Syntactic Complexity and Frequency in the Neurocognitive Language System

Prominent neurobiological models of language follow the widely accepted assumption that language comprehension requires two principal mechanisms: a lexicon storing the sound-to-meaning mapping of words, primarily involving bilateral temporal regions, and a combinatorial processor for syntactically structured items, such as phrases and sentences, localized in a left-lateralized network linking left inferior frontal gyrus (LIFG) and posterior temporal areas. However, recent research showing that the processing of simple phrasal sequences may engage only bilateral temporal areas, together with the claims of distributional approaches to grammar, raises the question of whether frequent phrases are stored alongside individual words in temporal areas. In this fMRI study, we varied the frequency of words and of short and long phrases in English. If frequent phrases are indeed stored, then only less frequent items should generate selective left frontotemporal activation, because memory traces for such items would be weaker or not available in temporal cortex. Complementary univariate and multivariate analyses revealed that, overall, simple words (verbs) and long phrases engaged LIFG and temporal areas, whereas short phrases engaged bilateral temporal areas, suggesting that syntactic complexity is a key factor for LIFG activation. Although we found a robust frequency effect for words in temporal areas, no frequency effects were found for the two phrasal conditions. These findings support the conclusion that long and short phrases are analyzed, respectively, in the left frontal network and in a bilateral temporal network but are not retrieved from memory in the same way as simple words during spoken language comprehension.This research was supported by an Advanced Investigator grant to W. M. W. from the European Research Council (AdG 230570 NEUROLEX) and by MRC Cognition and Brain Sciences Unit (CBSU) funding to W. M. W. (U.1055.04.002.00001.01). Com- puting resources were provided by the MRC CBSU

Tonotopic representation of loudness in the human cortex

A prominent feature of the auditory system is that neurons show tuning to audio frequency; each neuron has a characteristic frequency (CF) to which it is most sensitive. Furthermore, there is an orderly mapping of CF to position, which is called tonotopic organization and which is observed at many levels of the auditory system. In a previous study (Thwaites et al., 2016) we examined cortical entrainment to two auditory transforms predicted by a model of loudness, instantaneous loudness and short-term loudness, using speech as the input signal. The model is based on the assumption that neural activity is combined across CFs (i.e. across frequency channels) before the transform to short-term loudness. However, it is also possible that short-term loudness is determined on a channel-specific basis. Here we tested these possibilities by assessing neural entrainment to the overall and channel-specific instantaneous loudness and the overall and channel-specific short-term loudness. The results showed entrainment to channel-specific instantaneous loudness at latencies of 45 and 100 ms (bilaterally, in and around Heschl's gyrus). There was entrainment to overall instantaneous loudness at 165 ms in dorso-lateral sulcus (DLS). Entrainment to overall short-term loudness occurred primarily at 275 ms, bilaterally in DLS and superior temporal sulcus. There was only weak evidence for entrainment to channel-specific short-term loudness.This work was supported by an ERC Advanced Grant (230570, ‘Neurolex’) to WMW, by MRC Cognition and Brain Sciences Unit (CBU) funding to WMW (U.1055.04.002.00001.01), and by EPSRC grant RG78536 to JS and BM

Domain-specific and Domain-general Processing in Left Perisylvian Cortex: Evidence from Russian

The processing of words containing inflectional affixes triggers morphophonological parsing and affix-related grammatical information processing. Increased perceptual complexity related to stem-affix parsing is hypothesized to create predominantly domain-general processing demands, whereas grammatical processing primarily implicates domain-specific linguistic demands. Exploiting the properties of Russian morphology and syntax, we designed an fMRI experiment to separate out the neural systems supporting these two demand types, contrasting inflectional complexity, syntactic (phrasal) complexity, and derivational complexity in three comparisons: (a) increase in parsing demands while controlling for grammatical complexity (inflections vs. phrases), (b) increase in grammatical processing demands, and (c) combined demands of morphophonological parsing and grammatical processing (inflections and phrases vs. derivations). Left inferior frontal and bilateral temporal areas are most active when the two demand types are combined, with inflectional and phrasal complexity contrasting strongly with derivational complexity (which generated only bilateral temporal activity). Increased stem-affix parsing demands alone did not produce unique activations, whereas grammatical structure processing activated bilateral superior and middle temporal areas. Selective left frontotemporal language system engagement for short phrases and inflections seems to be driven by simultaneous and interdependent domain-general and domain-specific processing demands

Relating Dynamic Brain States to Dynamic Machine States: Human and Machine Solutions to the Speech Recognition Problem

There is widespread interest in the relationship between the neurobiological systems supporting human cognition and emerging computational systems capable of emulating these capacities. Human speech comprehension, poorly understood as a neurobiological process, is an important case in point. Automatic Speech Recognition (ASR) systems with near-human levels of performance are now available, which provide a computationally explicit solution for the recognition of words in continuous speech. This research aims to bridge the gap between speech recognition processes in humans and machines, using novel multivariate techniques to compare incremental ‘machine states’, generated as the ASR analysis progresses over time, to the incremen- tal ‘brain states’, measured using combined electro- and magneto-encephalography (EMEG), generated as the same inputs are heard by human listeners. This direct comparison of dynamic human and machine internal states, as they respond to the same incrementally delivered sensory input, revealed a significant correspondence between neural response patterns in human superior temporal cortex and the structural properties of ASR-derived phonetic models. Spatially coherent patches in human temporal cortex responded selectively to individual phonetic features defined on the basis of machine-extracted regularities in the speech to lexicon mapping process. These results demonstrate the feasibility of relating human and ASR solutions to the problem of speech recognition, and suggest the potential for further studies relating complex neural computations in human speech comprehension to the rapidly evolving ASR systems that address the same problem domain.This research was supported financially by an Advanced Investigator grant to WMW from the European Research Council (AdG 230570 NEUROLEX), by MRC Cognition and Brain Sciences Unit (CBSU) funding to WMW (U.1055.04.002.00001.01), and by a European Research Council Advanced Investigator grant under the European Community’s Horizon 2020 Research and Innovation Programme (2014-2020 ERC Grant agreement no 669820) to Lorraine K. Tyler. LS was partly supported by the NIHR Biomedical Research Centre and Biomedical Unit in Dementia based at Cambridge University Hospital NHS Foundation Trust

The influence of 2-hop network density on spoken word recognition

The final publication is available at Springer via http://dx.doi.org/10.3758/s13423-016-1103-9The influence of 2-hop density on spoken word recognition was investigated. 2-hop density measures the density of connections among the phonological neighbors (i.e., 1-hop neighbors) and phonological neighbors of those neighbors (i.e., 2-hop neighbors) of a target word. In both naming and lexical decision tasks, words with low 2-hop density were recognized more quickly than words with high 2-hop density. Because stimuli were selected such that the number of 1-hop and 2-hop neighbors were matched across both sets of words, the results suggest that spoken word recognition is influenced by the amount of connectivity among distant neighbors of the target word—a result that is not easily accommodated by current models of spoken word recognition. A diffusion of activation framework is proposed to account for the present finding

Effects of Place of Articulation Changes on Auditory Neural Activity: A Magnetoencephalography Study

In casual speech, phonemic segments often assimilate such that they adopt features from adjacent segments, a typical feature being their place of articulation within the vocal tract (e.g., labial, coronal, velar). Place assimilation (e.g., from coronal /n/ to labial /m/: rainbow→*raimbow) alters the surface form of words. Listeners' ability to perceptually compensate for such changes seems to depend on the phonemic context, on whether the adjacent segment (e.g., the /b/ in “rainbow”) invites the particular change. Also, some assimilations occur frequently (e.g., /n/→/m/), others are rare (e.g., /m/→/n/). We investigated the effects of place assimilation, its contextual dependency, and its frequency on the strength of auditory evoked mismatch negativity (MMN) responses, using pseudowords. Results from magnetoencephalography (MEG) revealed that the MMN was modulated both by the frequency and contextual appropriateness of assimilations

Preserved cognitive functions with age are determined by domain-dependent shifts in network responsivity

Healthy ageing has disparate effects on different cognitive domains. The neural basis of these differences, however, is largely unknown. We investigated this question by using Independent Components Analysis to obtain functional brain components from 98 healthy participants aged 23-87 years from the population-based Cam-CAN cohort. Participants performed two cognitive tasks that show age-related decrease (fluid intelligence and object naming) and a syntactic comprehension task that shows age-related preservation. We report that activation of task-positive neural components predicts inter-individual differences in performance in each task across the adult lifespan. Furthermore, only the two tasks that show performance declines with age show age-related decreases in task-positive activation of neural components and decreasing default mode (DM) suppression. Our results suggest that distributed, multi-component brain responsivity supports cognition across the adult lifespan, and the maintenance of this, along with maintained DM deactivation, characterizes successful ageing and may explain differential ageing trajectories across cognitive domains

Multiple determinants of lifespan memory differences

Memory problems are among the most common complaints as people grow older. Using structural equation modeling of commensurate scores of anterograde memory from a large (N = 315), population-derived sample (www.cam-can.org), we provide evidence for three memory factors that are supported by distinct brain regions and show differential sensitivity to age. Associative memory and item memory are dramatically affected by age, even after adjusting for education level and fluid intelligence, whereas visual priming is not. Associative memory and item memory are differentially affected by emotional valence, and the age-related decline in associative memory is faster for negative than for positive or neutral stimuli. Gray-matter volume in the hippocampus, parahippocampus and fusiform cortex, and a white-matter index for the fornix, uncinate fasciculus and inferior longitudinal fasciculus, show differential contributions to the three memory factors. Together, these data demonstrate the extent to which differential ageing of the brain leads to differential patterns of memory loss

Does Sleep Improve Your Grammar? : Preferential Consolidation of Arbitrary Components of New Linguistic Knowledge

We examined the role of sleep-related memory consolidation processes in learning new form-meaning mappings. Specifically, we examined a Complementary Learning Systems account, which implies that sleep-related consolidation should be more beneficial for new hippocampally dependent arbitrary mappings (e.g. new vocabulary items) relative to new systematic mappings (e.g. grammatical regularities), which can be better encoded neocortically. The hypothesis was tested using a novel language with an artificial grammatical gender system. Stem-referent mappings implemented arbitrary aspects of the new language, and determiner/suffix+natural gender mappings implemented systematic aspects (e.g. tib scoiffesh + ballerina, tib mofeem + bride; ked jorool + cowboy, ked heefaff + priest). Importantly, the determiner-gender and the suffix-gender mappings varied in complexity and salience, thus providing a range of opportunities to detect beneficial effects of sleep for this type of mapping. Participants were trained on the new language using a word-picture matching task, and were tested after a 2-hour delay which included sleep or wakefulness. Participants in the sleep group outperformed participants in the wake group on tests assessing memory for the arbitrary aspects of the new mappings (individual vocabulary items), whereas we saw no evidence of a sleep benefit in any of the tests assessing memory for the systematic aspects of the new mappings: Participants in both groups extracted the salient determiner-natural gender mapping, but not the more complex suffix-natural gender mapping. The data support the predictions of the complementary systems account and highlight the importance of the arbitrariness/systematicity dimension in the consolidation process for declarative memories

Behavioural plasticity in Argyrodes antipodiana.

In this thesis I look at the behavioural plasticity of A.antipodiana while foraging and interacting with conspecifics. The aim is to see how an animal, with apparently limited intelligence is able to deal with environmental variability. I conclude that A.antipodiana appears to have been able to overcome these limitations very effectively by using four methods, of which the first two effectively reduce the amount of learning necessary. Firstly, A.antipodiana simplifies the situation as much as possible by ignoring some of the variability. For example, in social situations, A.antipodiana appears to recognize only a few categories of conspecifics, rather than recognize conspecifics as individuals. Consequently, the social groups of A.antipodiana are not characterized by complicating factors such as aggressive orders or dominance hierarchies. Secondly, A.antipodiana largely ignores current situations and instead bases much of its behavioural decisions on its internal conditions. For example, in male-male conflicts, males respond to their opponents largely according to their own size, level of hunger, the number of contests in which they have participated, or their past experience of winning or losing; rather than to the characteristics of their opponents. Likewise in social interactions A.antipodiana is more inclined to be aggressive towards conspecifics if it itself is foraging, and less inclined if it is feeding with the host. When the situation cannot be simplified, A.antipodiana adopts a third response which is to use the simplest method possible for adjusting its behaviour. For example, A.antipodiana's ability to develop araneophagic skills appears to be governed by a critical period. Thus rather than developing araneophagic skills through trial and error with practise, the ability appears to be simply 'switched on' if required. The fourth means by which A.antipodiana compensates for limited intelligence is to be very selective in the areas in which it does use learning. For example, A.antipodiana seems to use problem solving when foraging, but only when it is actually trying to reach the food bundle upon which the host is feeding. A.antipodiana also seems to learn to move more stealthfully on the host's web. There is even evidence that A.antipodiana may behave with intent when males are competing for opportunities to copulate. Thus the intelligence of A.antipodiana appears to be severely limited. However, despite these limitations, by reducing the amount of learning necessary, and then channelling what learning it has to very restricted, but very poignant areas of its behaviour, A.antipodiana, is able to respond very effectively to its variable environment