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.

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.

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.

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.

Müller argues that general cognitive skills and linguistic skills are not necessarily independent. However, cross-linguistic evidence from an inherited specific language disorder affecting productive rules suggests significant degrees of modularity, innateness, and universality of language. Confident claims about the overall nature of such a complex system still await more interdisciplinary research.

Because Müiller fails to distinguish between two senses of the term “autonomy,” there is a danger that his results will be misinterpreted by both linguists and neuroscientists. Although he may very well have been successful in refuting one sense of autonomy, he may actually have helped to provide an explanation for the correctness of autonomy in its other sense.

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 study of signed languages has inspired scientific' speculation regarding foundations of human language. Relationships between the acquisition of sign language in apes and man are discounted on logical grounds. Evidence from the differential hreakdown of sign language and manual pantomime places limits on the degree of overlap between language and nonlanguage motor systems. Evidence from functional magnetic resonance imaging reveals neural areas of convergence and divergence underlying signed and spoken languages.

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)

Preadaptation for language is an unnecessary assumption because intermediate stages of linguistic ability are possible and adaptive. Language could have evolved through gradual selection from structures exhibiting few features associated with modern structures. Without physical evidence pertaining to language ability in prehabilis hominids, it remains possible that selective pressures for language use preceded and necessitated modern neurolinguistic structures.

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)

Clahsen's claim that output forms of productive processes are never listed in the lexicon is a consequence of the rule/list fallacy, empirically incorrect, and not necessary for the hypothesis that the human language faculty has a dual structure, that is, a lexicon and a set of rules.

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.

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.

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)

Müller attempts to downplay cases of dissociation between language and cognition as evidence against the modularity of language. We review cases of associations between verbal and nonverbal abilities as further evidence against the notion of language as an autonomous subsystem. We also point out a discrepancy between his proposal of homologies between nonhuman primates' communication and human language and recent proposals on the evolution of language.

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.

Wilkins & Wakefield assign importance to motor systems but skip from anatomy to cognitive structure with little attention to behavior. Organisms, no matter how sophisticated, that do not behave in accord with what they know will fall by the evolutionary wayside. Facts about behavior can supplement the authors' theory, whose hierarchical structures can accommodate an evolutionary scenario in which a million years or more of functionally varied utterances mainly limited to single words is followed by an explosion of linguistic diversity (...) with the development in the last 50,000 years or so of syntactically organized multiple word utterances. (shrink)

If POT (parieto-occipital-temporal junction) reorganization came earlier in australopithecines than in Homo, it is likely that the selective pressures were different, and not necessarily directed toward language. The brain endocast evidence for the POT in A. afarensis is actually better than it is for early Homo.

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)

We agree with Müller's epigenetic view of evolution and ontogeny and applaud his multilevel perspective. With him, we stress the importance in ontogeny of progressive specialisation rather than prewired structures. However, we argue that he slips from “speech” to “language” and that, in seeking homologies, these two levels need to be kept separate in the analysis of evolution and ontogeny.

Three issues are addressed in this commentary. (1) Wilkins & Wakefield are commended for placing the complex behavior they discuss within an evolutionary matrix. (2) They err on a number of points in regard to their treatment of this complex behavior. These involve (a) their emphasis on the evolution of conceptual structure rather than language, (b) their equation of meaning with reference, (c) their minimalist view of learning theory, and (d) their separation of the evolution of speech from that of (...) language. (3) They adopt a framework for neural preconditions that has been clearly discredited on both behavioral and neuroanatomical grounds. Finally, recent research involving modern neuroimaging techniques casts further doubt on the authors' postulated functions for the neural areas they discuss. (shrink)

This paper presents a theory of the syntactic aspects of human sentence production. An important characteristic of unprepared speech is that overt pronunciation of a sentence can be initiated before the speaker has completely worked out the meaning content he or she is going to express in that sentence. Apparently, the speaker is able to build up a syntactically coherent utterance out of a series of syntactic fragments each rendering a new part of the meaning content. This incremental, left‐to‐right mode (...) of sentence production is the central capability of the proposed Incremental Procedural Grammar (IPG). Certain other properties of spontaneous speech, as derivable from speech errors, hesitations, self‐repairs, and language pathology, are accounted for as well.The psychological plausibility thus gained by the grammar appears compatible with a satisfactory level of linguistic plausibility in that sentences receive structural descriptions which are in line with current theories of grammar. More importantly, an explanation for the existence of configurational conditions on transformations and other linguistics rules is proposed.The basic design feature of IPG which gives rise to these psychologically and linguistically desirable properties, is the “Procedures + Stack” concept. Sentences are built not by a central constructing agency which overlooks the whole process but by a team of syntactic procedures (modules) which work—in parallel—on small parts of the sentence, have only a limited overview, and whose sole communication channel is a stack.IPG covers object complement constructions, interrogatives, and word order in main and subordinate clauses. It handles unbounded dependencies, crossserial dependencies and coordination phenomena such as gapping and conjunction reduction. It is also capable of generating self‐repairs and elliptical answers to questions. IPG has been implemented as an incremental Dutch sentence generator written in LISP. (shrink)

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.

Segregated neural circuits that effect particular domain-specific behaviors can be differentiated from neuroanatomical structures implicated in many different aspects of behavior. The basal ganglionic components of circuits regulating nonlinguistic motor behavior, speech, and syntax all function in a similar manner. Hence, it is unlikely that special properties and evolutionary mechanisms are associated with the neural bases of human language.

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.

The synthetic position adopted by Müller is weakened by a large number of overinterpretations and misrepresentations, together with a caricatured view of innateness and modularity.

Müller argues that double dissociations do not imply underlying modularity of the cognitive system, citing neural networks as examples of fully distributed systems that can give rise to double dissociations. We challenge this claim, noting that suchdouble dissociations typically do not “scale-up,” and that even some singledissociations can be difficult to account for in a distributed system.

Both autonomy and local specificity are compatible with observed interconnectivity at the cell level when considering two different levels: cell assemblies and brain systems. Early syntactic structuring processes in particular are likely to representan autonomous module in the language/brain 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.

This commentary confronts one of the central tenets advanced in Wilkins & Wakefield's target article: By adopting a very narrow perspective on language, the authors have effectively limited discussion of earlier linguistic capabilities thought to be at least facilitative of, if not prerequisite to language defined as a An alternative conceptualization for describing semiogenesis is offered.

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.

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)

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.