The cerebellum: Its role in language and related cognitive and affective functions

The traditional view on the cerebellum as the sole coordinator of motor function has been substantially redefined during the past decades. Neuroanatomical, neuroimaging and clinical studies have extended the role of the cerebellum to the modulation of cognitive and affective processing. Neuroanatomical studies have demonstrated cerebellar connectivity with the supratentorial association areas involved in higher cognitive and affective functioning, while functional neuroimaging and clinical studies have provided evidence of cerebellar involvement in a variety of cognitive and affective tasks. This paper reviews the recently acknowledged role of the cerebellum in linguistic and related cognitive and behavioral–affective functions. In addition, typical cerebellar syndromes such as the cerebellar cognitive affective syndrome (CCAS) and the posterior fossa syndrome (PFS) will be briefly discussed and the current hypotheses dealing with the presumed neurobiological mechanisms underlying the linguistic, cognitive and affective modulatory role of the cerebellum will be reviewed.     

Neuroimaging studies of the cerebellum: language, learning and memory

During the decade following a functional neuroimaging study of language that showed cerebellar involvement in a cognitive task, PET and fMRI studies have continued to provide evidence that the role of the cerebellum extends beyond that of motor control and that this structure contributes in some way to cognitive operations. In this review, we describe neuroimaging evidence for cerebellar involvement in working memory, implicit and explicit learning and memory, and language, and we discuss some of the problems and limitations faced by researchers who use neuroimaging to investigate cerebellar function. We also raise a set of outstanding questions that need to be addressed through further neuroimaging and behavioral experiments before differing functional accounts of cerebellar involvement in cognition can be resolved.     

Solving the Mystery of the Human Cerebellum

The mystery of the human cerebellum is this: Why did it enlarge so dramatically in the last million years of human evolution, concomitantly with the greater enlargement of the cerebral cortex? A solution to this mystery was proposed in the 20th century as a result of research by several groups of scientists who investigated the contributions of the cerebellum to the cerebral cortex. In contrast to the 19th century investigations, which were focused on the motor functions of the cerebellum, the focus of the subsequent investigations was expanded to include some mental functions because evidence was produced that the cerebellum contributes to cognition. It was proposed that the combination in the cerebellum of motor and mental capabilities enables the cerebellum to confer on humans some adaptive advantages of great value, and this ability would explain why the human cerebellum has continued to enlarge so dramatically. A valuable adaptive advantage that is included in the proposal is the possibility that the cerebellum couples the motor function of articulating speech to the mental function that selects the language to be spoken, thus helping to produce fluent human speech and language. The validity of this proposal about linguistic processing has not yet been verified. Therefore the mystery of cerebellar enlargement in humans is not yet solved and requires further research.     

The Role of the Cerebellum in Cognition and Emotion: Personal Reflections Since 1982 on the Dysmetria of Thought Hypothesis,and Its Historical Evolution from Theory to Therapy

The cognitive neuroscience of the cerebellum is now an established multidisciplinary field of investigation. This essay traces the historical evolution of this line of inquiry from an emerging field to its current status, with personal reflections over almost three decades on this journey of discovery. It pays tribute to early investigators who recognized the wider role of the cerebellum beyond motor control, traces the origins of new terms and concepts including the dysmetria of thought theory, the universal cerebellar transform, and the cerebellar cognitive affective syndrome, and places these developments within the broader context of the scientific efforts of a growing community of cerebellar cognitive neuroscientists. This account considers the converging evidence from theoretical, anatomical, physiological, clinical, and functional neuroimaging approaches that have resulted in the transition from recognizing the cerebellar incorporation into the distributed neural circuits subserving cognition and emotion, to a hopeful new era of treatment of neurocognitive and neuropsychiatric manifestations of cerebellar diseases, and to cerebellar-based interventions for psychiatric disorders.     

Long-term associative learning predicts verbal short-term memory performance

Studies using tests such as digit span and nonword repetition have implicated short-term memory across a range of developmental domains. Such tests ostensibly assess specialized processes for the short-term manipulation and maintenance of information that are often argued to enable long-term learning. However, there is considerable evidence for an influence of long-term linguistic learning on performance in short-term memory tasks that brings into question the role of a specialized short-term memory system separate from long-term knowledge. Using natural language corpora, we show experimentally and computationally that performance on three widely used measures of short-term memory (digit span, nonword repetition, and sentence recall) can be predicted from simple associative learning operating on the linguistic environment to which a typical child may have been exposed. The findings support the broad view that short-term verbal memory performance reflects the application of long-term language knowledge to the experimental setting.     

The cerebellum and the timing of coordinated eye and hand tracking.

The cerebellum is known to have important functions in motor control, coordination, motor learning, and timing. It may have other "higher" functions as well, up to and including cognitive processing independent of motor behavior. In this article, we will review some of the evidence from functional imaging, lesion studies, electrophysiological recordings, and anatomy which support the theory that the cerebellum provides a "forward model" of the motor system. This forward model would be used for control of movement; it could also underlie a cerebellar role in coordination. In this role, the forward model would generate time-specific signals predicting the motion of each motor effector, essential for predictive control of, for example, eye and hand movements. Data are presented from human eye and hand tracking that support this. Tracking performance is better if eye and hand follow the same spatial trajectory, but better still if the eye leads the hand by about 75 to 100 ms. This suggests that information from the ocular control system feeds into the manual control system to assist its tracking.     

Architecture and organizational principles of Broca's region

Identifying cortical areas for language and speech processing is a prerequisite for cognitive neuroscience and clinical research. Although Broca's region is one of the essential nodes in the language network, its anatomical constituents are ill-defined and multiple definitions of Broca's region exist. Sanides' concept of microstructural gradations interpreted Broca's region as developing from neighboring motor, dorsolateral-prefrontal, and insular cortices. Recent mapping approaches based on cytoarchitecture, transmitter receptor distribution, and connectivity revealed a highly differentiated segregation of this region far beyond Brodmann's classical scheme. This novel segregational concept of structural and functional architecture more adequately reflects the various functions of Broca's region in cognitive and/or linguistic processes.     

Ontogenesis and Phylogenesis in the Analysis of Chinese Classifiers： Remarks on Philosophical Method

Philosophical analysis is commonly assumed to involve decomposing the meaning of a sentence or an expression into a set of conceptually basic constituent parts. This essay challenges this traditional view by examining the potential semantic roles that classifier phrases play in Chinese. It is suggested that the conceptual resources necessary for justifying claims about the semantical status of natural language classifier phrases should be informed in part by methods that accommodate ontogenic and evolutionary contexts. Evidence is provided for the view that many Chinese classifiers （but not all） are features regimented in the grammar of Chinese that have no functional role in normal adult communication, but which play an ontogenetic role in the child＇s development of linguistic competency. Furthermore, it is suggested that this ontogenetic role has features in common with the phylogenetic processes by which Chinese or its classical variants came about, as a later product of mechanisms that evolved in the species in accordance with varying demands for successful communication.     

Children with Specific Language Impairment are not impaired in the acquisition and retention of Pavlovian delay and trace conditioning of the eyeblink response

Three converging lines of evidence have suggested that cerebellar abnormality is implicated in developmental language and literacy problems. First, some brain imaging studies have linked abnormalities in cerebellar grey matter to dyslexia and specific language impairment (SLI). Second, theoretical accounts of both dyslexia and SLI have postulated impairments of procedural learning and automatisation of skills, functions that are known to be mediated by the cerebellum. Third, motor learning has been shown to be abnormal in some studies of both disorders. We assessed the integrity of face related regions of the cerebellum using Pavlovian eyeblink conditioning in 7–11 year-old children with SLI. We found no relationship between oral language skills or literacy skills with either delay or trace conditioning in the children. We conclude that this elementary form of associative learning is intact in children with impaired language or literacy development.     

Thought beyond language: neural dissociation of algebra and natural language

A central question in cognitive science is whether natural language provides combinatorial operations that are essential to diverse domains of thought. In the study reported here, we addressed this issue by examining the role of linguistic mechanisms in forging the hierarchical structures of algebra. In a 3-T functional MRI experiment, we showed that processing of the syntax-like operations of algebra does not rely on the neural mechanisms of natural language. Our findings indicate that processing the syntax of language elicits the known substrate of linguistic competence, whereas algebraic operations recruit bilateral parietal brain regions previously implicated in the representation of magnitude. This double dissociation argues against the view that language provides the structure of thought across all cognitive domains.     

Lexical entries and rules of language: a multidisciplinary study of German inflection

Following much work in linguistic theory, it is hypothesized that the language faculty has a modular structure and consists of two basic components, a lexicon of (structured) entries and a computational system of combinatorial operations to form larger linguistic expressions from lexical entries. This target article provides evidence for the dual nature of the language faculty by describing recent results of a multidisciplinary investigation of German inflection. We have examined: (1) its linguistic representation, focussing on noun plurals and verb inflection (participles), (2) processes involved in the way adults produce and comprehend inflected words, (3) brain potentials generated during the processing of inflected words, and (4) the way children acquire and use inflection. It will be shown that the evidence from all these sources converges and supports the distinction between lexical entries and combinatorial operations. Our experimental results indicate that adults have access to two distinct processing routes, one accessing (irregularly) inflected entries from the mental lexicon and another involving morphological decomposition of (regularly) inflected words into stem + affix representations. These two processing routes correspond to the dual structure of the linguistic system. Results from event-related potentials confirm this linguistic distinction at the level of brain structures. In children's language, we have also found these two processes to be clearly dissociated; regular and irregular inflection are used under different circumstances, and the constraints under which children apply them are identical to those of the adult linguistic system. Our findings will be explained in terms of a linguistic model that maintains the distinction between the lexicon and the computational system but replaces the traditional view of the lexicon as a simple list of idiosyncrasies with the notion of internally structured lexical representations.     

Understanding the meaning of words and sentences: The role of non-linguistic processes

This special issue on linguistic and non-linguistic processes in the comprehending of words and sentences represents the outcome of a three-day workshop at the University of Tübingen, Germany. The workshop focused on the contribution of non-linguistic processes in language comprehension and hence brought together researchers from the domains of linguistic and non-linguistic cognition.     

Cerebellar output: motor and cognitive channels

The input to the cerebellum has long been known to originate from widespread regions of the cerebral cortex including the frontal, parietal and temporal lobes. The output of the cerebellum, however, was thought to project mainly to the primary motor cortex. Recent anatomical observations have challenged this view. It is now apparent that cerebellar output goes to multiple cortical areas, including not only the primary motor cortex, but also areas of premotor and prefrontal cortex. In fact, there is growing evidence that each of the areas of cerebral cortex that project to the cerebellum is also the target of cerebellar output. The cerebellar output to individual cortical areas originates from distinct clusters of neurons in the deep nuclei which we have termed `output channels'. The individual output channels to the cortical areas we have examined display little or no overlap. Physiological recordings in awake trained primates indicate that neurons in different output channels appear to be involved in distinct aspects of behavior, and in both motor and cognitive functions. These observations indicate that the cerebellar influence on the cerebral cortex is more extensive than previously recognized.     

Theories of early language acquisition

What features of brain processing and neural development support linguistic development in young children? To what extent is the profile and timing of linguistic development in young children determined by a pre-ordained genetic programme? Does the environment play a crucial role in determining the patterns of change observed in children growing up? Recent experimental, neuroimaging and computational studies of developmental change in children promise to contribute to a deeper understanding of how the brain becomes wired up for language. In this review, the muttidisciplinary perspectives of cognitive neuroscience, experimental psycholinguistics and neural network modelling are brought to bear on four distinct areas in the study of language acquisition: early speech perception, word recognition, word learning and the acquisition of grammatical inflections. It is suggested that each area demonstrates how linguistic development can be driven by the interaction of general learning mechanisms, highly sensitive to particular statistical regularities in the input, with a richly structured environment which provides the necessary ingredients for the emergence of linguistic representations that support mature language processing. Similar epigenetic principles, guiding the emergence of linguistic structure, apply to all these domains, offering insights into phenomena ranging from the precocity of young infant's sensitivity to speech contrasts to the complexities of the problem facing the young child learning the arabic plural.     

Conventions and Their Role in Language

Two of the most fundamental questions about language are these: what are languages?; and, what is it to know a given language? Many philosophers who have reflected on these questions have presented answers that attribute a central role to conventions. In one of its boldest forms such a view runs as follows. Languages are either social entities constituted by networks of social conventions or abstract objects where when a particular community speaks a given language they do so in virtue of the conventions operative within that community. Consequently, for an individual to know a given language is for them to be party to the relevant conventions. Call this view conventionalism. In this article my aim is to evaluate conventionalism. I will argue that although there are linguistic conventions and that they do play an important role in language development and communication conventionalism should be rejected in favour of a more psychologistically orientated position.     

The neurology of syntax: language use without Broca's area

A new view of the functional role of the left anterior cortex in language use is proposed. The experimental record indicates that most human linguistic abilities are not localized in this region. In particular, most of syntax (long thought to be there) is not located in Broca's area and its vicinity (operculum, insula, and subjacent white matter). This cerebral region, implicated in Broca's aphasia, does have a role in syntactic processing, but a highly specific one: It is the neural home to receptive mechanisms involved in the computation of the relation between transformationally moved phrasal constituents and their extraction sites (in line with the Trace-Deletion Hypothesis). It is also involved in the construction of higher parts of the syntactic tree in speech production. By contrast, basic combinatorial capacities necessary for language processing--for example, structure-building operations, lexical insertion--are not supported by the neural tissue of this cerebral region, nor is lexical or combinatorial semantics. The dense body of empirical evidence supporting this restrictive view comes mainly from several angles on lesion studies of syntax in agrammatic Broca's aphasia. Five empirical arguments are presented: experiments in sentence comprehension, cross-linguistic considerations (where aphasia findings from several language types are pooled and scrutinized comparatively), grammaticality and plausibility judgments, real-time processing of complex sentences, and rehabilitation. Also discussed are recent results from functional neuroimaging and from structured observations on speech production of Broca's aphasics. Syntactic abilities are nonetheless distinct from other cognitive skills and are represented entirely and exclusively in the left cerebral hemisphere. Although more widespread in the left hemisphere than previously thought, they are clearly distinct from other human combinatorial and intellectual abilities. The neurological record (based on functional imaging, split-brain and right-hemisphere-damaged patients, as well as patients suffering from a breakdown of mathematical skills) indicates that language is a distinct, modularly organized neurological entity. Combinatorial aspects of the language faculty reside in the human left cerebral hemisphere, but only the transformational component (or algorithms that implement it in use) is located in and around Broca's area.     

Language and connectionism: the developing interface

After a difficult initial period in which connectionism was perceived as either irrelevant or antithetical to linguistic theory, connectionist concepts are now beginning to be brought to bear on basic issues concerning the structure, acquisition, and processing of language, both normal and disordered. This article describes some potential points of further contact between connectionism and linguistic theory. I consider how connectionist concepts may be relevant to issues concerning the representation of linguistic knowledge; the role of a priori constraints on acquisition; and the poverty of the stimulus argument. I then discuss whether these models contribute to the development of explanatory theories of language.     

Individual differences in the acquisition of non‐linguistic audio‐visual associations in 5 year olds

Audio‐visual associative learning – at least when linguistic stimuli are employed – is known to rely on core linguistic skills such as phonological awareness. Here we ask whether this would also be the case in a task that does not manipulate linguistic information. Another question of interest is whether executive skills, often found to support learning, may play a larger role in a non‐linguistic audio‐visual associative task compared to a linguistic one. We present a new task that measures learning when having to associate non‐linguistic auditory signals with novel visual shapes. Importantly, our novel task shares with linguistic processes such as reading acquisition the need to associate sounds with arbitrary shapes. Yet, rather than phonemes or syllables, it uses novel environmental sounds – therefore limiting direct reliance on linguistic abilities. Five‐year‐old French‐speaking children (N = 76, 39 girls) were assessed individually in our novel audio‐visual associative task, as well as in a number of other cognitive tasks evaluating linguistic abilities and executive functions. We found phonological awareness and language comprehension to be related to scores in the audio‐visual associative task, while no correlation with executive functions was observed. These results underscore a key relation between foundational language competencies and audio‐visual associative learning, even in the absence of linguistic input in the associative task.