Publications

Abstract

Variation in plasma levels of cortisol, an essential hormone in the stress response, is associated in population-based studies with cardio-metabolic, inflammatory and neuro-cognitive traits and diseases. Heritability of plasma cortisol is estimated at 30-60% but no common genetic contribution has been identified. The CORtisol NETwork (CORNET) consortium undertook genome wide association meta-analysis for plasma cortisol in 12,597 Caucasian participants, replicated in 2,795 participants. The results indicate that <1% of variance in plasma cortisol is accounted for by genetic variation in a single region of chromosome 14. This locus spans SERPINA6, encoding corticosteroid binding globulin (CBG, the major cortisol-binding protein in plasma), and SERPINA1, encoding α1-antitrypsin (which inhibits cleavage of the reactive centre loop that releases cortisol from CBG). Three partially independent signals were identified within the region, represented by common SNPs; detailed biochemical investigation in a nested sub-cohort showed all these SNPs were associated with variation in total cortisol binding activity in plasma, but some variants influenced total CBG concentrations while the top hit (rs12589136) influenced the immunoreactivity of the reactive centre loop of CBG. Exome chip and 1000 Genomes imputation analysis of this locus in the CROATIA-Korcula cohort identified missense mutations in SERPINA6 and SERPINA1 that did not account for the effects of common variants. These findings reveal a novel common genetic source of variation in binding of cortisol by CBG, and reinforce the key role of CBG in determining plasma cortisol levels. In turn this genetic variation may contribute to cortisol-associated degenerative diseases.

Abstract

Alphabetic orthographies show more or less ambiguous relations between spelling and sound patterns. In transparent orthographies, like Italian, the pronunciation can be predicted from the spelling and vice versa. Opaque orthographies, like English, often display unpredictable spelling–sound correspondences. In this paper we present a computational analysis of word-initial bi-directional spelling–sound correspondences for Dutch, English, French, German, and Hungarian, stated in entropy values for various grain sizes. This allows us to position the five languages on the continuum from opaque to transparent orthographies, both in spelling-to-sound and sound-to-spelling directions. The analysis is based on metrics derived from information theory, and therefore independent of any specific theory of visual word recognition as well as of any specific theoretical approach of orthography.

Abstract

Hierarchical predictive coding has been identified as a possible unifying principle of brain function, and recent work in cognitive neuroscience has examined how it may be affected by age–related changes. Using language comprehension as a test case, the present study aimed to dissociate age-related changes in prediction generation versus internal model adaptation following a prediction error. Event-related brain potentials (ERPs) were measured in a group of older adults (60–81 years; n = 40) as they read sentences of the form “The opposite of black is white/yellow/nice.” Replicating previous work in young adults, results showed a target-related P300 for the expected antonym (“white”; an effect assumed to reflect a prediction match), and a graded N400 effect for the two incongruous conditions (i.e. a larger N400 amplitude for the incongruous continuation not related to the expected antonym, “nice,” versus the incongruous associated condition, “yellow”). These effects were followed by a late positivity, again with a larger amplitude in the incongruous non-associated versus incongruous associated condition. Analyses using linear mixed-effects models showed that the target-related P300 effect and the N400 effect for the incongruous non-associated condition were both modulated by age, thus suggesting that age-related changes affect both prediction generation and model adaptation. However, effects of age were outweighed by the interindividual variability of ERP responses, as reflected in the high proportion of variance captured by the inclusion of by-condition random slopes for participants and items. We thus argue that – at both a neurophysiological and a functional level – the notion of general differences between language processing in young and older adults may only be of limited use, and that future research should seek to better understand the causes of interindividual variability in the ERP responses of older adults and its relation to cognitive performance.

Abstract

Speech comprehension involves extensive use of prediction. Linguistic prediction may be guided by the semantics or syntax, but also by the performance characteristics of the speech signal, such as disfluency. Previous studies have shown that listeners, when presented with the filler uh, exhibit a disfluency bias for discourse-new or unknown referents, drawing inferences about the source of the disfluency. The goal of the present study is to study the contrast between native and non-native disfluencies in speech comprehension. Experiment 1 presented listeners with pictures of high-frequency (e.g., a hand) and low-frequency objects (e.g., a sewing machine) and with fluent and disfluent instructions. Listeners were found to anticipate reference to low-frequency objects when encountering disfluency, thus attributing disfluency to speaker trouble in lexical retrieval. Experiment 2 showed that, when participants listened to disfluent non-native speech, no anticipation of low-frequency referents was observed. We conclude that listeners can adapt their predictive strategies to the (non-native) speaker at hand, extending our understanding of the role of speaker identity in speech comprehension.

Abstract

Where native speakers supposedly are fluent by default, non-native speakers often have to strive hard to achieve a native-like fluency level. However, disfluencies (such as pauses, fillers, repairs, etc.) occur in both native and non-native speech and it is as yet unclear ow luency raters weigh the fluency characteristics of native and non-native speech. Two rating experiments compared the way raters assess the luency of native and non-native speech. The fluency characteristics of native and non- native speech were controlled by using phonetic anipulations in pause (Experiment 1) and speed characteristics (Experiment 2). The results show that the ratings on manipulated native and on-native speech were affected in a similar fashion. This suggests that there is no difference in the way listeners weigh the fluency haracteristics of native and non-native speakers.

Abstract

A central motif in neuronal networks is convergence, linking several input neurons to one target neuron. In visual cortex, convergence renders target neurons responsive to complex stimuli. Yet, convergence typically sends multiple stimuli to a target, and the behaviorally relevant stimulus must be selected. We used two stimuli, activating separate electrocorticographic V1 sites, and both activating an electrocorticographic V4 site equally strongly. When one of those stimuli activated one V1 site, it gamma synchronized (60-80 Hz) to V4. When the two stimuli activated two V1 sites, primarily the relevant one gamma synchronized to V4. Frequency bands of gamma activities showed substantial overlap containing the band of interareal coherence. The relevant V1 site had its gamma peak frequency 2-3 Hz higher than the irrelevant V1 site and 4-6 Hz higher than V4. Gamma-mediated interareal influences were predominantly directed from V1 to V4. We propose that selective synchronization renders relevant input effective, thereby modulating effective connectivity.

Abstract

Relational grammar was proposed in Suppes (1976) as a semantical grammar for natural language. Fragments considered so far are restricted to distributive notions. In this article, relational grammar is extended to collective notions.

Abstract

There are two competing hypotheses for the origin of the Indo-European language family. The conventional view places the homeland in the Pontic steppes about 6000 years ago. An alternative hypothesis claims that the languages spread from Anatolia with the expansion of farming 8000 to 9500 years ago. We used Bayesian phylogeographic approaches, together with basic vocabulary data from 103 ancient and contemporary Indo-European languages, to explicitly model the expansion of the family and test these hypotheses. We found decisive support for an Anatolian origin over a steppe origin. Both the inferred timing and root location of the Indo-European language trees fit with an agricultural expansion from Anatolia beginning 8000 to 9500 years ago. These results highlight the critical role that phylogeographic inference can play in resolving debates about human prehistory.

Abstract

This study reports three experiments on how children learning Mandarin Chinese comprehend and use aspect markers. These experiments examine the role of lexical aspect in children's acquisition of grammatical aspect. Results provide converging evidence for children's early sensitivity to (1) the association between atelic verbs and the imperfective aspect markers zai, -zhe, and -ne, and (2) the association between telic verbs and the perfective aspect marker -le. Children did not show a sensitivity in their use or understanding of aspect markers to the difference between stative and activity verbs or between semelfactive and activity verbs. These results are consistent with Slobin's (1985) basic child grammar hypothesis that the contrast between process and result is important in children's early acquisition of temporal morphology. In contrast, they are inconsistent with Bickerton's (1981, 1984) language bioprogram hypothesis that the distinctions between state and process and between punctual and nonpunctual are preprogrammed into language learners. We suggest new ways of looking at the results in the light of recent probabilistic hypotheses that emphasize the role of input, prototypes and connectionist representations.

Abstract

In this study, we investigated the level of visual processing at which surface colour information improves the naming of colour diagnostic and noncolour diagnostic objects. Continuous electroencephalograms were recorded while participants performed a visual object naming task in which coloured and black-and-white versions of both types of objects were presented. The black-and-white and the colour presentations were compared in two groups of event-related potentials (ERPs): (1) The P1 and N1 components, indexing early visual processing; and (2) the N300 and N400 components, which index late visual processing. A colour effect was observed in the P1 and N1 components, for both colour and noncolour diagnostic objects. In addition, for colour diagnostic objects, a colour effect was observed in the N400 component. These results suggest that colour information is important for the naming of colour and noncolour diagnostic objects at different levels of visual processing. It thus appears that the visual system uses colour information, during naming of both object types, at early visual stages; however, for the colour diagnostic objects naming, colour information is also recruited during the late visual processing stages.

Abstract

In this study, we used event-related potentials (ERPs) to evaluate the contribution of surface color and color knowledge information in object identification. We constructed two color-object verification tasks – a surface and a knowledge verification task – using high color diagnostic objects; both typical and atypical color versions of the same object were presented. Continuous electroencephalogram was recorded from 26 subjects. A cluster randomization procedure was used to explore the differences between typical and atypical color objects in each task. In the color knowledge task, we found two significant clusters that were consistent with the N350 and late positive complex (LPC) effects. Atypical color objects elicited more negative ERPs compared to typical color objects. The color effect found in the N350 time window suggests that surface color is an important cue that facilitates the selection of a stored object representation from long-term memory. Moreover, the observed LPC effect suggests that surface color activates associated semantic knowledge about the object, including color knowledge representations. We did not find any significant differences between typical and atypical color objects in the surface color verification task, which indicates that there is little contribution of color knowledge to resolve the surface color verification. Our main results suggest that surface color is an important visual cue that triggers color knowledge, thereby facilitating object identification.

Abstract

The present study used electrophysiological and behavioral measures to investigate the perception of an English stop consonant contrast by native English listeners and by native Dutch listeners who were highly proficient in English. A /ba/-/pa/ continuum was created from a naturally produced /pa/ token by removing successive periods of aspiration, thus reducing the voice onset time. Although aspiration is a relevant cue for distinguishing voiced and unvoiced labial stop consonants (/b/ and /p/) in English, prevoicing is the primary cue used to distinguish between these categories in Dutch. In the electrophysiological experiment, participants listened to oddball sequences containing the standard /pa/ stimulus and one of three deviant stimuli while the mismatch-negativity response was measured. Participants then completed an identification task on the same stimuli. The results showed that native English participants were more sensitive to reductions in aspiration than native Dutch participants, as indicated by shifts in the category boundary, by differing within-group patterns of mismatch-negativity responses, and by larger mean evoked potential amplitudes in the native English group for two of the three deviant stimuli. This between-group difference in the sensorineural processing of aspiration cues indicates that native language experience alters the way in which the acoustic features of speech are processed in the auditory brain, even following extensive second-language training.

Abstract

It has been fifty years since Levelt’s monograph On Binocular Rivalry (1965) was published, but its four propositions that describe the relation between stimulus strength and the phenomenology of binocular rivalry remain a benchmark for theorists and experimentalists even today. In this review, we will revisit the original conception of the four propositions and the scientific landscape in which this happened. We will also provide a brief update concerning distributions of dominance durations, another aspect of Levelt’s monograph that has maintained a prominent presence in the field. In a critical evaluation of Levelt’s propositions against current knowledge of binocular rivalry we will then demonstrate that the original propositions are not completely compatible with what is known today, but that they can, in a straightforward way, be modified to encapsulate the progress that has been made over the past fifty years. The resulting modified, propositions are shown to apply to a broad range of bistable perceptual phenomena, not just binocular rivalry, and they allow important inferences about the underlying neural systems. We argue that these inferences reflect canonical neural properties that play a role in visual perception in general, and we discuss ways in which future research can build on the work reviewed here to attain a better understanding of these properties

Abstract

Automated processing of large amounts of scientific data, especially across domains, requires that the data can be selected and parsed without human intervention. Precise characterization of that data, as in typing, is needed once the processing goes beyond the realm of domain specific or local research group assumptions. The Research Data Alliance (RDA) Data Type Registries Working Group (DTR-WG) was assembled to address this issue through the creation of a Data Type Registry methodology, data model, and prototype. The WG was approved by the RDA Council during March of 2013 and will complete its work in mid-2014, in between the third and fourth RDA Plenaries.

Abstract

This study investigates how inaccurate phoneme processing affects recognition of partially onset-overlapping pairs like DAFFOdil-DEFIcit and of minimal pairs like flash-flesh in second-language listening. Two cross-modal priming experiments examined differences between native (L1) and second-language (L2) listeners at two stages of lexical processing: first, the activation of intended and mismatching lexical representations and second, the competition between those lexical representations. Experiment 1 shows that truncated primes like daffo- and defi- activated lexical representations of mismatching words (either deficit or daffodil) more for L2 listeners than for L1 listeners. Experiment 2 shows that for minimal pairs, matching primes (prime: flash, target: FLASH) facilitated recognition of visual targets for L1 and L2 listeners alike, whereas mismatching primes (flesh, FLASH) inhibited recognition consistently for L1 listeners but only in a minority of cases for L2 listeners; in most cases, for them, primes facilitated recognition of both words equally strongly. Thus, L1 and L2 listeners' results differed both at the stages of lexical activation and competition. First, perceptually difficult phonemes activated mismatching words more for L2 listeners than for L1 listeners, and second, lexical competition led to efficient inhibition of mismatching competitors for L1 listeners but in most cases not for L2 listeners.

Abstract

Background: According to decades of research on affective motivation in the human brain, approach motivational states are supported primarily by the left hemisphere and avoidance states by the right hemisphere. The underlying cause of this specialization, however, has remained unknown. Here we conducted a first test of the Sword and Shield Hypothesis (SSH), according to which the hemispheric laterality of affective motivation depends on the laterality of motor control for the dominant hand (i.e., the "sword hand," used preferentially to perform approach actions) and the nondominant hand (i.e., the "shield hand," used preferentially to perform avoidance actions). Methodology/Principal Findings: To determine whether the laterality of approach motivation varies with handedness, we measured alpha-band power (an inverse index of neural activity) in right- and left-handers during resting-state electroencephalography and analyzed hemispheric alpha-power asymmetries as a function of the participants' trait approach motivational tendencies. Stronger approach motivation was associated with more left-hemisphere activity in right-handers, but with more right-hemisphere activity in left-handers. Conclusions: The hemispheric correlates of approach motivation reversed between right- and left-handers, consistent with the way they typically use their dominant and nondominant hands to perform approach and avoidance actions. In both right- and left-handers, approach motivation was lateralized to the same hemisphere that controls the dominant hand. This covariation between neural systems for action and emotion provides initial support for the SSH

Abstract

Three eye-tracking experiments investigated how phonological reductions (e.g., ‘‘puter’’ for ‘‘computer’’) modulate phonological competition. Participants listened to sentences extracted from a pontaneous speech corpus and saw four printed words: a target (e.g., ‘‘computer’’), a competitor similar to the canonical form (e.g., ‘‘companion’’), one similar to the reduced form (e.g.,
‘‘pupil’’), and an unrelated distractor. In Experiment 1, we presented canonical and reduced forms in a syllabic and in a sentence context. Listeners directed
their attention to a similar degree to both competitors independent of the
target’s spoken form. In Experiment 2, we excluded reduced forms and
presented canonical forms only. In such a listening situation, participants
showed a clear preference for the ‘‘canonical form’’ competitor. In Experiment 3, we presented canonical forms intermixed with reduced forms in a sentence context and replicated the competition pattern of Experiment 1. These data suggest that listeners penalize acoustic mismatches less strongly when listeningto reduced speech than when listening to fully articulated speech. We conclude
that flexibility to adjust to speech-intrinsic factors is a key feature of the spoken word recognition system.

Abstract

This study examined the influence of background language variation on speech recognition. English listeners performed an English sentence recognition task in either “pure” background conditions in which all trials had either English or Dutch background babble or in mixed background conditions in which the background language varied across trials (i.e., a mix of English and Dutch or one of these background languages mixed with quiet trials). This design allowed the authors to compare performance on identical trials across pure and mixed conditions. The data reveal that speech-in-speech recognition is sensitive to contextual variation in terms of the target-background language (mis)match depending on the relative ease/difficulty of the test trials in relation to the surrounding trials.

Abstract

In listeners' daily communicative exchanges, they most often hear casual speech, in which words are often produced with fewer segments, rather than the careful speech used in most psycholinguistic experiments. Three experiments examined phonological competition during the recognition of reduced forms such as [pjutər] for computer using a target-absent variant of the visual world paradigm. Listeners' eye movements were tracked upon hearing canonical and reduced forms as they looked at displays of four printed words. One of the words was phonologically similar to the canonical pronunciation of the target word, one word was similar to the reduced pronunciation, and two words served as unrelated distractors. When spoken targets were presented in isolation (Experiment 1) and in sentential contexts (Experiment 2), competition was modulated as a function of the target word form. When reduced targets were presented in sentential contexts, listeners were probabilistically more likely to first fixate reduced-form competitors before shifting their eye gaze to canonical-form competitors. Experiment 3, in which the original /p/ from [pjutər] was replaced with a “real” onset /p/, showed an effect of cross-splicing in the late time window. We conjecture that these results fit best with the notion that speech reductions initially activate competitors that are similar to the phonological surface form of the reduction, but that listeners nevertheless can exploit fine phonetic detail to reconstruct strongly reduced forms to their canonical counterparts.

Abstract

In traditional theories of language comprehension, syntactic and semantic processing are inextricably linked. This assumption has been challenged by the ‘Semantic Illusion Effect’ found in studies using Event Related brain Potentials. Semantically anomalous sentences did not produce the expected increase in N400 amplitude but rather one in P600 amplitude. To explain these findings, complex models have been devised in which an independent semantic processing stream can arrive at a sentence interpretation that may differ from the interpretation prescribed by the syntactic structure of the sentence. We review five such multi-stream models and argue that they do not account for the full range of relevant results because they assume that the amplitude of the N400 indexes some form of semantic integration. Based on recent evidence we argue that N400 amplitude might reflect the retrieval of lexical information from memory. On this view, the absence of an N400-effect in Semantic Illusion sentences can be explained in terms of priming. Furthermore, we suggest that semantic integration, which has previously been linked to the N400 component, might be reflected in the P600 instead. When combined, these functional interpretations result in a single-stream account of language processing that can explain all of the Semantic Illusion data.

Abstract

This study examined whether speech-on-speech masking is sensitive to variation in the degree of similarity between the target and the masker speech. Three experiments investigated whether speech-in-speech recognition varies across different background speech languages (English vs Dutch) for both English and Dutch targets, as well as across variation in the semantic content of the background speech (meaningful vs semantically anomalous sentences), and across variation in listener status vis-à-vis the target and masker languages (native, non-native, or unfamiliar). The results showed that the more similar the target speech is to the masker speech (e.g., same vs different language, same vs different levels of semantic content), the greater the interference on speech recognition accuracy. Moreover, the listener’s knowledge of the target and the background language modulate the size of the release from masking. These factors had an especially strong effect on masking effectiveness in highly unfavorable listening conditions. Overall this research provided evidence that that the degree of target-masker similarity plays a significant role in speech-in-speech recognition. The results also give insight into how listeners assign their resources differently depending on whether they are listening to their first or second language

Abstract

This study examined the temporal dynamics of spoken word recognition in noise and background speech. In two visual-world experiments, English participants listened to target words while looking at four pictures on the screen: a target (e.g. candle), an onset competitor (e.g. candy), a rhyme competitor (e.g. sandal), and an unrelated distractor (e.g. lemon). Target words were presented in quiet, mixed with broadband noise, or mixed with background speech. Results showed that lexical competition changes throughout the observation window as a function of what is presented in the background. These findings suggest that, rather than being strictly sequential, stream segregation and lexical competition interact during spoken word recognition

Abstract

A major finding in studies of early vocabulary acquisition has been that children tend to learn a lot of nouns early but make do with relatively few verbs, among which semantically general-purpose verbs like do, make, get, have, give, come, go, and be play a prominent role. The preponderance of nouns is explained in terms of nouns labelling concrete objects beings “easier” to learn than verbs, which label relational categories. Nouns label “natural categories” observable in the world, verbs label more linguistically and culturally specific categories of events linking objects belonging to such natural categories (Gentner 1978, 1982; Clark 1993). This view has been challenged recently by data from children learning certain non-Indo-European languges like Korean, where children have an early verb explosion and verbs dominate in early child utterances. Children learning the Mayan language Tzeltal also acquire verbs early, prior to any noun explosion as measured by production. Verb types are roughly equivalent to noun types in children’s beginning production vocabulary and soon outnumber them. At the one-word stage children’s verbs mostly have the form of a root stripped of affixes, correctly segmented despite structural difficulties. Quite early (before the MLU 2.0 point) there is evidence of productivity of some grammatical markers (although they are not always present): the person-marking affixes cross-referencing core arguments, and the completive/incompletive aspectual distinctions. The Tzeltal facts argue against a natural-categories explanation for childre’s early vocabulary, in favor of a view emphasizing the early effects of language-specific properties of the input. They suggest that when and how a child acquires a “verb” category is centrally influenced by the structural properties of the input, and that the semantic structure of the language - where the referential load is concentrated - plays a fundamental role in addition to distributional facts.

Abstract

When Tzeltal children in the Mayan community of Tenejapa, in southern Mexico, begin speaking, their production vocabulary consists predominantly of verb roots, in contrast to the dominance of nouns in the initial vocabulary of first‐language learners of Indo‐European languages. This article proposes that a particular Tzeltal conversational feature—known in the Mayanist literature as "dialogic repetition"—provides a context that facilitates the early analysis and use of verbs. Although Tzeltal babies are not treated by adults as genuine interlocutors worthy of sustained interaction, dialogic repetition in the speech the children are exposed to may have an important role in revealing to them the structural properties of the language, as well as in socializing the collaborative style of verbal interaction adults favor in this community.

Abstract

A study is presented on the effects of discourse–semantic and lexical–syntactic information during spoken sentence processing. Event-related brain potentials (ERPs) were registered while subjects listened to discourses that ended in a sentence with a temporary syntactic ambiguity. The prior discourse–semantic information biased toward one analysis of the temporary ambiguity, whereas the lexical-syntactic information allowed only for the alternative analysis. The ERP results show that discourse–semantic information can momentarily take precedence over syntactic information, even if this violates grammatical gender agreement rules.

Abstract

Two experiments are reported that provide evidence on task-induced effects during
visual lexical processing in a primetarget semantic priming paradigm. The research focuses on target expectancy effects by manipulating the proportion of semantically related and unrelated word pairs. In Experiment 1, a lexical decision task was used and reaction times (RTs) and event-related brain potentials (ERPs) were obtained. In Experiment 2, subjects silently read the stimuli, without any additional task demands, and ERPs were recorded. The RT and ERP results of Experiment 1 demonstrate that an expectancy mechanism contributed to the priming effect when a high proportion of related word pairs was presented. The ERP results of Experiment 2 show that in the absence of extraneous task requirements, an expectancy mechanism is not active. However, a standard ERP semantic priming effect was obtained in Experiment 2. The combined results show that priming effects due to relatedness proportion are induced by task demands and are not a standard aspect of online lexical processing.

Abstract

This work was supported by the Max Planck Institute for Psycholinguistics and the Nederlandse Organisatie voor Wetenschappelijk Onderzoek (NWO; MPI 56-384, The Dynamics of Multilingual Processing, awarded to M Gullberg and P Indefrey).

Abstract

This is a Spanish translation of Brown 1997.

Abstract

Linguistic expressions of time often draw on spatial language, which raises the question of whether cultural specificity in spatial language and cognition is reflected in thinking about time. In the Mayan language Tzeltal, spatial language relies heavily on an absolute frame of reference utilizing the overall slope of the land, distinguishing an “uphill/downhill” axis oriented from south to north, and an orthogonal “crossways” axis (sunrise-set) on the basis of which objects at all scales are located. Does this absolute system for calculating spa-tial relations carry over into construals of temporal relations? This question was explored in a study where Tzeltal consultants produced temporal expressions and performed two different non-linguistic temporal ordering tasks. The results show that at least five distinct schemata for conceptualizing time underlie Tzeltal linguistic expressions: (i) deictic ego-centered time, (ii) time as an ordered sequence (e.g., “first”/“later”), (iii) cyclic time (times of the day, seasons), (iv) time as spatial extension or location (e.g., “entering/exiting July”), and (v) a time vector extending uphillwards into the future. The non-linguistic task results showed that the “time moves uphillwards” metaphor, based on the absolute frame of reference prevalent in Tzeltal spatial language and thinking and important as well in the linguistic expressions for time, is not strongly reflected in responses on these tasks. It is argued that systematic and consistent use of spatial language in an absolute frame of reference does not necessarily transfer to consistent absolute time conceptualization in non-linguistic tasks; time appears to be more open to alternative construals.

Abstract

Speaking begins with the formulation of an intended preverbal message and linguistic encoding of this information. The transition from thought to speech occurs incrementally, with cascading planning at subsequent levels of production. In this article, we aim to specify the mechanisms that support incremental message preparation. We contrast two hypotheses about the mechanisms responsible for incorporating message-level information into a linguistic plan. According to the Initial Preparation view, messages can be encoded as fluent utterances if all information is ready before speaking begins. By contrast, on the Continuous Incrementality view, messages can be continually prepared and updated throughout the production process, allowing for fluent production even if new information is added to the message while speaking is underway. Testing these hypotheses, eye-tracked speakers in two experiments produced unscripted, conjoined noun phrases with modifiers. Both experiments showed that new message elements can be incrementally incorporated into the utterance even after articulation begins, consistent with a Continuous Incrementality view of message planning, in which messages percolate to linguistic encoding immediately as that information becomes available in the mind of the speaker. We conclude by discussing the functional role of incremental message planning in conversational speech and the situations in which this continuous incremental planning would be most likely to be observed

Abstract

Genes of the Major Histocompatibility Complex (MHC) have recently been shown to have neuronal functions in the thalamus and hippocampus. Common genetic variants in the Human Leukocyte Antigens (HLA) region, human homologue of the MHC locus, are associated with small effects on susceptibility to schizophrenia, while volumetric changes of the thalamus and hippocampus have also been linked to schizophrenia. We therefore investigated whether common variants of the HLA would affect volumetric variation of the thalamus and hippocampus. We analyzed thalamus and hippocampus volumes, as measured using structural magnetic resonance imaging, in 1.265 healthy participants. These participants had also been genotyped using genome-wide single nucleotide polymorphism (SNP) arrays. We imputed genotypes for single nucleotide polymorphisms at high density across the HLA locus, as well as HLA allotypes and HLA amino acids, by use of a reference population dataset that was specifically targeted to the HLA region. We detected a significant association of the SNP rs17194174 with thalamus volume (nominal P=0.0000017, corrected P=0.0039), as well as additional SNPs within the same region of linkage disequilibrium. This effect was largely lateralized to the left thalamus and is localized within a genomic region previously associated with schizophrenia. The associated SNPs are also clustered within a potential regulatory element, and a region of linkage disequilibrium that spans genes expressed in the thalamus, including HLA-A. Our data indicate that genetic variation within the HLA region influences the volume and asymmetry of the human thalamus. The molecular mechanisms underlying this association may relate to HLA influences on susceptibility to schizophrenia

Abstract

Epigenetic effects on psychiatric traits remain relatively under-studied, and it remains unclear what the sizes of individual epigenetic effects may be, or how they vary between different clinical populations. The gene LRRTM1 (chromosome 2p12) has previously been linked and associated with schizophrenia in a parent-of-origin manner in a set of affected siblings (LOD = 4.72), indirectly suggesting a disruption of paternal imprinting at this locus in these families. From the same set of siblings that originally showed strong linkage at this locus, we analyzed 99 individuals using 454-bisulfite sequencing, from whole blood DNA, to measure the level of DNA methylation in the promoter region of LRRTM1. We also assessed seven additional loci that would be informative to compare. Paternal identity-by-descent sharing at LRRTM1, within sibling pairs, was linked to their similarity of methylation at the gene's promoter. Reduced methylation at the promoter showed a significant association with schizophrenia. Sibling pairs concordant for schizophrenia showed more similar methylation levels at the LRRTM1 promoter than diagnostically discordant pairs. The alleles of common SNPs spanning the locus did not explain this epigenetic linkage, which can therefore be considered as largely independent of DNA sequence variation and would not be detected in standard genetic association analysis. Our data suggest that hypomethylation at the LRRTM1 promoter, particularly of the paternally inherited allele, was a risk factor for the development of schizophrenia in this set of siblings affected with familial schizophrenia, and that had previously showed linkage at this locus in an affected-sib-pair context.

Abstract

The Hmong Diaspora is one of the widest modern human migrations. Mainly localised in South-East Asia, the United States of America, and metropolitan France, a small community has also settled the Amazonian forest of French Guiana. We have biologically analysed 62 individuals of this unique Guianese population through three complementary genetic markers: mitochondrial DNA (HVS-I/II and coding region SNPs), Y-chromosome (SNPs and STRs), and the Gm allotypic system. All genetic systems showed a high conservation of the Asian gene pool (Asian ancestry: mtDNA = 100.0%; NRY = 99.1%; Gm = 96.6%), without a trace of founder effect. When compared across various Asian populations, the highest correlations were observed with Hmong-Mien groups still living in South-East Asia (Fst < 0.05; P-value < 0.05). Despite a long history punctuated by exodus, the French Guianese Hmong have maintained their original genetic diversity.

Abstract

Several neurodevelopmental disorders are associated with preference for routine and challenging behavior following changes to routines. We examine individuals with Prader–Willi syndrome, who show elevated levels of this behavior, to better understand how previous experience of a routine can affect challenging behavior elicited by disruption to that routine. Play based challenges exposed 16 participants to routines, which were either adhered to or changed. Temper outburst behaviors, heart rate and movement were measured. As participants were exposed to routines for longer before a change (between 10 and 80 min; within participants), more temper outburst behaviors were elicited by changes. Increased emotional arousal was also elicited, which was indexed by heart rate increases not driven by movement. Further study will be important to understand whether current intervention approaches that limit exposure to changes, may benefit from the structured integration of flexibility to ensure that the opportunity for routine establishment is also limited.

Abstract

Nuclear receptors (NRs) form a large superfamily of transcription factors that participate in virtually every key biological process. They control development, fertility, gametogenesis and are misregulated in many cancers. Their enormous functional plasticity as transcription factors relates in part to NR-mediated interactions with hundreds of coregulatory proteins upon ligand (e.g., hormone) binding to their ligand binding domains (LBD), or following covalent modification. Some coregulator association relates to the distinct residues that shape a coactivator binding pocket termed AF-2, a surface groove that primarily determines the preference and specificity of protein–protein interactions. However, the highly conserved AF-2 pocket in the NR superfamily appears to be insufficient to account for NR subtype specificity leading to fine transcriptional modulation in certain settings. Additional protein–protein interaction surfaces, most notably on their LBD, may contribute to modulating NR function. NR coregulators and chaperones, normally much larger than the NR itself, may also bind to such interfaces. In the case of the androgen receptor (AR) LBD surface, structural and functional data highlighted the presence of another site named BF-3, which lies at a distinct but topographically adjacent surface to AF-2. AR BF-3 is a hot spot for mutations involved in prostate cancer and androgen insensitivity syndromes, and some FDA-approved drugs bind at this site. Structural studies suggested an allosteric relationship between AF-2 and BF-3, as occupancy of the latter affected coactivator recruitment to AF-2. Physiological relevant partners of AR BF-3 have not been described as yet. The newly discovered site is highly conserved among the steroid receptors subclass, but is also present in other NRs. Several missense mutations in the BF-3 regions of these human NRs are implicated in pathology and affect their function in vitro. The fact that AR BF-3 pocket is a druggable site evidences its pharmacological potential. Compounds that may affect allosterically NR function by binding to BF-3 open promising avenues to develop type-specific NR modulators.

Abstract

Heschl's gyrus (HG) is a core region of the auditory cortex whose morphology is highly variable across individuals. This variability has been linked to sound perception ability in both speech and music domains. Previous studies show that variations in morphological features of HG, such as cortical surface area and thickness, are heritable. To identify genetic variants that affect HG morphology, we conducted a genome-wide association scan (GWAS) meta-analysis in 3054 healthy individuals using HG surface area and thickness as quantitative traits. None of the single nucleotide polymorphisms (SNPs) showed association P values that would survive correction for multiple testing over the genome. The most significant association was found between right HG area and SNP rs72932726 close to gene DCBLD2 (3q12.1; P=2.77x10(-7)). This SNP was also associated with other regions involved in speech processing. The SNP rs333332 within gene KALRN (3q21.2; P=2.27x10(-6)) and rs143000161 near gene COBLL1 (2q24.3; P=2.40x10(-6)) were associated with the area and thickness of left HG, respectively. Both genes are involved in the development of the nervous system. The SNP rs7062395 close to the X-linked deafness gene POU3F4 was associated with right HG thickness (Xq21.1; P=2.38x10(-6)). This is the first molecular genetic analysis of variability in HG morphology

Abstract

This study examined whether musical training, ethnicity, and experience with a natural tone language influenced sensitivity to tone while listening to an artificial tone language. The language was designed with three tones, modeled after level-tone African languages. Participants listened to a 15-min random concatenation of six 3-syllable words. Sensitivity to tone was assessed using minimal pairs differing only in one syllable (nonword task: e.g., to-kà-su compared to ca-fí-to) or only in tone (tone task: e.g., to-kà-su compared to to-ká-su). Proficiency in an East Asian heritage language was the strongest predictor of success on the tone task. Asians without tone language experience were no better than other ethnic groups. We conclude by considering implications for research on second language learning, especially as approached through artificial language learning.

Abstract

Pediatric myelodysplastic syndrome (MDS) is a heterogeneous disease covering a spectrum ranging from aplasia (RCC) to myeloproliferation (RAEB(t)). In adult-type MDS there is increasing evidence for abnormal function of the bone-marrow microenvironment. Here, we extensively studied the mesenchymal stromal cells (MSCs) derived from children with MDS. MSCs were expanded from the bone-marrow of 17 MDS patients (RCC: n = 10 and advanced MDS: n = 7) and pediatric controls (n = 10). No differences were observed with respect to phenotype, differentiation capacity, immunomodulatory capacity or hematopoietic support. mRNA expression analysis by Deep-SAGE revealed increased IL-6 expression in RCC- and RAEB(t)-MDS. RCC-MDS MSC expressed increased levels of DKK3, a protein associated with decreased apoptosis. RAEB(t)-MDS revealed increased CRLF1 and decreased DAPK1 expressions. This pattern has been associated with transformation in hematopoietic malignancies. Genes reported to be differentially expressed in adult MDS-MSC did not differ between MSC of pediatric MDS and controls. An altered mRNA expression profile, associated with cell survival and malignant transformation, of MSC derived from children with MDS strengthens the hypothesis that the micro-environment is of importance in this disease. Our data support the understanding that pediatric and adult MDS are two different diseases. Further evaluation of the pathways involved might reveal additional therapy targets.

Abstract

An aberrant interaction between hematopoietic stem cells and mesenchymal stromal cells has been linked to disease and shown to contribute to the pathophysiology of hematologic malignancies in murine models. Juvenile myelomonocytic leukemia is an aggressive malignant disease affecting young infants. Here we investigated the impact of juvenile myelomonocytic leukemia on mesenchymal stromal cells. Mesenchymal stromal cells were expanded from bone marrow samples of patients at diagnosis (n=9) and after hematopoietic stem cell transplantation (n=7; from 5 patients) and from healthy children (n=10). Cells were characterized by phenotyping, differentiation, gene expression analysis (of controls and samples obtained at diagnosis) and in vitro functional studies assessing immunomodulation and hematopoietic support. Mesenchymal stromal cells from patients did not differ from controls in differentiation capacity nor did they differ in their capacity to support in vitro hematopoiesis. Deep-SAGE sequencing revealed differential mRNA expression in patient-derived samples, including genes encoding proteins involved in immunomodulation and cell-cell interaction. Selected gene expression normalized during remission after successful hematopoietic stem cell transplantation. Whereas natural killer cell activation and peripheral blood mononuclear cell proliferation were not differentially affected, the suppressive effect on monocyte to dendritic cell differentiation was increased by mesenchymal stromal cells obtained at diagnosis, but not at time of remission. This study shows that active juvenile myelomonocytic leukemia affects the immune response-related gene expression and function of mesenchymal stromal cells. In contrast, the differential gene expression of hematopoiesis-related genes could not be supported by functional data. Decreased immune surveillance might contribute to the therapy resistance and progression in juvenile myelomonocytic leukemia.

Abstract

Modulations of occipito-parietal α-band (8–14 Hz) power that are opposite in direction (α-enhancement vs. α-suppression) and origin of generation (ipsilateral vs. contralateral to the locus of attention) are a robust correlate of anticipatory visuospatial attention. Yet, the neural generators of these α-band modulations, their interdependence across homotopic areas, and their respective contribution to subsequent perception remain unclear. To shed light on these questions, we employed magnetoencephalography, while human volunteers performed a spatially cued detection task. Replicating previous findings, we found α-power enhancement ipsilateral to the attended hemifield and contralateral α-suppression over occipitoparietal sensors. Source localization (beamforming) analysis showed that α-enhancement and suppression were generated in 2 distinct brain regions, located in the dorsal and ventral visual streams, respectively. Moreover, α-enhancement and suppression showed different dynamics and contribution to perception. In contrast to the initial and transient dorsal α-enhancement, α-suppression in ventro-lateral occipital cortex was sustained and influenced subsequent target detection. This anticipatory biasing of ventrolateral extrastriate α-activity probably reflects increased receptivity in the brain region specialized in processing upcoming target features. Our results add to current models on the role of α-oscillations in attention orienting by showing that α-enhancement and suppression can be dissociated in time, space, and perceptual relevance.

Abstract

Rapid response to danger holds an evolutionary advantage. In this positron emission tomography study, phobics were exposed to masked visual stimuli with timings that either allowed awareness or not of either phobic, fear-relevant (e.g., spiders to snake phobics), or neutral images. When the timing did not permit awareness, the amygdala responded to both phobic and fear-relevant stimuli. With time for more elaborate processing, phobic stimuli resulted in an addition of an affective processing network to the amygdala activity, whereas no activity was found in response to fear-relevant stimuli. Also, right prefrontal areas appeared deactivated, comparing aware phobic and fear-relevant conditions. Thus, a shift from top-down control to an affectively driven system optimized for speed was observed in phobic relative to fear-relevant aware processing.

Abstract

Word meaning processing in the brain involves ventrolateral temporal cortex, but a semantic contribution of the dorsal stream, especially frontocentral sensorimotor areas, has been controversial. We here examine brain activation during passive reading of object-related nouns from different semantic categories, notably animal, food, and tool words, matched for a range of psycholinguistic features. Results show ventral stream activation in temporal cortex along with category-specific activation patterns in both ventral and dorsal streams, including sensorimotor systems and adjacent pFC. Precentral activation reflected action-related semantic features of the word categories. Cortical regions implicated in mouth and face movements were sparked by food words, and hand area activation was seen for tool words, consistent with the actions implicated by the objects the words are used to speak about. Furthermore, tool words specifically activated the right cerebellum, and food words activated the left orbito-frontal and fusiform areas. We discuss our results in the context of category-specific semantic deficits in the processing of words and concepts, along with previous neuroimaging research, and conclude that specific dorsal and ventral areas in frontocentral and temporal cortex index visual and affective–emotional semantic attributes of object-related nouns and action-related affordances of their referent objects.

Abstract

Although the typological contrast between Romance and Germanic languages as verb-framed versus satellite-framed (Talmy 1985) forms the background for many empirical studies on L2 acquisition, the inconclusive picture to date calls for more differentiated, fine-grained analyses. The present study goes beyond explanations based on this typological contrast and takes into account the sources from which spatial concepts are mainly derived in order to shape the trajectory traced by the entity in motion when moving through space: the entity in V-languages versus features of the ground in S-languages. It investigates why advanced French learners of English and German have difficulty acquiring the use of spatial concepts typical of the L2s to shape the trajectory, although relevant concepts can be expressed in their L1. The analysis compares motion event descriptions, based on the same sets of video clips, of L1 speakers of the three languages to L1 French-L2 English and L1 French-L2 German speakers, showing that the learners do not fully acquire the use of L2-specific spatial concepts. We argue that encoded concepts derived from the entity in motion vs. the ground lead to a focus on different aspects of motion events, in accordance with their compatibility with these sources, and are difficult to restructure in L2 acquisition.

Abstract

Can children’s handedness influence how they represent abstract concepts like kindness and intelligence? Here we show that from an early age, right-handers associate rightward space more strongly with positive ideas and leftward space with negative ideas, but the opposite is true for left-handers. In one experiment, children indicated where on a diagram a preferred toy and a dispreferred toy should go. Right-handers tended to assign the preferred toy to a box on the right and the dispreferred toy to a box on the left. Left-handers showed the opposite pattern. In a second experiment, children judged which of two cartoon animals looked smarter (or dumber) or nicer (or meaner). Right-handers attributed more positive qualities to animals on the right, but left-handers to animals on the left. These contrasting associations between space and valence cannot be explained by exposure to language or cultural conventions, which consistently link right with good. Rather, right- and left-handers implicitly associated positive valence more strongly with the side of space on which they can act more fluently with their dominant hands. Results support the body-specificity hypothesis (Casasanto, 2009), showing that children with different kinds of bodies think differently in corresponding ways.

Abstract

Learning a specific skill during childhood may partly determine the functional organization of the adult brain. This hypothesis led us to study oral language processing in illiterate subjects who, for social reasons, had never entered school and had no knowledge of reading or writing. In a brain activation study using PET and statistical parametric mapping, we compared word and pseudoword repetition in literate and illiterate subjects. Our study confirms behavioural evidence of different phonological processing in illiterate subjects. During repetition of real words, the two groups performed similarly and activated similar areas of the brain. In contrast, illiterate subjects had more difficulty repeating pseudowords correctly and did not activate the same neural structures as literates. These results are consistent with the hypothesis that learning the written form of language (orthography) interacts with the function of oral language. Our results indicate that learning to read and write during childhood influences the functional organization of the adult human brain.

Abstract

Last year was the 150th anniversary of Paul Broca's landmark case report on speech disorder that paved the way for subsequent studies of cortical localization of higher cognitive functions. However, many complex functions rely on the activity of distributed networks rather than single cortical areas. Hence, it is important to understand how brain regions are linked within large-scale networks and to map lesions onto connecting white matter tracts. To facilitate this network approach we provide a synopsis of classical neurological syndromes associated with frontal, parietal, occipital, temporal and limbic lesions. A review of tractography studies in a variety of neuropsychiatric disorders is also included. The synopsis is accompanied by a new atlas of the human white matter connections based on diffusion tensor tractography freely downloadable on http://www.natbrainlab.com. Clinicians can use the maps to accurately identify the tract affected by lesions visible on conventional CT or MRI. The atlas will also assist researchers to interpret their group analysis results. We hope that the synopsis and the atlas by allowing a precise localization of white matter lesions and associated symptoms will facilitate future work on the functional correlates of human neural networks as derived from the study of clinical populations. Our goal is to stimulate clinicians to develop a critical approach to clinico-anatomical correlative studies and broaden their view of clinical anatomy beyond the cortical surface in order to encompass the dysfunction related to connecting pathways.

Abstract

Specific language impairment (SLI), an unexpected failure to develop appropriate language skills despite adequate non-verbal intelligence, is a heterogeneous multifactorial disorder with a complex genetic basis. We identified a homozygous microdeletion of 21,379 bp in the ZNF277 gene (NM_021994.2), encompassing exon 5, in an individual with severe receptive and expressive language impairment. The microdeletion was not found in the proband’s affected sister or her brother who had mild language impairment. However, it was inherited from both parents, each of whom carries a heterozygous microdeletion and has a history of language problems. The microdeletion falls within the AUTS1 locus, a region linked to autistic spectrum disorders (ASDs). Moreover, ZNF277 is adjacent to the DOCK4 and IMMP2L genes, which have been implicated in ASD. We screened for the presence of ZNF277 microdeletions in cohorts of children with SLI or ASD and panels of control subjects. ZNF277 microdeletions were at an increased allelic frequency in SLI probands (1.1%) compared with both ASD family members (0.3%) and independent controls (0.4%). We performed quantitative RT-PCR analyses of the expression of IMMP2L, DOCK4 and ZNF277 in individuals carrying either an IMMP2L_DOCK4 microdeletion or a ZNF277 microdeletion. Although ZNF277 microdeletions reduce the expression of ZNF277, they do not alter the levels of DOCK4 or IMMP2L transcripts. Conversely, IMMP2L_DOCK4 microdeletions do not affect the expression levels of ZNF277. We postulate that ZNF277 microdeletions may contribute to the risk of language impairments in a manner that is independent of the autism risk loci previously described in this region.

Abstract

Lexicalized theories of syntax often assume that verb-structure regularities are mediated by lemmas, which abstract over variation in verb tense and aspect. German syntax seems to challenge this assumption, because verb position depends on tense and aspect. To examine how German speakers link these elements, a structural priming study was performed which varied syntactic structure, verb position (encoded by tense and aspect), and verb overlap. Abstract structural priming was found, both within and across verb position, but priming was larger when the verb position was the same between prime and target. Priming was boosted by verb overlap, but there was no interaction with verb position. The results can be explained by a lemma model where tense and aspect are linked to structural choices in German. Since the architecture of this lemma model is not consistent with results from English, a connectionist model was developed which could explain the cross-linguistic variation in the production system. Together, these findings support the view that language learning plays an important role in determining the nature of structural priming in different languages

Abstract

Linguistic adaptation is a phenomenon where language representations change in response to linguistic input. Adaptation can occur on multiple linguistic levels such as phonology (tuning of phonotactic constraints), words (repetition priming), and syntax (structural priming). The persistent nature of these adaptations suggests that they may be a form of implicit learning and connectionist models have been developed which instantiate this hypothesis. Research on implicit learning, however, has also produced evidence that explicit chunk knowledge is involved in the performance of these tasks. In this review, we examine how these interacting implicit and explicit processes may change our understanding of language learning and processing.

Abstract

Messenger ribonucleic acids (RNAs) contain a large number of cis-regulatory RNA elements that function in many types of post-transcriptional regulation. These cis-regulatory elements are often characterized by conserved structures and/or sequences. Although some classes are well known, given the wide range of RNA-interacting proteins in eukaryotes, it is likely that many new classes of cis-regulatory elements are yet to be discovered. An approach to this is to use computational methods that have the advantage of analysing genomic data, particularly comparative data on a large scale. In this study, a set of structural discovery algorithms was applied followed by support vector machine (SVM) classification. We trained a new classification model (CisRNA-SVM) on a set of known structured cis-regulatory elements from 3′-untranslated regions (UTRs) and successfully distinguished these and groups of cis-regulatory elements not been strained on from control genomic and shuffled sequences. The new method outperformed previous methods in classification of cis-regulatory RNA elements. This model was then used to predict new elements from cross-species conserved regions of human 3′-UTRs. Clustering of these elements identified new classes of potential cis-regulatory elements. The model, training and testing sets and novel human predictions are available at: http://mRNA.otago.ac.nz/CisRNA-SVM.

Abstract

This study examines the perception of paralinguistic intonational meanings deriving from Ohala’s Frequency Code (Experiment 1) and Gussenhoven’s Effort Code (Experiment 2) in British English and Dutch. Native speakers of British English and Dutch listened to a number of stimuli in their native language and judged each stimulus on four semantic scales deriving from these two codes: SELF-CONFIDENT versus NOT SELF-CONFIDENT, FRIENDLY versus NOT FRIENDLY (Frequency Code); SURPRISED versus NOT SURPRISED, and EMPHATIC versus NOT EMPHATIC (Effort Code). The stimuli, which were lexically equivalent across the two languages, differed in pitch contour, pitch register and pitch span in Experiment 1, and in pitch register, peak height, peak alignment and end pitch in Experiment 2. Contrary to the traditional view that the paralinguistic usage of intonation is similar across languages, it was found that British English and Dutch listeners differed considerably in the perception of “confident,” “friendly,” “emphatic,” and “surprised.” The present findings support a theory of paralinguistic meaning based on the universality of biological codes, which however acknowledges a languagespecific component in the implementation of these codes.

Abstract

While detecting genetic variations underlying brain structures helps reveal mechanisms of neural disorders, high data dimensionality poses a major challenge for imaging genomic association studies. In this work, we present the application of a recently proposed approach, parallel independent component analysis with reference (pICA-R), to investigate genomic factors potentially regulating gray matter variation in a healthy population. This approach simultaneously assesses many variables for an aggregate effect and helps to elicit particular features in the data. We applied pICA-R to analyze gray matter density (GMD) images (274,131 voxels) in conjunction with single nucleotide polymorphism (SNP) data (666,019 markers) collected from 1,256 healthy individuals of the Brain Imaging Genetics (BIG) study. Guided by a genetic reference derived from the gene GNA14, pICA-R identified a significant SNP-GMD association (r = −0.16, P = 2.34 × 10−8), implying that subjects with specific genotypes have lower localized GMD. The identified components were then projected to an independent dataset from the Mind Clinical Imaging Consortium (MCIC) including 89 healthy individuals, and the obtained loadings again yielded a significant SNP-GMD association (r = −0.25, P = 0.02). The imaging component reflected GMD variations in frontal, precuneus, and cingulate regions. The SNP component was enriched in genes with neuronal functions, including synaptic plasticity, axon guidance, molecular signal transduction via PKA and CREB, highlighting the GRM1, PRKCH, GNA12, and CAMK2B genes. Collectively, our findings suggest that GNA12 and GNA14 play a key role in the genetic architecture underlying normal GMD variation in frontal and parietal regions

Abstract

The goal of this study is to examine how the degree of vowel-to-vowel coarticulation varies as a function of prosodic factors such as nuclear-pitch accent (accented vs. unaccented), level of prosodic boundary (Prosodic Word vs. Intermediate Phrase vs. Intonational Phrase), and position-in-prosodic-domain (initial vs. final). It is hypothesized that vowels in prosodically stronger locations (e.g., in accented syllables and at a higher prosodic boundary) are not only coarticulated less with their neighboring vowels, but they also exert a stronger influence on their neighbors. Measurements of tongue position for English /a i/ over time were obtained with Carsten’s electromagnetic articulography. Results showed that vowels in prosodically stronger locations are coarticulated less with neighboring vowels, but do not exert a stronger influence on the articulation of neighboring vowels. An examination of the relationship between coarticulation and duration revealed that (a) accent-induced coarticulatory variation cannot be attributed to a duration factor and (b) some of the data with respect to boundary effects may be accounted for by the duration factor. This suggests that to the extent that prosodically conditioned coarticulatory variation is duration-independent, there is no absolute causal relationship from duration to coarticulation. It is proposed that prosodically conditioned V-to-V coarticulatory reduction is another type of strengthening that occurs in prosodically strong locations. The prosodically driven coarticulatory patterning is taken to be part of the phonetic signatures of the hierarchically nested structure of prosody.

Abstract

This study examines acoustic and aerodynamic characteristics of consonants in standard Korean and in Cheju, an endangered Korean language. The focus is on the well-known three-way distinction among voiceless stops (i.e., lenis, fortis, aspirated) and the two-way distinction between the voiceless fricatives /s/ and /s*/. While such a typologically unusual contrast among voiceless stops has long drawn the attention of phoneticians and phonologists, there is no single work in the literature that discusses a body of data representing a relatively large number of speakers. This study reports a variety of acoustic and aerodynamic measures obtained from 12 Korean speakers (four speakers of Seoul Korean and eight speakers of Cheju). Results show that, in addition to findings similar to those reported by others, there are three crucial points worth noting. Firstly, lenis, fortis, and aspirated stops are systematically differentiated from each other by the voice quality of the following vowel. Secondly, these stops are also differentiated by aerodynamic mechanisms. The aspirated and fortis stops are similar in supralaryngeal articulation, but employ a different relation between intraoral pressure and flow. Thirdly, our study suggests that the fricative /s/ is better categorized as “lenis” rather than “aspirated”. The paper concludes with a discussion of the implications of Korean data for theories of the voicing contrast and their phonological representations.

Abstract

Models of speech production assume that syllables play a functional role in the process of word-form encoding in speech production. In this study, we investigate this claim and specifically provide evidence about the level at which syllables come into play. We report two studies using an odd-man-out variant of the implicit priming paradigm to examine the role of the syllable during the process of word formation. Our results show that this modified version of the implicit priming paradigm can trace the emergence of syllabic structure during spoken word generation. Comparing these results to prior syllable priming studies, we conclude that syllables emerge at the interface between phonological and phonetic encoding. The results are discussed in terms of the WEAVER++ model of lexical access.

Abstract

The present study concerns individual differences in gesture production. We used correlational and multiple regression analyses to examine the relationship between individuals’ cognitive abilities and empathy levels and their gesture frequency and saliency. We chose predictor variables according to experimental evidence of the functions of gesture in speech production and communication. We examined 3 types of gestures: representational gestures, conduit gestures, and palm-revealing gestures. Higher frequency of representational gestures was related to poorer visual and spatial working memory, spatial transformation ability, and conceptualization ability; higher frequency of conduit gestures was related to poorer visual working memory, conceptualization ability, and higher levels of empathy; and higher frequency of palm-revealing gestures was related to higher levels of empathy. The saliency of all gestures was positively related to level of empathy. These results demonstrate that cognitive abilities and empathy levels are related to individual differences in gesture frequency and saliency

Abstract

Language and action systems are highly interlinked. A critical piece of evidence is that speech and its accompanying gestures are tightly synchronized. Five experiments were conducted to test 2 hypotheses about the synchronization of speech and gesture. According to the interactive view, there is continuous information exchange between the gesture and speech systems, during both their planning and execution phases. According to the ballistic view, information exchange occurs only during the planning phases of gesture and speech, but the 2 systems become independent once their execution has been initiated. In all experiments, participants were required to point to and/or name a light that had just lit up. Virtual reality and motion tracking technologies were used to disrupt their gesture or speech execution. Participants delayed their speech onset when their gesture was disrupted. They did so even when their gesture was disrupted at its late phase and even when they received only the kinesthetic feedback of their gesture. Also, participants prolonged their gestures when their speech was disrupted. These findings support the interactive view and add new constraints on models of speech and gesture production

Abstract

Koriat (1981) demonstrated that an association from the target to a preceding prime, in the absence of an association from the prime to the target, facilitates lexical decision and referred to this effect as "backward priming". Backward priming is of relevance, because it can provide information about the mechanism underlying semantic priming effects. Following Neely (1991), we distinguish three mechanisms of priming: spreading activation, expectancy, and semantic matching/integration. The goal was to determine which of these mechanisms causes backward priming, by assessing effects of backward priming on a language-relevant ERP component, the N400, and reaction time (RT). Based on previous work, we propose that the N400 priming effect reflects expectancy and semantic matching/integration, but in contrast with RT does not reflect spreading activation. Experiment 1 shows a backward priming effect that is qualitatively similar for the N400 and RT in a lexical decision task. This effect was not modulated by an ISI manipulation. Experiment 2 clarifies that the N400 backward priming effect reflects genuine changes in N400 amplitude and cannot be ascribed to other factors. We will argue that these backward priming effects cannot be due to expectancy but are best accounted for in terms of semantic matching/integration.

Abstract

Recent game-theoretic simulation and analytical models have demonstrated that cooperative strategies mediated by indicators of cooperative potential, or “tags,” can invade, spread, and resist invasion by noncooperators across a range of population-structure and cost-benefit scenarios. The plausibility of these models is potentially relevant for human evolutionary accounts insofar as humans possess some phenotypic trait that could serve as a reliable tag. Linguistic markers, such as accent and dialect, have frequently been either cursorily defended or promptly dismissed as satisfying the criteria of a reliable and evolutionarily viable tag. This paper integrates evidence from a range of disciplines to develop and assess the claim that speech accent mediated the evolution of tag-based cooperation in humans. Existing evidence warrants the preliminary conclusion that accent markers meet the demands of an evolutionarily viable tag and potentially afforded a cost-effective solution to the challenges of maintaining viable cooperative relationships in diffuse, regional social networks.

Abstract

Common wisdom has it that Buddhism enhances compassion and self-other integration. We put this assumption to empirical test by comparing practicing Taiwanese Buddhists with well-matched atheists. Buddhists showed more evidence of self-other integration in the social Simon task, which assesses the degree to which people co-represent the actions of a coactor. This suggests that self-other integration and task co-representation vary as a function of religious practice.

Abstract

Four cross-modal priming experiments and two forced-choice identification experiments investigated the use of suprasegmental cues to stress in the recognition of spoken English words, by native (English-speaking) and non- native (Dutch) listeners. Previous results had indicated that suprasegmental information was exploited in lexical access by Dutch but not by English listeners. For both listener groups, recognition of visually presented target words was faster, in comparison to a control condition, after stress-matching spoken primes, either monosyllabic (mus- from MUsic /muSEum) or bisyl labic (admi- from ADmiral/admiRAtion). For native listeners, the effect of stress-mismatching bisyllabic primes was not different from that of control primes, but mismatching monosyllabic primes produced partial facilitation. For non-native listeners, both bisyllabic and monosyllabic stress-mismatching primes produced partial facilitation. Native English listeners thus can exploit suprasegmental information in spoken-word recognition, but information from two syllables is used more effectively than information from one syllable. Dutch listeners are less proficient at using suprasegmental information in English than in their native language, but, as in their native language, use mono- and bisyllabic information to an equal extent. In forced-choice identification, Dutch listeners outperformed native listeners at correctly assigning a monosyllabic fragment (e.g., mus-) to one of two words differing in stress.

Abstract

Contemporary methods of computational cognitive modeling have recently been criticized by Addyman and French (2012) on the grounds that they have not kept up with developments in computer technology and human–computer interaction. They present a manifesto for change according to which, it is argued, modelers should devote more effort to making their models accessible, both to non-modelers (with an appropriate easy-to-use user interface) and modelers alike. We agree that models, like data, should be freely available according to the normal standards of science, but caution against confusing implementations with specifications. Models may embody theories, but they generally also include implementation assumptions. Cognitive modeling methodology needs to be sensitive to this. We argue that specification, replication and experimentation are methodological approaches that can address this issue.

Abstract

Speech processing can often take place in adverse listening conditions that involve the mixing of speech and background noise. In this study, we investigated processing dependencies between background noise and indexical speech features, using a speeded classification paradigm (Garner, 1974; Exp. 1), and whether background noise is encoded and represented in memory for spoken words in a continuous recognition memory paradigm (Exp. 2). Whether or not the noise spectrally overlapped with the speech signal was also manipulated. The results of Experiment 1 indicated that background noise and indexical features of speech (gender, talker identity) cannot be completely segregated during processing, even when the two auditory streams are spectrally nonoverlapping. Perceptual interference was asymmetric, whereby irrelevant indexical feature variation in the speech signal slowed noise classification to a greater extent than irrelevant noise variation slowed speech classification. This asymmetry may stem from the fact that speech features have greater functional relevance to listeners, and are thus more difficult to selectively ignore than background noise. Experiment 2 revealed that a recognition cost for words embedded in different types of background noise on the first and second occurrences only emerged when the noise and the speech signal were spectrally overlapping. Together, these data suggest integral processing of speech and background noise, modulated by the level of processing and the spectral separation of the speech and noise.

Abstract

In three experiments, listeners detected vowel or consonant targets in lists of CV syllables constructed from five vowels and five consonants. Responses were faster in a predictable context (e.g., listening for a vowel target in a list of syllables all beginning with the same consonant) than in an unpredictable context (e.g., listening for a vowel target in a list of syllables beginning with different consonants). In Experiment 1, the listeners’ native language was Dutch, in which vowel and consonant repertoires are similar in size. The difference between predictable and unpredictable contexts was comparable for vowel and consonant targets. In Experiments 2 and 3, the listeners’ native language was Spanish, which has four times as many consonants as vowels; here effects of an unpredictable consonant context on vowel detection were significantly greater than effects of an unpredictable vowel context on consonant detection. This finding suggests that listeners’ processing of phonemes takes into account the constitution of their language’s phonemic repertoire and the implications that this has for contextual variability.

Abstract

Single nucleotide polymorphisms (SNPs) within the MIR137, TCF4, and ZNF804A genes show genome-wide association to schizophrenia. However, the biological basis for the associations is unknown. Here, we tested the effects of these genes on brain structure in 1300 healthy adults. Using volumetry and voxel-based morphometry, neither gene-wide effects—including the combined effect of the genes—nor single SNP effects—including specific psychosis risk SNPs—were found on total brain volume, grey matter, white matter, or hippocampal volume. These results suggest that the associations between these risk genes and schizophrenia are unlikely to be mediated via effects on macroscopic brain structure.

Abstract

In most of the world, people have regular exposure to multiple accents. Therefore, learning to quickly process accented speech is a prerequisite to successful communication. In this paper, we examine work on the perception of accented speech across the lifespan, from early infancy to late adulthood. Unfamiliar accents initially impair linguistic processing by infants, children, younger adults, and older adults, but listeners of all ages come to adapt to accented speech. Emergent research also goes beyond these perceptual abilities, by assessing links with production and the relative contributions of linguistic knowledge and general cognitive skills. We conclude by underlining points of convergence across ages, and the gaps left to face in future work.

Abstract

The present study investigated the neural correlates of infant discrimination of very similar linguistic varieties (Quebecois and Parisian French) using functional Near InfraRed Spectroscopy. In line with previous behavioral and electrophysiological data, there was no evidence that 3-month-olds discriminated the two regional accents, whereas 5-month-olds did, with the locus of discrimination in left anterior perisylvian regions. These neuroimaging results suggest that a developing language network relying crucially on left perisylvian cortices sustains infants' discrimination of similar linguistic varieties within this early period of infancy.

Abstract

There are increasing reports that individual variation in behavioral and neurophysiological measures of infant speech processing predicts later language outcomes, and specifically concurrent or subsequent vocabulary size. If such findings are held up under scrutiny, they could both illuminate theoretical models of language development and contribute to the prediction of communicative disorders. A qualitative, systematic review of this emergent literature illustrated the variety of approaches that have been used and highlighted some conceptual problems regarding the measurements. A quantitative analysis of the same data established that the bivariate relation was significant, with correlations of similar strength to those found for well-established nonlinguistic predictors of language. Further exploration of infant speech perception predictors, particularly from a methodological perspective, is recommended.

Abstract

Typically, the point vowels [i,ɑ,u] are acoustically more peripheral in infant-directed speech (IDS) compared to adult-directed speech (ADS). If caregivers seek to highlight lexically relevant contrasts in IDS, then two sounds that are contrastive should become more distinct, whereas two sounds that are surface realizations of the same underlying sound category should not. To test this prediction, vowels that are phonemically contrastive ([i-ɪ] and [eɪ-ε]), vowels that map onto the same underlying category ([æ- ] and [ε- ]), and the point vowels [i,ɑ,u] were elicited in IDS and ADS by American English mothers of two age groups of infants (four- and eleven-month-olds). As in other work, point vowels were produced in more peripheral positions in IDS compared to ADS. However, there was little evidence of hyperarticulation per se (e.g. [i-ɪ] was hypoarticulated). We suggest that across-the-board lexically based hyperarticulation is not a necessary feature of IDS.

Abstract

Social tolerance is a core aspect of primate social relationships with implications for the evolution of cooperation, prosociality and social learning. We measured the social tolerance of bonobos in an experiment recently validated with chimpanzees to allow for a comparative assessment of group-level tolerance, and found that the bonobo group studied here exhibited lower social tolerance on average than chimpanzees. Furthermore, following the Relational Model [de Waal, 1996], we investigated whether bonobos responded to an increased potential for social conflict with tolerance, conflict avoidance or conflict escalation, and found that only behaviours indicative of conflict escalation differed across conditions. Taken together, these findings contribute to the current debate over the level of social tolerance of bonobos and lend support to the position that the social tolerance of bonobos may not be notably high compared with other primates.

Abstract

Studies of animal behavior consistently demonstrate that the social environment impacts cooperation, yet the effect of social dynamics has been largely excluded from studies of human cooperation. Here, we introduce a novel approach inspired by nonhuman primate research to address how social hierarchies impact human cooperation. Participants competed to earn hierarchy positions and then could cooperate with another individual in the hierarchy by investing in a common effort. Cooperation was achieved if the combined investments exceeded a threshold, and the higher ranked individual distributed the spoils unless control was contested by the partner. Compared to a condition lacking hierarchy, cooperation declined in the presence of a hierarchy due to a decrease in investment by lower ranked individuals. Furthermore, hierarchy was detrimental to cooperation regardless of whether it was earned or arbitrary. These findings mirror results from nonhuman primates and demonstrate that hierarchies are detrimental to cooperation. However, these results deviate from nonhuman primate findings by demonstrating that human behavior is responsive to changing hierarchical structures and suggests partnership dynamics that may improve cooperation. This work introduces a controlled way to investigate the social influences on human behavior, and demonstrates the evolutionary continuity of human behavior with other primate species.

Abstract

In the wild, chimpanzees (Pan troglodytes) are often faced with clumped food resources that they may know how to access but abstain from doing so due to social pressures. To better understand how social settings influence resource acquisition, we tested fifteen semi-wild chimpanzees from two social groups alone and in the presence of others. We investigated how resource acquisition was affected by relative social dominance, whether collaborative problem solving or (active or passive) sharing occurred amongst any of the dyads, and whether these outcomes were related to relationship quality as determined from six months of observational data. Results indicated that chimpanzees, regardless of rank, obtained fewer rewards when tested in the presence of others compared to when they were tested alone. Chimpanzees demonstrated behavioral inhibition; chimpanzees who showed proficient skill when alone often abstained from solving the task when in the presence of others. Finally, individuals with close social relationships spent more time together in the problem solving space, but collaboration and sharing were infrequent and sessions in which collaboration or sharing did occur contained more instances of aggression. Group living provides benefits and imposes costs, and these findings highlight that one cost of group living may be diminishing productive individual behaviors.

Abstract

Researchers have struggled to obtain a clear account of the evolution of prosocial behaviour despite a great deal of recent effort. The aim of this review is to take a brief step back from addressing the question of evolutionary origins of prosocial behaviour in order to identify contextual factors that are contributing to variation in the expression of prosocial behaviour and hindering progress towards identifying phylogenetic patterns. Most available data come from the Primate Order, and the choice of contextual factors to consider was informed by theory and practice, including the nature of the relationship between the potential donor and recipient, the communicative behaviour of the recipients, and features of the prosocial task including whether rewards are visible and whether the prosocial choice creates an inequity between actors. Conclusions are drawn about the facilitating or inhibiting impact of each of these factors on the expression of prosocial behaviour, and areas for future research are highlighted. Acknowledging the impact of these contextual features on the expression of prosocial behaviours should stimulate new research into the proximate mechanisms that drive these effects, yield experimental designs that better control for potential influences on prosocial expression, and ultimately allow progress towards reconstructing the evolutionary origins of prosocial behaviour.

Abstract

The social factors that influence cooperation have remained largely uninvestigated but have the potential to explain much of the variation in cooperative behavior observed in the natural world. We show here that certain dimensions of the social environment, namely the size of the social group, the degree of social tolerance expressed, the structure of the dominance hierarchy, and the patterns of dispersal, may influence the emergence and stability of cooperation in predictable ways. Furthermore, the social environment experienced by a species over evolutionary time will have shaped their cognition to provide certain strengths and strategies that are beneficial in their species‟ social world. These cognitive adaptations will in turn impact the likelihood of cooperating in a given social environment. Experiments with one primate species, the cottontop tamarin, illustrate how social dynamics may influence emergence and stability of cooperative behavior in this species. We then take a more general viewpoint and argue that the hypotheses presented here require further experimental work and the addition of quantitative modeling to obtain a better understanding of how social dynamics influence the emergence and stability of cooperative behavior in complex systems. We conclude by pointing out subsequent specific directions for models and experiments that will allow relevant advances in the understanding of the emergence of cooperation.

Abstract

Recent debates have questioned the extent to which culturally-transmitted norms drive behavioral variation in resource sharing across human populations. We shed new light on this discussion by examining the group-level variation in the social dynamics and resource sharing of chimpanzees, a species that is highly social and forms long-term community associations but differs from humans in the extent to which cultural norms are adopted and enforced. We rely on theory developed in primate socioecology to guide our investigation in four neighboring chimpanzee groups at a sanctuary in Zambia. We used a combination of experimental and observational approaches to assess the distribution of resource holding potential in each group. In the first assessment, we measured the proportion of the population that gathered in a resource-rich zone, in the second we assessed naturally occurring social spacing via social network analysis, and in the third we assessed the degree to which benefits were equally distributed within the group. We report significant, stable group-level variation across these multiple measures, indicating that group-level variation in resource sharing and social tolerance is not necessarily reliant upon human-like cultural norms.

Abstract

Rhythmic categories such as morae in Japanese or stress units in English play a role in the perception of spoken
language. We examined this role in Japanese, since recent evidence suggests that morae may intervene as
structural units in word recognition. First, we found that traditional puns more often substituted part of a mora
than a whole mora. Second, when listeners reconstructed distorted words, e.g. panorama from panozema, responses
were faster and more accurate when only a phoneme was distorted (panozama, panorema) than when a
whole CV mora was distorted (panozema). Third, lexical decisions on the same nonwords were better predicted
by duration and number of phonemes from nonword uniqueness point to word end than by number of morae. Our
results indicate no role for morae in early spoken-word processing; we propose that rhythmic categories constrain
not initial lexical activation but subsequent processes of speech segmentation and selection among word candidates.

Abstract

Native American English and non-native(Dutch)listeners identified either the consonant or the vowel in all possible American English CV and VC syllables. The syllables were embedded in multispeaker babble at three signal-to-noise ratios(0, 8, and 16 dB). The phoneme identification
performance of the non-native listeners was less accurate than that of the native listeners. All listeners were adversely affected by noise. With these isolated syllables, initial segments were harder to identify than final segments. Crucially, the effects of language background and noise did not interact; the performance asymmetry between the native and non-native groups was not significantly different across signal-to-noise ratios. It is concluded that the frequently reported disproportionate difficulty of non-native listening under disadvantageous conditions is not due to a disproportionate increase in phoneme misidentifications.

Abstract

Recognizing spoken language involves automatic activation of multiple candidate words. The process of selection between candidates is made more efficient by inhibition of embedded words (like egg in beg) that leave a portion of the input stranded (here, b). Results from European languages suggest that this inhibition occurs when consonants are stranded but not when syllables are stranded. The reason why leftover syllables do not lead to inhibition could be that in principle they might themselves be words; in European languages, a syllable can be a word. In Sesotho (a Bantu language), however, a single syllable cannot be a word. We report that in Sesotho, word recognition is inhibited by stranded consonants, but stranded monosyllables produce no more difficulty than stranded bisyllables (which could be Sesotho words). This finding suggests that the viability constraint which inhibits spurious embedded word candidates is not sensitive to language-specific word structure, but is universal.

Abstract

Languages differ in the constitution of their phonemic repertoire and in the relative distinctiveness of phonemes within the repertoire. In the present study, we asked whether such differences constrain spoken-word recognition, via two word reconstruction experiments, in which listeners turned non-words into real words by changing single sounds. The experiments were carried out in Dutch (which has a relatively balanced vowel-consonant ratio and many similar vowels) and in Spanish (which has many more consonants than vowels and high distinctiveness among the vowels). Both Dutch and Spanish listeners responded significantly faster and more accurately when required to change vowels as opposed to consonants; when allowed to change any phoneme, they more often altered vowels than consonants. Vowel information thus appears to constrain lexical selection less tightly (allow more potential candidates) than does consonant information, independent of language-specific phoneme repertoire and of relative distinctiveness of vowels.

Abstract

Spraak is continu, er zijn geen betrouwbare signalen waardoor de luisteraar weet waar het ene woord eindigt en het volgende begint. Voor volwassen luisteraars is het segmenteren van gesproken taal in afzonderlijke woorden dus niet onproblematisch, maar voor een kind dat nog geen woordenschat bezit, vormt de continuïteit van spraak een nog grotere uitdaging. Desalniettemin produceren de meeste kinderen hun eerste herkenbare woorden rond het begin van het tweede levensjaar. Aan deze vroege spraakproducties gaat een formidabele perceptuele prestatie vooraf. Tijdens het eerste levensjaar - met name gedurende de tweede helft - ontwikkelt de spraakperceptie zich van een algemeen fonetisch discriminatievermogen tot een selectieve gevoeligheid voor de fonologische contrasten die in de moedertaal voorkomen. Recent onderzoek heeft verder aangetoond dat kinderen, lang voordat ze ook maar een enkel woord kunnen zeggen, in staat zijn woorden die kenmerkend zijn voor hun moedertaal te onderscheiden van woorden die dat niet zijn. Bovendien kunnen ze woorden die eerst in isolatie werden aangeboden herkennen in een continue spraakcontext. Het dagelijkse taalaanbod aan een kind van deze leeftijd maakt het in zekere zin niet gemakkelijk, bijvoorbeeld doordat de meeste woorden niet in isolatie voorkomen. Toch wordt het kind ook wel houvast geboden, onder andere doordat het woordgebruik beperkt is.

Abstract

To examine whether lexically stored knowledge about spelling influences phoneme evaluation, we conducted three experiments with a low-level phonetic judgement task: phoneme goodness rating. In each experiment, listeners heard phonetic tokens varying along a continuum centred on /s/, occurring finally in isolated word or nonword tokens. An effect of spelling appeared in Experiment 1: Native English speakers’ goodness ratings for the best /s/ tokens were significantly higher in words spelled with S (e.g., bless) than in words spelled with C (e.g., voice). No such difference appeared when nonnative speakers rated the same materials in Experiment 2, indicating that the difference could not be due to acoustic characteristics of the S- versus C-words. In Experiment 3, nonwords with lexical neighbours consistently spelled with S (e.g., pless) versus with C (e.g., floice) failed to elicit orthographic neighbourhood effects; no significant difference appeared in native English speakers’ ratings for the S-consistent versus the C-consistent sets. Obligatory influence of lexical knowledge on phonemic processing would have predicted such neighbourhood effects; the findings are thus better accommodated by models in which phonemic decisions draw strategically upon lexical information.