language

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in Neurobiology

Introduction A critical analysis of recent research in language development requires consideration of two hotly contested issues. Proponents of innateness argue that our ability to acquire a language is determined by genetic factors, and mediated by a form of neural organization that is unique to our species. Proponents of domain specificity argue that this ability is also separate and ‘special’, constituting what Chomsky [l] has termed a ‘mental organ’. Because the first claim has to be true at some level of analysis (i.e. we are the only species that can acquire a language in its full-fledged form; see [a*]), the real debate revolves around the mental organ claim [3,4-l. h-e the mental structures that support language ‘modular’, discontinuous and dissociable from all other perceptual and cognitive systems? Does the brain of a newborn child contain neural structures that are destined to mediate language, and language alone? The domain-specificity view can be contrasted with an approach in which language is viewed as an innate system, but one that involves a reconfiguration of mental and neural systems that exist in other species [5*-7-l, and which continue to serve at least some non-linguistic functions in our own [W] In this review I will consider several areas of research on the biological foundations of language learning, focusing on the case for and against domain specificity.

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Innateness

La Jolla, California,

plastic mix of neural systems

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Bates

specificity

Two recent examples illustrate the confusion between innateness and domain specificity. Petitto and Marentette [9**] published an influential paper demonstrating that deaf infants exposed to sign language ‘babble’ with their hands, producing meaningless but systematic actions that are not observed in hearing children. Furthermore, this form of manual babbling occur:, around %lO months

1992, 2:180-185

of age, the point at which vocal babbling appears in the hearing child. The authors conclude that language learning involves innate abilities that are independent of modality (i.e. vocal or manual); they also claim that these abilities are specific to language, providing support for Chomsky’s mental organ claim [ 11. Their first conclusion is clearly supported by the evidence, but the second is not. We have known for more than 100 years that children begin to imitate novel actions (i.e. actions that are not already in their repertoire) at around E&10 months. The more systematic the adult input, the more systematic the child’s imitation is likely to be. Petitto and Marentette’s demonstration of babbling in the visual modality constitutes a particularly beautiful example of this interesting but well established fact. The kind of imitation that underlies babbling is undoubtedly based upon abilities that are innate, and particularly well developed in our species (human children imitate far better and more often than any other primate) [2*,10*], but proof of its existence does not, in itself, constitute evidence in favor of the notion that language is ‘special’. A somewhat different example appeared in a letter to Nuture by Gopnik [ 11-1 (for further details see [12*] ), describing preliminary results from a study of grammatical abilities in a family of individuals suffering from some kind of genetically based disorder (see also [ 13.1). Members of this family had difficulty with particular aspects of grammar, including regular verb inflections (e.g. the ‘-ed’ in verbs like ‘walked’ and ‘kissed’). By contrast, they had less trouble with irregular forms like ‘came’ or ‘gave’. This pattern was offered as an example of an innate and domain-specific disorder, termed ‘feature-blind dysphasia’, and has been cited as evidence in favor of Pinker’s claim that regular and irregular forms are handled by separate mental and perhaps neural mechanisms [ 14**,15*]. Shortly after Gopnik’s letter appeared, Nature published a rebuttal by Vargha-Khadem and Passingham [ 16**] (see also [17-l), who had studied the same family for a num-

Abbreviations MRI-magnetic

180

resonance

imaging; SLlLspecific @

Current

Biology

language impairment;

Ltd ISSN 0959-4388

WMGWilliams

syndrome.

language

ber of years. These authors point out that the members of this family suffer from a much broader range of linguistic deficits than one might conclude from Gopnik’s description. Their peculiar grammatical symptoms are only the tip of an iceberg, one by-product of a disorder with repercussions in many different areas of language and cognition, providing further evidence for innateness, but none for domain-specificity [ 1W]

language

learning

in neural

networks

There is a branch of language acquisition research called ‘learnability theory’ [19*], which uses formal analysis to determine the range of conditions under which different kinds of grammars can (in principle) be learned. Until recently, most of this research had been based upon the assumption that language learning in humans is similar to language learning in serial digital computers, where apriori hypotheses about grammatical rules are tested against strings of input symbols, based on some combination of positive evidence (‘here is a sentence in the target language’), and negative evidence (‘here is a sentence that is not permitted by the target language’). A famous proof by Gold [20] showed that a broad class of grammars (including generative grammars of the sort described by Chomsky) could not be learned by a system of this kind unless negative evidence was available in abundance, or strong innate constraints were placed upon the kinds of hypotheses that the system would consider. As we know that human children are rarely given explicit negative evidence, the learnability theory seems to require the conclusion that children have an extensive store of innate and domain-specific grammatical knowledge. In the past two years, this conclusion has been challenged by major breakthroughs in the application of a different kind of computer architecture (called neural networks, connectionism, and/or parallel distributed processing) to classic problems in language learnability. Because connectionism makes a very different set of assumptions about the way that knowledge is represented and acquired, Gold’s pessimistic conclusions about lan guage learnability do not necessarily apply. This new era began in 1986 with a simulation by Rumelhart and Mc Clelland [21] on the acquisition of the English past tense, showing that connection&t networks go through stages that are very similar to the ones displayed by children who are acquiring English (producing and then recovering from rule-like overgeneralizations like ‘corned’ and ‘wented’, in the absence of negative evidence). This simulation has-been severely criticized (see especially Pinker and Prince [22]). Since then, a number of new works have appeared that get around these criticisms, replieating and extending the Rumelhart-McClelland findings in several new directions [ 18**,23*“-270*]. The most rem cent example comes from Marchman [18**], who has ‘lesioned’ neural networks at various points during learning of the past tense (randomly eliminating between 2 %-44 % of the connections in the network). These simulations capture some classic ‘critical period effects in language learning (e.g. smaller earlier lesions lead to bet-

development

Bates

ter outcomes; later larger lesions lead to persistent problems in grammar), showing that such effects can occur in the absence of ‘special’ maturational constraints (compare with Newport [28*] and Elman [ 24**]). In addition, Marchman’s damaged systems found it more difficult to acquire regular verbs (e.g. ‘walked’) than irregulars (e.g. ‘came’), proving that the specific pattern of deficits described by Gopnik and by Pinker can result from nonspecific forms of brain damage in a general-purpose learning device. Such research on language learning in neural networks is still in its infancy, and we do not know how far it can go. But it promises to be an important tool, helping us to determine just how much innate knowledge has to be in place for certain kinds of learning to occur. / Electrophysiological

studies

.’ , ,L -.T; L,,_’

Z Two laboratories have begun to@produce exciting evidence on the event-related brain potentials associated with stages of language development in the normal child. Molfese [ .Z$P] has uncovered some electrophysiological correlates of the words that l-year-old children do and do not know. Mills, Coffey and Neville [30=*,31*=] have taken the method a step further, demonstrating changes in event-related potentials to familiar and unfamiliar words at different stages in early language development, controlling for chronological age. For example, there are components in the event-related potentials that distinguish lo-month-olds, who have begun to understand words, from IO-month-olds who are still unable to comprehend speech. Additional components in event-related potentials associated with comprehension of familiar words can distinguish between 13.montholds who can and cannot produce those words. In short, the neural systems that mediate language appear to change over time, and as a function of language learning in addition to chronological age.

Children

with focal brain

injury

It has been known for some time that children show better recovery from focal brain injury than adults with analogous lesions [32,33,34-l. As reviewed by Satz et al. [35-l, however, the pendulum has swung back and forth between claims of equipotentiality (i.e. the left hemisphere is in no way specialized for language) and specificity (i.e. left-hemisphere damage always leaves residual damage). New retrospective studies of older children with this etiology suggest that many do show complete recovery in language [36*]; others show relatively subtle deficits, which may be due to ancillary factors [37*]. By contrast, prospective studies of brain-injured infants in the first stages of language learning show significant delays with both left- and right-hemisphere damage [38=,39**,40*], but the pattern of deficits observed in the early stages does not map in any obvious way onto the brain-behavior correlations observed in brain-dam aged adults. For example, comprehension deficits occur

181

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Cognitive

neuroscience

more often with right-hemisphere injury (and not with damage to Wernicke’s area); expressive deficits are more severe and/or more persistent with left posterior injury (and not with damage to Broca’s area). There is also no straightforward effect of lesion size. Indeed, there is some reason to believe that very large lesions lead to an early ‘compensatory switch’ to the undamaged hemisphere, and hence to better short-term outcomes (see also [31**]). My colleagues and I conclude that there is enormous plasticity in the brain regions that can support language, but that recovery occurs after an initial delay, and the areas responsible for language learning are not necessarily the same regions that mediate use and maintenance of language in adults.

Specific language impairment By definition, specific language impairment (SLI) refers to delays in receptive and/or expressive language development in children with no other known form of neurological or cognitive impairment. Recent studies of XI suggest that this definition may not be accurate. Although these children do not suffer from global forms of mental retardation, they show subtle impairments in aspects of cogmtion and/or perception that are not specific to language. For example, new findings [41-l corroborate claims by Tallal and her colleagues [42] that many children with XI experience difficulty in processing rapid transitions in acoustic information (including non-linguistic stimuli). This may help to explain new studies comparing SLI in English, Italian and Hebrew [43”,44*], showing that the specific areas of grammar that are most delayed vary from one language to another, and that the most vulnerable elements within each language appear to be those that are low in ‘phonological substance’ (i.e. salience). The subtle deficits associated with SLI may also transcend the acoustic modality, affecting certain kinds of manual gesture [45*]. The brain bases of this syndrome are still unknown; magnetic resonance imaging (MRI) studies of SLI show no signs of frank lesions, although subtle differences in symmetry have been uncovered [46~*]. Taken together, these studies suggest that SLI may not be a purely linguistic (or acoustic) phenomenon. Nor is it a congenital variant of the aphasias observed in adults with focal brain injury.

Williams

syndrome

and other ‘savants’

The strongest evidence to date in favor of domain specificity comes from rare cases in which language appears to be remarkably spared despite severe limitations in other cognitive domains. Etiologies associated with this unusual profile include spina bifida and hydrocephalus, and a rare form of mental retardation called Williams syndrome (WMS) [47**], The dissociations observed in WMS prove that language can ‘decouple’ from mental age at some point in development. Nevertheless, recent studies of WMS place constraints on the conclusion that

language is a separate mental system from the beginning. First, it is clear that language development is seriously delayed in infants and pre-school children with WMS, suggesting that certain cognitive prerequisites must be in place before language can be acquired [48]. Second, studies of older children with WMS demonstrate peculiar islands of sparing in some non-linguistic domains (e.g. face recognition), suggesting that their spared language may be based on some deviant form of information processing that operates beyond the boundaries of ‘language proper’. Comparisons of WMS and Down syndrome children matched for mental age suggest that the term ‘mental retardation’ is misleading. Each group shows a distinct and complementary profile of cognitive impairments, reminiscent in many respects of the contrasting profiles shown by adult patients with left- versus right-hemisphere dam age. In spite of this, MRI studies of these two groups yield no evidence of a left-right difference [49*]. Instead, there are group differences in brain morphology along an anterior-posterior axis, including differences in cerebellarqerebral ratios, and differences in the proportional size of neo-cerebellar versus paleo-cerebellar structures These landmark studies challenge traditional ideas about the neural structures that mediate language and related cognitive systems.

Conclusion The research of recent years has led to several interesting conclusions. First, neural network simulations of language learning have demonstrated that a great deal of language can be acquired by a general-purpose learning device. Second, electrophysiological studies have shown that the neural systems subserving language change with development, reflecting language level rather than chronological age. Third, studies of language acquisition in children with focal brain injury demonstrate a great deal of plasticity in the brain regions that can subserve language learning. Fourth, studies of specific language delay suggest that SLI is not so specific after all. Fifth, comparative studies of mental retardation challenge any simple view of the relation between language and cogmtion, and raise new questions about the neural substrate of linguistic and cognitive profiles. A great deal has been learnt in the past few years about the biological foundations of language development; evidence for innateness is good, but evidence for a domain-specific ‘mental organ’ is difficult to find. Instead, language learning appears to be based on a relatively plastic mix of neural systems that also serve other functions,

Acknowledgements This review was visiting scientist Research (CNR), riod came from

prepared while the author was in residence as a at the Institute of Psychology, National Council of Rome, Italy. Partial supportfor this sabbatical pethe CNR and from an NIDCD grant ‘Cross-linguistic

language development Bates studies of aphasia’. I am grateful earlier draft of the manuscript.

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36. .

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41. .

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43. ..

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ROMA, LEONARD L: Interpreting

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46. ..

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Bates

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47. ..

48.

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Brain Morphology on Magnetic Resonance Images in Williams Syndrome and Down Syndrome. Arch New01 1990, 47:523_553. Although the contrasting behavioral protiles displayed by children with Williams versus Down syndrome resemble the contrasts associated with right versus left hemisphere damage, no left-right differences between these groups were uncovered on MRL Group cliffe~:ryces were obselved along an anterior-posterior dimension, inclttiding p?;portional differences in the structure of the cerebellum. II,, *;/ ., 49. ..

JERNGANT, BELISIGI U: Anomalous

JERNICANT, HESSEUNK MD, SOWELL E, TALLU P: Cerebral

Structure on Magnetic Resonance Imaging in Language and Learning-Impaired Children. Arch New-01 1991, 48:53!+545. Children diagnosed with specific language impairment at age 4 fail to display normal left-right aswntmetries in MRI after age 10.

E Bates, Departments of Psychology and Cognitive Science, University of California, San Diego, La Jolla, California 92093, USA.

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Language development.

Recent research suggests that our ability to learn language is innate, but not necessarily domain-specific. That is, language development appears to b...
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