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.

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)

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 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)

Connectionist models aiming to reveal the mechanisms of atypical development must in their undamaged form constitute plausible models of normal development and follow a developmental trajectory that matches empirical data. Constructivist models that adapt their structure to the learning task satisfy this demand. They are therefore more informative in the study of atypical development than the static models employed by Thomas & Karmiloff-Smith (T&K-S).

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.

Neuropsychological deficits have been widely used to elucidate normal cognitive functioning. Can patients with such deficits also be used to understand conscious visual experience? In this paper, we ask what it would be like to be a patient with apperceptive agnosia . Philosophical analyses of such questions have suggested that subjectively experiencing what another person experiences would be impossible. Although such roadblocks into the conscious experience of others exist, the experimental study of both patients and neurologically normal subjects can be (...) used to understand visual processing mechanisms. In order to understand the visual processes damaged in apperceptive agnosia, we first review this syndrome and present a case study of one such patient, patient J.W. We then review several theoretical accounts of apperceptive agnosia, and we conclude that studies of the patients themselves may not allow us to discriminate between the various explanations of the syndrome. To test these accounts, we have simulated apperceptive agnosia in neurologically normal subjects. The implications of our results for understanding both apperceptive agnosia and normal visual processing are discussed. (shrink)

It is often assumed that similar domain-specific behavioural impairments found in cases of adult brain damage and developmental disorders correspond to similar underlying causes, and can serve as convergent evidence for the modular structure of the normal adult cognitive system. We argue that this correspondence is contingent on an unsupported assumption that atypical development can produce selective deficits while the rest of the system develops normally (Residual Normality), and that this assumption tends to bias data collection in the field. Based (...) on a review of connectionist models of acquired and developmental disorders in the domains of reading and past tense, as well as on new simulations, we explore the computational viability of Residual Normality and the potential role of development in producing behavioural deficits. Simulations demonstrate that damage to a developmental model can produce very different effects depending on whether it occurs prior to or following the training process. Because developmental disorders typically involve damage prior to learning, we conclude that the developmental process is a key component of the explanation of endstate impairments in such disorders. Further simulations demonstrate that in simple connectionist learning systems, the assumption of Residual Normality is undermined by processes of compensation or alteration elsewhere in the system. We outline the precise computational conditions required for Residual Normality to hold in development, and suggest that in many cases it is an unlikely hypothesis. We conclude that in developmental disorders, inferences from behavioural deficits to underlying structure crucially depend on developmental conditions, and that the process of ontogenetic development cannot be ignored in constructing models of developmental disorders. Key Words: Acquired and developmental disorders; connectionist models; modularity; past tense; reading. (shrink)

Learning the structure of event sequences is a ubiquitous problem in cognition and particularly in language. One possible solution is to learn a probabilistic generative model of sequences that allows making predictions about upcoming events. Though appealing from a neurobiological standpoint, this approach is typically not pursued in connectionist modeling. Here, we investigated a sequential version of the restricted Boltzmann machine, a stochastic recurrent neural network that extracts high-order structure from sensory data through unsupervised generative learning and can encode contextual (...) information in the form of internal, distributed representations. We assessed whether this type of network can extract the orthographic structure of English monosyllables by learning a generative model of the letter sequences forming a word training corpus. We show that the network learned an accurate probabilistic model of English graphotactics, which can be used to make predictions about the letter following a given context as well as to autonomously generate high-quality pseudowords. The model was compared to an extended version of simple recurrent networks, augmented with a stochastic process that allows autonomous generation of sequences, and to non-connectionist probabilistic models. We conclude that sequential RBMs and stochastic simple recurrent networks are promising candidates for modeling cognition in the temporal domain. (shrink)

Understanding how conceptual structures inform stone tool production and use would help us resolve the issue of a pongid-hominid dichotomy in brain organisation and cognitive ability. Evidence from ideational apraxia suggests that the planning of linguistic and manipulative behaviours is not colocalized in homologous circuits. An alternative account in terms of the evolutionary expansion of the whole prefrontal-premotor area may be more plausible.

Evidence is presented from a polyglot savant to suggest that double dissociations between linguistic and nonverbal abilities are more important than Müller's target article implies. It is also argued that the special nature of syntax makes its assimilation to other aspects of language or to nonhuman communication systems radically implausible.

Müller's review of the neuroscientific evidence undermines nativist claims for autonomous syntax and the argument from the poverty of the stimulus. Generativists will appeal to data from language acquisition, but here too there is growing evidence against the nativist position. Epigenetic naturalism, the developmental alternative to nativism, can be extended to epigenetic socionaturalism, acknowledging the importance of sociocultural processes in language and cognitive development.

Words become associated following repeated co-occurrence episodes. This process might be further determined by the semantic characteristics of the words. The present study focused on how semantic and episodic factors interact in incidental formation of word associations. First, we found that human participants associate semantically related words more easily than unrelated words; this advantage increased linearly with repeated co-occurrence. Second, we developed a computational model, SEMANT, suggesting a possible mechanism for this semantic-episodic interaction. In SEMANT, episodic associations are implemented through (...) lateral connections between nodes in a pre-existent self-organized map of word semantics. These connections are strengthened at each instance of concomitant activation, proportionally with the amount of the overlapping activity waves of activated nodes. In computer simulations SEMANT replicated the dynamics of associative learning in humans and led to testable predictions concerning normal associative learning as well as impaired learning in a diffuse semantic system like that characteristic of schizophrenia. (shrink)

Neuropsychological results are increasingly cited in cognitive theories although their methodology has been severely criticised. The book argues for an eclectic approach but particularly stresses the use of single-case studies. A range of potential artifacts exists when inferences are made from such studies to the organisation of normal function – for example, resource differences among tasks, premorbid individual differences, and reorganisation of function. The use of “strong” and “classical” dissociations minimises potential artifacts. The theoretical convergence between findings from fields where (...) cognitive neuropsychology is well developed and those from the normal literature strongly suggests that the potential artifacts are not critical. The fields examined in detail in this respect are short-term memory, reading, writing, the organisation of input and output speech systems, and visual perception. Functional dissociation data suggest that not only are input systems organised modularly, but so are central systems. This conclusion is supported by findings on impairment of knowledge, visual attention, supervisory functions, memory, and consciousness. (shrink)

Most words in English are ambiguous between different interpretations; words can mean different things in different contexts. We investigate the implications of different types of semantic ambiguity for connectionist models of word recognition. We present a model in which there is competition to activate distributed semantic representations. The model performs well on the task of retrieving the different meanings of ambiguous words, and is able to simulate data reported by Rodd, Gaskell, and Marslen‐Wilson [J. Mem. Lang. 46 (2002) 245] on (...) how semantic ambiguity affects lexical decision performance. In particular, the network shows a disadvantage forwords with multiple unrelated meanings (e.g., bark) that coexists with a benefit for words with multiple related word senses (e.g., twist). The ambiguity disadvantage arises because of interference between the different meanings, while the sense benefit arises because of differences in the structure of the attractor basins formed during learning. Words with few senses develop deep, narrow attractor basins, while words with many senses develop shallow, broad basins. We conclude that the mental representations of word meanings can be modelled as stable states within a high‐dimensional semantic space, and that variations in the meanings of words shape the landscape of this space. (shrink)