Language and learning: the cognitive revolution at 60-odd.

Around the middle of the last century, the prevailing psychological paradigm of behaviourism was challenged by what is now known as the cognitive revolution. Behaviourists viewed learning as changes in patterns of behaviour through reinforcement. By contrast, advocates of the cognitive approach argued that such behavioural changes were outward manifestations of computational operations on mental representations. Here we consider the current state of the cognitive revolution, focusing on the two most contentious issues in the debate: language and learning. The cognitive approach has proved to be extremely fruitful in both fields. Although contemporary learning theory has almost completely embraced the cognitive approach, the study of language has witnessed a clear empiricist trend to revert back to a kind of neo-behaviourism. Many contemporary authors contend that language is a means of communication that is learned solely through the observation of external events, and culturally transmitted to successive generations. Here, we argue that learning and language can only be properly understood from a cognitive perspective, where the mind is conceived of as a biologically underpinned computational system. As is the case in learning theory, there is abundant evidence showing that language is subserved by an autonomous cognitive system in the mind. We conclude that the cognitive revolution has fundamentally changed our understanding of the mind.

Vocal learning in songbirds: the role of syllable order in song recognition.

Songbird vocal learning has interesting behavioural and neural parallels with speech acquisition in human infants. Zebra finch males sing one unique song that they imitate from conspecific males, and both sexes learn to recognize their father's song. Although males copy the stereotyped syllable sequence of their father's song, the role of sequential information in recognition remains unclear. Here, we investigated father's song recognition after changing the serial order of syllables (switching the middle syllables, first and last syllables, or playing all syllables in inverse order). Behavioural approach and call responses of adult male and female zebra finches to their father's versus unfamiliar songs in playback tests demonstrated significant recognition of father's song with all syllable-order manipulations. We then measured behavioural responses to normal versus inversed-order father's song. In line with our first results, the subjects did not differentiate between the two. Interestingly, when males' strength of song learning was taken into account, we found a significant correlation between song imitation scores and the approach responses to the father's song. These findings suggest that syllable sequence is not essential for recognition of father's song in zebra finches, but that it does affect responsiveness of males in proportion to the strength of vocal learning. This article is part of the theme issue 'Vocal learning in animals and humans'.

Memory-specific correlated neuronal activity in higher-order auditory regions of a parrot.

Male budgerigars (Melopsittacus undulatus) are open-ended learners that can learn to produce new vocalisations as adults. We investigated neuronal activation in male budgerigars using the expression of the protein products of the immediate early genes zenk and c-fos in response to exposure to conspecific contact calls (CCs: that of the mate or an unfamiliar female) in three subregions (CMM, dNCM and vNCM) of the caudomedial pallium, a higher order auditory region. Significant positive correlations of Zenk expression were found between these subregions after exposure to mate CCs. In contrast, exposure to CCs of unfamiliar females produced no such correlations. These results suggest the presence of a CC-specific association among the subregions involved in auditory memory. The caudomedial pallium of the male budgerigar may have functional subdivisions that cooperate in the neuronal representation of auditory memory.

The slings and arrows of comparative linguistics.

In this Formal Comment the authors respond to objections to their previous Essay, reiterating that comparative linguistics is not an easy undertaking.

Meaningful syntactic structure in songbird vocalizations?

The faculty of language is thought to be uniquely human. Recently, it has been claimed that songbirds are able to associate meaning with sound, comparable to the way that humans do. In human language, the meaning of expressions (semantics) is dependent on a mind-internal hierarchical structure (syntax). Meaning is associated with structure through the principle of compositionality, whereby the meaning of a complex expression is a function of the meaning of its constituent parts and the mode of composition. We argue that while recent experimental findings on songbird call sequences offer exciting novel insights into animal communication, despite claims to the contrary, they are quite unlike what we find in human language. There are indeed remarkable behavioral and neural parallels in auditory-vocal imitation learning between songbirds and human infants that are absent in our closest evolutionary relatives, the great apes. But so far, there is no convincing evidence of syntax-determined meaning in nonhuman animals.

The growth of language: Universal Grammar, experience, and principles of computation.

Human infants develop language remarkably rapidly and without overt instruction. We argue that the distinctive ontogenesis of child language arises from the interplay of three factors: domain-specific principles of language (Universal Grammar), external experience, and properties of non-linguistic domains of cognition including general learning mechanisms and principles of efficient computation. We review developmental evidence that children make use of hierarchically composed structures ('Merge') from the earliest stages and at all levels of linguistic organization. At the same time, longitudinal trajectories of development show sensitivity to the quantity of specific patterns in the input, which suggests the use of probabilistic processes as well as inductive learning mechanisms that are suitable for the psychological constraints on language acquisition. By considering the place of language in human biology and evolution, we propose an approach that integrates principles from Universal Grammar and constraints from other domains of cognition. We outline some initial results of this approach as well as challenges for future research.

Brains for birds and babies: Neural parallels between birdsong and speech acquisition.

Language as a computational cognitive mechanism appears to be unique to the human species. However, there are remarkable behavioral similarities between song learning in songbirds and speech acquisition in human infants that are absent in non-human primates. Here we review important neural parallels between birdsong and speech. In both cases there are separate but continually interacting neural networks that underlie vocal production, sensorimotor learning, and auditory perception and memory. As in the case of human speech, neural activity related to birdsong learning is lateralized, and mirror neurons linking perception and performance may contribute to sensorimotor learning. In songbirds that are learning their songs, there is continual interaction between secondary auditory regions and sensorimotor regions, similar to the interaction between Wernicke's and Broca's areas in human infants acquiring speech and language. Taken together, song learning in birds and speech acquisition in humans may provide useful insights into the evolution and mechanisms of auditory-vocal learning.

Language, mind and brain.

Language serves as a cornerstone of human cognition. However, our knowledge about its neural basis is still a matter of debate, partly because 'language' is often ill-defined. Rather than equating language with 'speech' or 'communication', we propose that language is best described as a biologically determined computational cognitive mechanism that yields an unbounded array of hierarchically structured expressions. The results of recent brain imaging studies are consistent with this view of language as an autonomous cognitive mechanism, leading to a view of its neural organization, whereby language involves dynamic interactions of syntactic and semantic aspects represented in neural networks that connect the inferior frontal and superior temporal cortices functionally and structurally.

What do animals learn in artificial grammar studies?

Artificial grammar learning is a popular paradigm to study syntactic ability in nonhuman animals. Subjects are first trained to recognize strings of tokens that are sequenced according to grammatical rules. Next, to test if recognition depends on grammaticality, subjects are presented with grammar-consistent and grammar-violating test strings, which they should discriminate between. However, simpler cues may underlie discrimination if they are available. Here, we review stimulus design in a sample of studies that use particular sounds as tokens, and that claim or suggest their results demonstrate a form of sequence rule learning. To assess the extent of acoustic similarity between training and test strings, we use four simple measures corresponding to cues that are likely salient. All stimulus sets contain biases in similarity measures such that grammatical test stimuli resemble training stimuli acoustically more than do non-grammatical test stimuli. These biases may contribute to response behaviour, reducing the strength of grammatical explanations. We conclude that acoustic confounds are a blind spot in artificial grammar learning studies in nonhuman animals.

Sex differences in behavioural and neural responsiveness to mate calls in a parrot.

Vocalisation in songbirds and parrots has become a prominent model system for speech and language in humans. We investigated possible sex differences in behavioural and neural responsiveness to mate calls in the budgerigar, a vocally-learning parrot. Males and females were paired for 5 weeks and then separated, after which we measured vocal responsiveness to playback calls (a call of their mate versus a call of an unfamiliar conspecific). Both sexes learned to recognise mate calls during the pairing period. In males, but not females, mate calls evoked significantly fewer vocal responses than unfamiliar calls at one month after separation. Furthermore, in females, there was significantly greater molecular neuronal activation in response to mate calls compared to silence in the caudomedial mesopallium (CMM), a higher-order auditory region, in both brain hemispheres. In males, we found right-sided dominance of molecular neuronal activation in response to mate calls in the CMM. This is the first evidence suggesting sex differences in functional asymmetry of brain regions related to recognition of learned vocalisation in birds. Thus, sex differences related to recognition of learned vocalisations may be found at the behavioural and neural levels in avian vocal learners as it is in humans.

Structures, Not Strings: Linguistics as Part of the Cognitive Sciences.

There are many questions one can ask about human language: its distinctive properties, neural representation, characteristic uses including use in communicative contexts, variation, growth in the individual, and origin. Every such inquiry is guided by some concept of what 'language' is. Sharpening the core question--what is language?--and paying close attention to the basic property of the language faculty and its biological foundations makes it clear how linguistics is firmly positioned within the cognitive sciences. Here we will show how recent developments in generative grammar, taking language as a computational cognitive mechanism seriously, allow us to address issues left unexplained in the increasingly popular surface-oriented approaches to language.

Evolution cannot explain how minds work.

Following Jerry Hogan, I argue that questions of function and evolution, and questions of mechanism should be seen as logically distinct. Evolution is concerned with a historical reconstruction of traits, while the actual underlying mechanisms are the domain of cognitive neuroscience and psychology. Functional and evolutionary considerations may be used to generate hypotheses regarding the underlying mechanisms. But these hypotheses may be false and should always be tested empirically. Many researchers still hold that common descent implies cognitive closeness. Studies on birds suggest that evolutionary convergence may be the rule rather than the exception in animal cognition. Neurocognitive differences between classes of individuals are often thought to be the result of adaptive specialisation. In the case of learning and memory, however, empirical results are more consistent with a 'general process' interpretation, without qualitative differences between different taxa. Evolutionary psychology (EP) argues that the mind of modern humans was formed as a result of selection pressures in the Stone Age. The empirical data are often overinterpreted, and EP is mostly based upon an outdated view of evolutionary biology. In human speech and language, both neurogenetic homology and evolutionary convergence are involved regarding speech, but human language has a unique combinatorial complexity. This article is part of a Special Issue entitled: In Honor of Jerry Hogan.

Learning-related brain hemispheric dominance in sleeping songbirds.

There are striking behavioural and neural parallels between the acquisition of speech in humans and song learning in songbirds. In humans, language-related brain activation is mostly lateralised to the left hemisphere. During language acquisition in humans, brain hemispheric lateralisation develops as language proficiency increases. Sleep is important for the formation of long-term memory, in humans as well as in other animals, including songbirds. Here, we measured neuronal activation (as the expression pattern of the immediate early gene ZENK) during sleep in juvenile zebra finch males that were still learning their songs from a tutor. We found that during sleep, there was learning-dependent lateralisation of spontaneous neuronal activation in the caudomedial nidopallium (NCM), a secondary auditory brain region that is involved in tutor song memory, while there was right hemisphere dominance of neuronal activation in HVC (used as a proper name), a premotor nucleus that is involved in song production and sensorimotor learning. Specifically, in the NCM, birds that imitated their tutors well were left dominant, while poor imitators were right dominant, similar to language-proficiency related lateralisation in humans. Given the avian-human parallels, lateralised neural activation during sleep may also be important for speech and language acquisition in human infants.