience to propose an interdisciplinary cognitive science of technology. The foundation of this effort is an evolutionarily motivated definition of technology that highlights three key features: material production, social collaboration, and cultural reproduction. This broad scope respects the complexity of the subject but poses a challenge for theoretical unification. Addressing this challenge requires a comparative approach to reduce the diversity of real-world technological cognition to a smaller number of recurring processes and relationships. To this end, a synthetic perceptual-motor hypothesis (PMH) for the evolutionary-developmental-cultural construction of technological cognition is advanced as an initial target for investigation.

The measurement, evolution, and neural representation of action grammars of human behavior.

Human behaviors from toolmaking to language are thought to rely on a uniquely evolved capacity for hierarchical action sequencing. Testing this idea will require objective, generalizable methods for measuring the structural complexity of real-world behavior. Here we present a data-driven approach for extracting action grammars from basic ethograms, exemplified with respect to the evolutionarily relevant behavior of stone toolmaking. We analyzed sequences from the experimental replication of ~ 2.5 Mya Oldowan vs. ~ 0.5 Mya Acheulean tools, finding that, while using the same "alphabet" of elementary actions, Acheulean sequences are quantifiably more complex and Oldowan grammars are a subset of Acheulean grammars. We illustrate the utility of our complexity measures by re-analyzing data from an fMRI study of stone toolmaking to identify brain responses to structural complexity. Beyond specific implications regarding the co-evolution of language and technology, this exercise illustrates the general applicability of our method to investigate naturalistic human behavior and cognition.

Emergence of perceptuomotor relationships during paleolithic stone toolmaking learning: intersections of observation and practice.

Stone toolmaking is a human motor skill which provides the earliest archeological evidence motor skill and social learning. Intentionally shaping a stone into a functional tool relies on the interaction of action observation and practice to support motor skill acquisition. The emergence of adaptive and efficient visuomotor processes during motor learning of such a novel motor skill requiring complex semantic understanding, like stone toolmaking, is not understood. Through the examination of eye movements and motor skill, the current study sought to evaluate the changes and relationship in perceptuomotor processes during motor learning and performance over 90 h of training. Participants' gaze and motor performance were assessed before, during and following training. Gaze patterns reveal a transition from initially high gaze variability during initial observation to lower gaze variability after training. Perceptual changes were strongly associated with motor performance improvements suggesting a coupling of perceptual and motor processes during motor learning.

Technological variation in the earliest Oldowan from Gona, Afar, Ethiopia.

Inter-site technological variation in the archaeological record is one of the richest potential sources of information about Plio-Pleistocene hominid behavior and evolution. However, appropriate methods for describing and comparing Oldowan assemblages have yet to be agreed upon, and interpretation of the early record remains highly controversial. Particularly salient is disagreement over whether the Oldowan is a single technological phenomenon or is more accurately divided into multiple regional and/or chronological traditions, perhaps including a less developed Pre-Oldowan phase in the late Pliocene. Some of this disagreement reflects theoretical and methodological differences between research traditions and some is more directly evidential. Here we present a framework for describing and interpreting Oldowan variation and apply it to three Pliocene assemblages (EG-10, EG-12, and OGS-7) from Gona, all dated to c. 2.6 million years (Ma). Results indicate proficient knapping and a full range of Oldowan reduction strategies in these earliest known occurrences, consistent with the idea of an Oldowan "technological stasis" from 2.6-1.6 Ma. Patterns of variation in raw material selection and predominant reduction strategy at each site clearly indicate the importance of cultural transmission in the Oldowan, but confounding ecological and economic variation continue to render interpretation in terms of multiple tool making traditions or species inappropriate. We propose that cultural transmission and ecological adaptation should be recognized as complementary, rather than mutually exclusive, mechanisms in future attempts to explain Oldowan technological variation.

Co-occurrence of Acheulian and Oldowan artifacts with Homo erectus cranial fossils from Gona, Afar, Ethiopia.

Although stone tools generally co-occur with early members of the genus Homo, they are rarely found in direct association with hominins. We report that both Acheulian and Oldowan artifacts and Homo erectus crania were found in close association at 1.26 million years (Ma) ago at Busidima North (BSN12), and ca. 1.6 to 1.5 Ma ago at Dana Aoule North (DAN5) archaeological sites at Gona, Afar, Ethiopia. The BSN12 partial cranium is robust and large, while the DAN5 cranium is smaller and more gracile, suggesting that H. erectus was probably a sexually dimorphic species. The evidence from Gona shows behavioral diversity and flexibility with a lengthy and concurrent use of both stone technologies by H. erectus, confounding a simple "single species/single technology" view of early Homo.

The evolutionary neuroscience of tool making.

The appearance of the first intentionally modified stone tools over 2.5 million years ago marked a watershed in human evolutionary history, expanding the human adaptive niche and initiating a trend of technological elaboration that continues to the present day. However, the cognitive foundations of this behavioral revolution remain controversial, as do its implications for the nature and evolution of modern human technological abilities. Here we shed new light on the neural and evolutionary foundations of human tool making skill by presenting functional brain imaging data from six inexperienced subjects learning to make stone tools of the kind found in the earliest archaeological record. Functional imaging of this complex, naturalistic task was accomplished through positron emission tomography with the slowly decaying radiological tracer (18)flouro-2-deoxyglucose. Results show that simple stone tool making is supported by a mosaic of primitive and derived parietofrontal perceptual-motor systems, including recently identified human specializations for representation of the central visual field and perception of three-dimensional form from motion. In the naïve tool makers reported here, no activation was observed in prefrontal executive cortices associated with strategic action planning or in inferior parietal cortex thought to play a role in the representation of everyday tool use skills. We conclude that uniquely human capacities for sensorimotor adaptation and affordance perception, rather than abstract conceptualization and planning, were central factors in the initial stages of human technological evolution.

Cognitive demands of lower paleolithic toolmaking.

Stone tools provide some of the most abundant, continuous, and high resolution evidence of behavioral change over human evolution, but their implications for cognitive evolution have remained unclear. We investigated the neurophysiological demands of stone toolmaking by training modern subjects in known Paleolithic methods ("Oldowan", "Acheulean") and collecting structural and functional brain imaging data as they made technical judgments (outcome prediction, strategic appropriateness) about planned actions on partially completed tools. Results show that this task affected neural activity and functional connectivity in dorsal prefrontal cortex, that effect magnitude correlated with the frequency of correct strategic judgments, and that the frequency of correct strategic judgments was predictive of success in Acheulean, but not Oldowan, toolmaking. This corroborates hypothesized cognitive control demands of Acheulean toolmaking, specifically including information monitoring and manipulation functions attributed to the "central executive" of working memory. More broadly, it develops empirical methods for assessing the differential cognitive demands of Paleolithic technologies, and expands the scope of evolutionary hypotheses that can be tested using the available archaeological record.

Stone tools, language and the brain in human evolution.

Long-standing speculations and more recent hypotheses propose a variety of possible evolutionary connections between language, gesture and tool use. These arguments have received important new support from neuroscientific research on praxis, observational action understanding and vocal language demonstrating substantial functional/anatomical overlap between these behaviours. However, valid reasons for scepticism remain as well as substantial differences in detail between alternative evolutionary hypotheses. Here, we review the current status of alternative 'gestural' and 'technological' hypotheses of language origins, drawing on current evidence of the neural bases of speech and tool use generally, and on recent studies of the neural correlates of Palaeolithic technology specifically.

Stone toolmaking and the evolution of human culture and cognition.

Although many species display behavioural traditions, human culture is unique in the complexity of its technological, symbolic and social contents. Is this extraordinary complexity a product of cognitive evolution, cultural evolution or some interaction of the two? Answering this question will require a much better understanding of patterns of increasing cultural diversity, complexity and rates of change in human evolution. Palaeolithic stone tools provide a relatively abundant and continuous record of such change, but a systematic method for describing the complexity and diversity of these early technologies has yet to be developed. Here, an initial attempt at such a system is presented. Results suggest that rates of Palaeolithic culture change may have been underestimated and that there is a direct relationship between increasing technological complexity and diversity. Cognitive evolution and the greater latitude for cultural variation afforded by increasingly complex technologies may play complementary roles in explaining this pattern.