Conceptualizing intelligence in its biological context, as the expression of manifold adaptations, compels a rethinking of measuring this characteristic in humans, relying also on animal studies of analogous skills. Mental manipulation, as an extension of object manipulation, provides a continuous, biologically based concept for studying G as it pertains to individual differences in humans and other species.

The origin of human language is one of the most fascinating and most difficult problems of evolution. Here I argue that pre-hunt communication was the starting context of the evolution of human language. Hunting of big game created a shared interest; animals and hunting actions are easy to imitate; the need to plan created a pressure for increasing complexity; and finally, cultural inheritance of hunting tools and know-how made the transition unique. I further argue that this “first step” was actually (...) a two-stage process where first indexical and iconic signs evolved to coordinate recruitment for the hunt; then later, in the second stage, the complexity of this communication system increased as a response to the increased demand to coordinate group-hunting effort (including division of labor). I provide a review of the fossil record and show that the available evidence is fully compatible with the theory. (shrink)

This book proposes a theory of human cognitive evolution, drawing from paleontology, linguistics, anthropology, cognitive science, and especially neuropsychology. The properties of humankind's brain, culture, and cognition have coevolved in a tight iterative loop; the main event in human evolution has occurred at the cognitive level, however, mediating change at the anatomical and cultural levels. During the past two million years humans have passed through three major cognitive transitions, each of which has left the human mind with a new way (...) of representing reality and a new form of culture. Modern humans consequently have three systems of memory representation that were not available to our closest primate relatives: mimetic skill, language, and external symbols. These three systems are supported by new types of “hard” storage devices, two of which (mimetic and linguistic) are biological, one technological. Full symbolic literacy consists of a complex of skills for interacting with the external memory system. The independence of these three uniquely human ways of representing knowledge is suggested in the way the mind breaks down after brain injury and confirmed by various other lines of evidence. Each of the three systems is based on aninventivecapacity, and the products of those capacities – such as languages, symbols, gestures, social rituals, and images – continue to be invented and vetted in the social arena. Cognitive evolution is not yet complete: the externalization of memory has altered the actual memory architecture within which humans think. This is changing the role of biological memory and the way in which the human brain deploys its resources; it is also changing the form of modern culture. (shrink)

Fundamental to spatial knowledge in all species are the representations underlying object recognition, object search, and navigation through space. But what sets humans apart from other species is our ability to express spatial experience through language. This target article explores the language ofobjectsandplaces, asking what geometric properties are preserved in the representations underlying object nouns and spatial prepositions in English. Evidence from these two aspects of language suggests there are significant differences in the geometric richness with which objects and places (...) are encoded. When an object is named (i.e., with count nouns), detailed geometric properties – principally the object's shape (axes, solid and hollow volumes, surfaces, and parts) – are represented. In contrast, when an object plays the role of either “figure” (located object) or “ground” (reference object) in a locational expression, only very coarse geometric object properties are represented, primarily the main axes. In addition, the spatial functions encoded by spatial prepositions tend to be nonmetric and relatively coarse, for example, “containment,” “contact,” “relative distance,” and “relative direction.” These properties are representative of other languages as well. The striking differences in the way language encodes objects versus places lead us to suggest two explanations: First, there is a tendency for languages to level out geometric detail from both object and place representations. Second, a nonlinguistic disparity between the representations of “what” and “where” underlies how language represents objects and places. The language of objects and places converges with and enriches our understanding of corresponding spatial representations. (shrink)

Wilkins & Wakefield's identification of anatomical features in the Koobi Fora endocast, which may be thought to carry some functional significance in relation to organization for language, raises fundamental problems of method: attention is drawn to some limitations of the evidence, of endocasts and of the neuroanatomical map used to interpret them.

The syntactic structures of natural languages reflect conceptual categories more directly than they reflect communicative categories. This fact supports the main premise of the target article, namely, that the most important event in language evolution was the development of a hierarchical conceptual structure.

Wilkins & Wakefield's intriguing model of language evolution is deficient in evidence of human uniqueness in metaphorical matching, amodal representation, reference, conceptual structure, hierarchical organization, linguistic comprehension, sign use, laterality, and handedness. Primates show communicative reference, laterality, and handedness, and apes in particular show hierarchical organization, conceptual structure, cross-modal abilities, sign use, and displaced reference.

Inferences made from endocasts of fossil skulls cannot provide information on the function of particular neocortical areas or the subcortical pathways to prefrontal cortex that form part of the neural substrate for speech, syntax, and certain aspects of cognition. The neural bases of syntax cannot be disassociated from “communication.” Manual motor control was probably a preadaptive factor in the evolution of humansyntactic ability, but neurophysiological data on living humans show that speech motor control and syntax are more closely linked. The (...) evolution of fully modern speech occurred fairly recently; thoughHomo habilismay have had some degree of speech and syntactic ability, it was not fully modern. Stone tools are uncertain indices of language or cognition. (shrink)

ProbablyHomo habilisis two species not one; similarly for Pan troglodytes. Although amenable to training, in naturePan paniscusmay be a “specialized insular dwarf.” Language is uniquely human, but symbolic behavior and intelligence are widespread among animals with little respect for phylogenetic closeness toHomo sapiens.

Representation must be prior to communication in evolution. Wilkins & Wakefield's target article gives the impression that communicative pressures play a secondary role. We suggest that their evolutionary precursor is compatible with protolanguage rather than language itself. The difference between these two communicative systems should not be underestimated: only the former can be trivially reappropriated from a representational system.

Contra Wilkins & Wakefield, we argue that an evolutionarily inspired approach to language must consider different facets of language (i.e., more than syntax and semantics), and must explore the possibility of nonhuman precursors. Several examples are discussed, illustrating the power of the comparative approach in illuminating our understanding of language evolution.

Wilkins & Wakefield suggest that changes in the hominid brain made it uniquely “preadaptive” for language, yet no precursor functions served as adaptive substrates to the emergence of language. We present contrary evidence that the ability to discriminate and process rapid and complex auditory information is a cross-species function subserving communication processes including, but not limited to, human speech perception. We suggest that auditory temporal processing served as an evolutionary precursor to speech processing and consequent language development in humans.

Hominid-like morphology in habiline cranial endocasts does not necessarily imply the presence of language capacity. The cortical zone in question is not associated exclusively with language in humans, and its emergence in habilines might indicate the evolution of other cognitive functions special to humans that were preconditions for the later evolution of language.

The precise manner in which language serves its communicative function suggests that natural selection, rather than exaptation or reappropriation, played the major role in its evolution. Natural selection is more readily invoked, I suggest, if it is assumed that language originated as a system of manual gestures, and later switched to an oral mode.

Wilkins & Wakefield (1995) provide a thoughtful contribution to our understanding of language origins. In this commentary I attempt to define the relationship between object-manipulation and primate brain function further by reviewing research on aimed throwing and the production and use of stone tools by tufted capuchin monkeys (Cebtis apella). I propose that examining the relation between brain function and object-manipulation inCebuswill provide insight into the preconditions of language in our hominid ancestors.

Chomskyian claims of a genetically hard-wired and cognitively autonomous “universal grammar” are being promoted by generative linguistics as facts about language to the present day. The related doctrine of an evolutionary discontinuity in language emergence, however, is based on misconceptions about the notions of homology and preadaptation. The obvious lack of equivalence between symbolic communicative capacities in existing nonhuman primates and human language does not preclude common roots. Normal and disordered language development is strongly influenced by the genome, but there (...) is no evidence for the existence of specific genes underlying “universal grammar.” In the mature brain, stages of language processing can be distinguished and “first-pass” syntactic analyses appear to precede semantic decoding. However, this partial seriality – as well as behavioral and clinical dissociations between lexical and functional categories – can be best described, not in terms of serially activated and discrete modules, but in terms of classes of cell assemblies that differ in their distributional properties. Whereas cell assemblies involved in semantic interpretation (“content word” assemblies) are widely distributed and are generally less vulnerable to focal lesion, those involved in structural decoding (functor assemblies) are primarily distributed within the left perisylvian cortices and are selectively vulnerable to left perisylvian lesion. These distributional differences are explained in terms of the perceptuomotor components involved in the acquisition of relevant representations. The emerging “motivated” or “toposemantic” brain regional specificity can only be accommodated with “soft” and maturational versions of modularity. The failure to reproduce double dissociations in current connectionist models is due to overly simple neuroscientific assumptions, notably that of overall equipotentiality. Linguistic models should not be expected to be “implemented” in the brain, but need to be constrained by neuroscientific evidence on how biological brains function. (shrink)

Contrary to Müller's claims, and in support of modular theories, genetic factors play a substantial and significant role in language. The finding that some children with specific language impairment (SLI) have nonlinguistic impairments may reflect improper diagnosis of SLI or impairments that are secondary to linguistic impairments. Thus, such findings do not argue against the modularity thesis. The lexical/functional distinction appears to be innate and specifically linguistic and could be instantiated in either symbolic or connectionist systems.

The belief that syntax is an innate, autonomous, species-specific module is highly questionable. Syntax demonstrates the mosaic nature of evolutionary change, in that it made use of (and led to the enhancement of) numerous preexisting neurocognitive features. It is best understood as an emergent characteristic of the explosion of semantic complexity that occurred during hominid evolution.

Müller's target article aims to summarize approaches to the question of how language elements (phonemes, morphemes, etc.) and rules are laid down in the brain. However, it suffers from being too vague about basic assumptions and empirical predictions of neurobiological models, and the empirical evidence available to test the models is not appropriately evaluated. (1) In a neuroscientific model of language, different cortical localizations of words can only be based on biological principles. These need to be made explicit. (2) Evidence (...) for and against word class differences could be evaluated more rigorously. (3) All (and only) humans are able to learn languages with complex syntactic structures; it is, therefore, not appropriate to deny innateness and universality of syntactic principles. The real question appears to be the following: Which neurobiological principles are the linguistic principles based on? (shrink)

I enlarge on the theme that the brain mechanisms required for languageand other aspects of the human mind evolved through selective changes in the regulatory genes governing growth. Extension of the period of postnatal growth increases the role of the environment in structuring the brain, and spatiotemporal programming (heterochrony) ofgrowth might explain hierarchical representation, hemispheric specialization, and perhaps sex differences.

The concepts of the innateness, universality, species-specificity, and autonomy of the human language capacity have had an extreme impact on the psycholinguistic debate for over thirty years. These concepts are evaluated from several neurobiological perspectives, with an emphasis on the emergence of language and its decay due to brain lesion and progressive brain disease.Evidence of perceptuomotor homologies and preadaptations for human language in nonhuman primates suggests a gradual emergence of language during hominid evolution. Regarding ontogeny, the innate component of language (...) capacity is likely to be polygenic and shared with other developmental domains. Dissociations between verbal and nonverbal development are probably rooted in the perceptuomotor specializations of neural substrates rather than the autonomy of a grammar module. Aphasiologicaldata often assumed to suggest modular linguistic subsystems can be accounted for in terms of a neurofunctional model incorporating perceptuomotor-based regional specializationsand distributivity of representations. Thus, dissociations between grammatical functors and content words are due to different conditions of acquisition and resulting differences in neural representation. Human brains are characterized by multifactorial interindividual variability, and strict universality of functional organization is biologically unrealistic.A theoretical alternative is proposed according to which (1) linguistic specialization of brain areas is due to epigenetic and probabilistic maturational events, not to genetic ”hard-wiring,” and (2) linguistic knowledge is neurally represented in distributed cell assemblies whose topography reflects the perceptuomotor modalities involved in the acquisition and use of a given item of knowledge. (shrink)

The cumulative open-endedness of human cultures represents a major break with the social traditions of nonhuman species. As traditions are altered and the modifications retained along the cultural lineage, human populations are capable of producing complex traits that no individual could have figured out on its own. For cultures to produce increasingly complex traditions, improvements and modifications must be kept for the next generations to build upon. High-fidelity transmission would thus act as a ratchet, retaining modifications and allowing the historical (...) buildup of complex traditions. Mechanisms acting against slippage are important, of course, but cultures also need to move forward for the ratchet to retain anything important. In this article, I argue that studies of modification-generating processes and the many ways they shape cumulative culture have been overlooked. Key to a better understanding of cultural modification processes is taking seriously that cultural traditions consist of complex, hierarchically structured recipes. Taking such structures seriously and assessing the different ways they can vary in cultural design space, a novel picture for the onset of cumulative cultural evolution emerges. I argue that a possible impediment for cumulative culture in nonhuman animals may in fact reside not so much in the fidelity of their social transmission but rather in the constraints, internal and external, on their capacity to modify complex, hierarchically structured cultural recipes. (shrink)

Evidence of the pervasiveness of neural reuse in the human brain has forced a revision of the standard conception of modularity in the cognitive sciences. One persistent line of argument against such revision, however, cites the evidence of cognitive dissociations. While this article takes the dissociations seriously, it contends that the traditional modular account is not the best explanation. The key to the puzzle is neural redundancy. The article offers both a philosophical analysis of the relation between reuse and redundancy (...) as well as a plausible solution to the problem of dissociations. (shrink)

This response clarifies the nature of reappropriation and the definition of language. It explicates the relationship between neural systems and language and between homology and evolutionary gradualism. Through a review of ape capacities in the realms of language and tool use, it distinguishes human language acquisition from nonhuman learning. Finally, it suggests the appropriate sorts of evidence on which to base further evolutionary arguments relevant to the origins of language.

This response to continuing commentary addresses brain-hand relationships in Cebus apella (as introduced in West-ergaard's commentary), the evolutionary and acquisition parallels between music and language (suggested by Lynch), and the potential behavioral linguistic consequences of the evolutionary neurobiology in Australopithecus africanus and Homo habilis (discussed by Tobias). Finally, we reiterate the importance of well informed, multidisciplinary approaches to the study of the emergence of human species-specific cognition, especially linguistic capacity.

This target article presents a plausible evolutionary scenario for the emergence of the neural preconditions for language in the hominid lineage. In pleistocene primate lineages there was a paired evolutionary expansion of frontal and parietal neocortex (through certain well-documented adaptive changes associated with manipulative behaviors) resulting, in ancestral hominids, in an incipient Broca's region and in a configurationally unique junction of the parietal, occipital, and temporal lobes of the brain (the POT). On our view, the development of the POT in (...) our ancestors resulted in the neuroanatomical substrate consistent with the ability for representations in modality-neutral association cortex and, as a result of structure-imposing interaction with Broca's area, the hierarchically structured Evidence from paleoneurology and comparative primate neuroanatomy is used to argue that Homo habilis (2.5–2 million years ago) was the first hominid to have the appropriate gross neuroanatomical configuration to support conceptual structure. We thus suggest that the neural preconditions for language are met in H. habilis. Finally, we advocate a theory of language acquisition that uses conceptual structure as input to the learning procedures, thus bridging the gap between it and language. (shrink)

How should biological behaviour be modelled? A relatively new approach is to investigate problems in neuroethology by building physical robot models of biological sensorimotor systems. The explication and justification of this approach are here placed within a framework for describing and comparing models in the behavioural and biological sciences. First, simulation models – the representation of a hypothesis about a target system – are distinguished from several other relationships also termed “modelling” in discussions of scientific explanation. Seven dimensions on which (...) simulation models can differ are defined and distinctions between them discussed: 1. Relevance: whether the model tests and generates hypotheses applicable to biology. 2. Level: the elemental units of the model in the hierarchy from atoms to societies. 3. Generality: the range of biological systems the model can represent. 4. Abstraction: the complexity, relative to the target, or amount of detail included in the model. 5. Structural accuracy: how well the model represents the actual mechanisms underlying the behaviour. 6. Performance match: to what extent the model behaviour matches the target behaviour. 7. Medium: the physical basis by which the model is implemented. No specific position in the space of models thus defined is the only correct one, but a good modelling methodology should be explicit about its position and the justification for that position. It is argued that in building robot models biological relevance is more effective than loose biological inspiration; multiple levels can be integrated; that generality cannot be assumed but might emerge from studying specific instances; abstraction is better done by simplification than idealisation; accuracy can be approached through iterations of complete systems; that the model should be able to match and predict target behaviour; and that a physical medium can have significant advantages. These arguments reflect the view that biological behaviour needs to be studied and modelled in context, that is, in terms of the real problems faced by real animals in real environments. Key Words: animal behaviour; levels; models; neuroethology; realism; robotics; simulation. (shrink)

One can disagree with Müller that it is neurobiologically questionable to suppose that human language is innate, specialized, and species-specific, yet agree that the precise brain mechanisms controlling language in any individual will be influenced by epigenesis and genetic variability, and that the interplay between inherited and acquired aspects of linguistic capacity deserves to be investigated.

Wilkins & Wakefield are clearly right to separate linguistic capacity from communicative ability, if only because other animal species have one without the other. But I question the abruptness of the demarcation they make between a period when hominids evolved enriched conceptual representation for other reasons entirely, and a subsequent later stage when language use became an adaptation.

In their 2002 seminal paper Hauser, Chomsky and Fitch hypothesize that recursion is the only human-specific and language-specific mechanism of the faculty of language. While debate focused primarily on the meaning of recursion in the hypothesis and on the human-specific and syntax-specific character of recursion, the present work focuses on the claim that recursion is language-specific. We argue that there are recursive structures in the domain of motor intentionality by way of extending John R. Searle’s analysis of intentional action. We (...) then discuss evidence from cognitive science and neuroscience supporting the claim that motor-intentional recursion is language-independent and suggest some explanatory hypotheses: linguistic recursion is embodied in sensory-motor processing; linguistic and motor-intentional recursions are distinct and mutually independent mechanisms. Finally, we propose some reflections about the epistemic status of HCF as presenting an empirically falsifiable hypothesis, and on the possibility of testing recursion in different cognitive domains. (shrink)

This article has two goals. First, it synthesizes and critically assesses current scientific knowledge about the cognitive bases of human tool use. Second, it shows how the cognitive traits reviewed help to explain why technological accumulation evolved so markedly in humans, and so modestly in apes.

It has been suggested that hierarchically structured symbols, a remarkable feature of human language, are produced via the operation of recursive combination. Recursive combination is frequently observed in human behavior, not only in language but also in action sequences, mind-reading, technology, et cetera.; in contrast, it is rarely observed in animals. Why is it that only humans use this operation? What is the adaptability of recursive combination? We aim (1) to identify the environmental feature(s) in which recursive combination is effective (...) for survival and reproduction, and that has facilitated the evolution of this ability, and (2) to demonstrate the possible evolutionary processes of recursive combination. To achieve this, we constructed an evolutionary simulation of agents that generated products using recursive combination and used the results to explore the types of fitness functions (that reflect the kinds of adaptive environments) that give rise to this ability. We identified two types of adaptability of the recursive combination: (1) diversifiability of production and (2) diversifiability of products. Through the former, recursive combination promotes robustness against failure of production caused by inaccurate manipulations or irreversible changes. In an environment in which diversified products are preferable, sharing a portion of the production process for these products entails producing multiple products in which recursive combination plays a key role. We suppose that recursive combination works as a driving force of material culture. Finally, we discuss the possible evolutionary scenarios of recursive combination that is later generalized to encompass many aspects of human cognition, including human language. (shrink)

Recent findings indicate that frontal brain asymmetry may be a marker of for depression. However, the psychological predispositions that account linkage between frontal brain asymmetry and depression are unclear. approach-withdrawal hypothesis is the primary framework that has been to account for the linkages between frontal brain asymmetry and or emotional disorders. We review evidence consistent with this and suggest several directions for its extension. One such direction is to constrain the approach-withdrawal hypothesis by linking frontal asymmetry to the known functions (...) of the prefrontal cortex. On this we propose that frontal brain asymmetry may be preferentially linked processes that promote the temporal continuity and shifting of or emotional priorities and the suppression of interference by sources of motivation or emotion. We review evidence from and neurobiological studies of depression that is broadly consistent with these predictions. We emphasise the need for future studies testing our hypotheses. (shrink)

The problem of how certain structure–function composites of high complexity could have evolved gradually and by natural selection has been with us at least since Charles Darwin admitted how difficult it was to explain, “his” theory, the origins of “organs of extreme perfection and complication” – such as the eyes of higher animals. Human language capacity is another evolutionary achievement of extraordinary perfection and complexity. Like other skilled human activities, it involves both central (neural) and peripheral (vocal and respiratory) complexes. (...) The reduction of these to simpler building stones to which evolutionary principles may be applied is staggeringly difficult. (shrink)

The anterior cingulate cortex (ACC)has been identified as part of a supervisoryattentional network for selecting alternativemotor programs in response to top-down corticalprocessing, particularly in situationsinvolving conflicting cognitive tasks.Bilateral lesions to the ACC may be causallyassociated with akinetic mutism, where patientsare unable to voluntarily initiate responses.The clinical and neuroanatomical evidence forthis presumed causal association is examined atlength. However, given the many reciprocalprojections between cerebral, motor, limbic andparalimbic structures within the executivesupervisory network, the association ofvoluntary behavior with a particular structure(the ACC) is (...) highly controversial and thereforepremature at this time. Also considered is theclaim that our subjective sense of voluntarycontrol and free will is simply due toour not having conscious access to theunderlying neural computations that precede our decisions and actions. On thecontrary, the distinction between voluntary and involuntary thoughts and actions mayrather be a matter of temporal and directionallag between parallel computations in differentneural areas. Finally, with reference toDennett, there is an extended discussion ofwhether patients with akinetic mutism are (i) conscious automata, (ii) non-intentional systems, and (iii) in azombie-like state. The relevance of (i)–(iii) for the cognitive neuroscientificliterature is then briefly addressed. (shrink)

The overall goal of this target article is to demonstrate a mechanism for an embodied cognition. The particular vehicle is a much-studied, but still widely debated phenomenon seen in 7–12 month-old-infants. In Piaget's classic “A-not-B error,” infants who have successfully uncovered a toy at location “A” continue to reach to that location even after they watch the toy hidden in a nearby location “B.” Here, we question the traditional explanations of the error as an indicator of infants' concepts of objects (...) or other static mental structures. Instead, we demonstrate that the A-not-B error and its previously puzzling contextual variations can be understood by the coupled dynamics of the ordinary processes of goal-directed actions: looking, planning, reaching, and remembering. We offer a formal dynamic theory and model based on cognitive embodiment that both simulates the known A-not-B effects and offers novel predictions that match new experimental results. The demonstration supports an embodied view by casting the mental events involved in perception, planning, deciding, and remembering in the same analogic dynamic language as that used to describe bodily movement, so that they may be continuously meshed. We maintain that this mesh is a pre-eminently cognitive act of “knowing” not only in infancy but also in everyday activities throughout the life span. Key Words: cognitive development; dynamical systems theory; embodied cognition; infant development; motor control; motor planning; perception and action. (shrink)