Publications

Abstract

To identify genetic variants associated with birth weight, we meta-analyzed six genome-wide association (GWA) studies (n = 10,623 Europeans from pregnancy/birth cohorts) and followed up two lead signals in 13 replication studies (n = 27,591). rs900400 near LEKR1 and CCNL1 (P = 2 x 10(-35)) and rs9883204 in ADCY5 (P = 7 x 10(-15)) were robustly associated with birth weight. Correlated SNPs in ADCY5 were recently implicated in regulation of glucose levels and susceptibility to type 2 diabetes, providing evidence that the well-described association between lower birth weight and subsequent type 2 diabetes has a genetic component, distinct from the proposed role of programming by maternal nutrition. Using data from both SNPs, we found that the 9% of Europeans carrying four birth weight-lowering alleles were, on average, 113 g (95% CI 89-137 g) lighter at birth than the 24% with zero or one alleles (P(trend) = 7 x 10(-30)). The impact on birth weight is similar to that of a mother smoking 4-5 cigarettes per day in the third trimester of pregnancy.

Abstract

Disruptions of the FOXP2 gene cause a rare speech and language disorder, a discovery that has opened up novel avenues for investigating the relevant neural pathways. FOXP2 shows remarkably high conservation of sequence and neural expression in diverse vertebrates, suggesting that studies in other species are useful in elucidating its functions. Here we describe how investigations of mice that carry disruptions of Foxp2 provide insights at multiple levels: molecules, cells, circuits and behaviour. Work thus far has implicated the gene in key processes including neurite outgrowth, synaptic plasticity, sensorimotor integration and motor-skill learning.

Abstract

Cumulative tool-based culture underwrote our species' evolutionary success and tool-based nut-cracking is one of the strongest candidates for cultural transmission in our closest relatives, chimpanzees. However the social learning processes that may explain both the similarities and differences between the species remain unclear. A previous study of nut-cracking by initially naïve chimpanzees suggested that a learning chimpanzee holding no hammer nevertheless replicated hammering actions it witnessed. This observation has potentially important implications for the nature of the social learning processes and underlying motor coding involved. In the present study, model and observer actions were quantified frame-by-frame and analysed with stringent statistical methods, demonstrating synchrony between the observer's and model's movements, cross-correlation of these movements above chance level and a unidirectional transmission process from model to observer. These results provide the first quantitative evidence for motor mimicking underlain by motor coding in apes, with implications for mirror neuron function.

Abstract

Previous research on language development shows that children are tuned early on to the language-specific semantic and syntactic encoding of events in their native language. Here we ask whether language-specificity is also evident in children's early representations in gesture accompanying speech. In a longitudinal study, we examined the spontaneous speech and cospeech gestures of eight Turkish-speaking children aged one to three and focused on their caused motion event expressions. In Turkish, unlike in English, the main semantic elements of caused motion such as Action and Path can be encoded in the verb (e.g. sok- ‘put in’) and the arguments of a verb can be easily omitted. We found that Turkish-speaking children's speech indeed displayed these language-specific features and focused on verbs to encode caused motion. More interestingly, we found that their early gestures also manifested specificity. Children used iconic cospeech gestures (from 19 months onwards) as often as pointing gestures and represented semantic elements such as Action with Figure and/or Path that reinforced or supplemented speech in language-specific ways until the age of three. In the light of previous reports on the scarcity of iconic gestures in English-speaking children's early productions, we argue that the language children learn shapes gestures and how they get integrated with speech in the first three years of life.

Abstract

As the popularity of sending messages electronically increases, so does the necessity of conveying messages more efficiently. One way of increasing efficiency is to abbreviate words and expressions by combining letters with numbers such as gr8 for “great,” using acronyms, such as lol for “laughing out loud,” or clippings such as msg for “message.” The present study compares the processing of shortcuts to the processing of closely matched pseudo-shortcuts. ERPs were recorded while participants were performing a lexical decision task. Response times showed that shortcuts were categorized more slowly as nonwords than pseudo-shortcuts. The ERP results showed no differences between shortcuts and pseudo-shortcuts at time windows 50–150 ms and 150–270 ms, but there were significant differences between 270 and 500 ms. These results suggest that at early stages of word recognition, the orthographic and phonological processing is similar for shortcuts and pseudo-shortcuts. However, at the time of lexical access, shortcuts diverge from pseudo-shortcuts, suggesting that shortcuts activate stored lexical representations.

Abstract

The current event-related brain potential study examined the processing of observed speech errors.
Participants were asked to detect errors in the speech of others while listening to the description of a visual
network. Networks consisted of colored drawings of objects connected by straight or curved lines. We
investigated the processing of two types of errors in the network descriptions, i.e., incorrect color and errors
in determiners usage (gender agreement violations). In the 100- to 300-ms and 300- to 550-ms time
windows, we found larger PMN and N400 amplitudes for both color and determiner error trials compared to
correct trials. Furthermore, color but not determiner errors led to larger P600 amplitudes compared to
correct color trials. Color errors also showed enhanced P600 amplitudes compared to determiner errors.
Taken together, processing erroneous network descriptions elicits different brain potentials than listening to
the corresponding correct utterances. Hence, speech is monitored for errors not only during speech
production but also during listening to the naturally occurring speech of others.

Abstract

For efficiency reasons, words in electronic messages are sometimes formed by combining letters with numbers, as in gr8 for “great.” The aim of this study was to investigate whether a digit incorporated into a letter—digit shortcut would retain its numerosity. A priming paradigm was used with letter—digit shortcuts (e.g., gr8) and matched pseudoshortcuts (e.g., qr8) as primes. The primes were presented simultaneously with sets of dots (targets) for which even/odd decisions were required, or they appeared 250 msec before target onset. When pseudoshortcuts were presented, decision latencies were shorter when the target and the digit in the prime were matched in parity than when they were mismatched. This main effect of match was not significant for shortcuts. The results suggest that the number concepts of digits combined with letters become activated but are quickly suppressed or deactivated when the digit is part of an existing shortcut.

Abstract

This study investigated how sentence formulation is influenced by a preceding discourse context. In two eye-tracking experiments, participants described pictures of two-character transitive events in Dutch (Experiment 1) and Chinese (Experiment 2). Focus was manipulated by presenting questions before each picture. In the Neutral condition, participants first heard ‘What is happening here?’ In the Object or Subject Focus conditions, the questions asked about the Object or Subject character (What is the policeman stopping? Who is stopping the truck?). The target response was the same in all conditions (The policeman is stopping the truck). In both experiments, sentence formulation in the Neutral condition showed the expected pattern of speakers fixating the subject character (policeman) before the object character (truck). In contrast, in the focus conditions speakers rapidly directed their gaze preferentially only to the character they needed to encode to answer the question (the new, or focused, character). The timing of gaze shifts to the new character varied by language group (Dutch vs. Chinese): shifts to the new character occurred earlier when information in the question can be repeated in the response with the same syntactic structure (in Chinese but not in Dutch). The results show that discourse affects the timecourse of linguistic formulation in simple sentences and that these effects can be modulated by language-specific linguistic structures such as parallels in the syntax of questions and declarative sentences.

Abstract

Researchers have long avoided neurophysiological experiments of overt speech production due to the suspicion that artifacts caused by muscle activity may lead to a bad signal-to-noise ratio in the measurements. However, the need to actually produce speech may influence earlier processing and qualitatively change speech production processes and what we can infer from neurophysiological measures thereof. Recently, however, overt speech has been successfully investigated using EEG, MEG, and fMRI. The aim of this Research Topic is to draw together recent research on the neurophysiological basis of language production, with the aim of developing and extending theoretical accounts of the language production process. In this Research Topic of Frontiers in Language Sciences, we invite both experimental and review papers, as well as those about the latest methods in acquisition and analysis of overt language production data. All aspects of language production are welcome: i.e., from conceptualization to articulation during native as well as multilingual language production. Focus should be placed on using the neurophysiological data to inform questions about the processing stages of language production. In addition, emphasis should be placed on the extent to which the identified components of the electrophysiological signal (e.g., ERP/ERF, neuronal oscillations, etc.), brain areas or networks are related to language comprehension and other cognitive domains. By bringing together electrophysiological and neuroimaging evidence on language production mechanisms, a more complete picture of the locus of language production processes and their temporal and neurophysiological signatures will emerge.

Abstract

The constraints that govern acceptable phoneme combinations in speech perception and production have considerable plasticity. We addressed whether sleep influences the acquisition of new constraints and their integration into the speech-production system. Participants repeated sequences of syllables in which two phonemes were artificially restricted to syllable onset or syllable coda, depending on the vowel in that sequence. After 48 sequences, participants either had a 90-min nap or remained awake. Participants then repeated 96 sequences so implicit constraint learning could be examined, and then were tested for constraint generalization in a forced-choice task. The sleep group, but not the wake group, produced speech errors at test that were consistent with restrictions on the placement of phonemes in training. Furthermore, only the sleep group generalized their learning to new materials. Polysomnography data showed that implicit constraint learning was associated with slow-wave sleep. These results show that sleep facilitates the integration of new linguistic knowledge with existing production constraints. These data have relevance for systems-consolidation models of sleep.

Abstract

Heterozygous mutations of the human FOXP2 gene are implicated in a severe speech and language disorder. Aetiological mutations of murine Foxp2 yield abnormal synaptic plasticity and impaired motor-skill learning in mutant mice, while knockdown of the avian orthologue in songbirds interferes with auditory-guided vocal learning. Here, we investigate influences of two distinct Foxp2 point mutations on vocalizations of 4-day-old mouse pups (Mus musculus). The R552H missense mutation is identical to that causing speech and language deficits in a large well-studied human family, while the S321X nonsense mutation represents a null allele that does not produce Foxp2 protein. We ask whether vocalizations, based solely on innate mechanisms of production, are affected by these alternative Foxp2 mutations. Sound recordings were taken in two different situations: isolation and distress, eliciting a range of call types, including broadband vocalizations of varying noise content, ultrasonic whistles and clicks. Sound production rates and several acoustic parameters showed that, despite absence of functional Foxp2, homozygous mutants could vocalize all types of sounds in a normal temporal pattern, but only at comparably low intensities. We suggest that altered vocal output of these homozygotes may be secondary to developmental delays and somatic weakness. Heterozygous mutants did not differ from wild-types in any of the measures that we studied (R552H ) or in only a few (S321X ), which were in the range of differences routinely observed for different mouse strains. Thus, Foxp2 is not essential for the innate production of emotional vocalizations with largely normal acoustic properties by mouse pups.

Abstract

An endophenotype for attention deficit/hyperactivity disorder (AD/HD) is executive functioning. In the autism and developmental literature executive dysfunctions has also been linked to theory of mind (ToM) and pragmatic language use. The central question of this review is whether deficits in ToM and pragmatic language use are common in AD/HD. AD/HD seems to be associated with pragmatic deficits, but not with ToM deficits. In this review we address how this pattern of findings might facilitate the understanding of the commonalities and differences between executive functioning, ToM, and pragmatic abilities. Based on the reviewed studies we conclude that ToM is not likely to be a potential endophenotype for AD/HD, while it is too early to draw such a conclusion for pragmatic language use.

Abstract

Reading and language abilities are heritable traits that are likely to share some genetic influences with each other. To identify pleiotropic genetic variants affecting these traits, we first performed a Genome-wide Association Scan (GWAS) meta-analysis using three richly characterised datasets comprising individuals with histories of reading or language problems, and their siblings. GWAS was performed in a total of 1862 participants using the first principal component computed from several quantitative measures of reading- and language-related abilities, both before and after adjustment for performance IQ. We identified novel suggestive associations at the SNPs rs59197085 and rs5995177 (uncorrected p≈10−7 for each SNP), located respectively at the CCDC136/FLNC and RBFOX2 genes. Each of these SNPs then showed evidence for effects across multiple reading and language traits in univariate association testing against the individual traits. FLNC encodes a structural protein involved in cytoskeleton remodelling, while RBFOX2 is an important regulator of alternative splicing in neurons. The CCDC136/FLNC locus showed association with a comparable reading/language measure in an independent sample of 6434 participants from the general population, although involving distinct alleles of the associated SNP. Our datasets will form an important part of on-going international efforts to identify genes contributing to reading and language skills.

Abstract

BACKGROUND: It is unclear to what extent non-clinical psychotic experiences during childhood and adolescence share underlying aetiological mechanisms with schizophrenia. One candidate mechanism for schizophrenia involves the epigenetic status of the developing fetus, which depends on the internal folate-status of mother and child. Our study examines the relationships between multiple determinants of perinatal folate-status and development of psychotic experiences in adolescence. METHODS: Study participants were up to 5344 mother-child pairs from the Avon Longitudinal Study of Parents and their Children, UK, with information on maternal and/or child MTHFR C677T genotype, maternal folate intake (supplementation at 18/32- weeks gestation; dietary intake at 32- weeks gestation) and psychosis-like symptoms (PLIKS) for children assessed at age 12. RESULTS: Nominal evidence was observed that maternal folate supplementation at 18 weeks increased the odds of PLIKS in children (odds ratio(OR)=1.34; 95%-CI:[1.00;1.76]) and, consistent with this, that children of MTHFR C667T TT homozygous mothers had decreased odds of PLIKS (OR=0.72; 95%CI:[0.50;1.02]; recessive model) with strongest effects in boys (OR=0.44, 95%-CI:[0.22;0.79]; sex-specific p=0.029). None of the reported effects remained significant when corrected for multiple testing. CONCLUSIONS: Overall, this study found no support that maternal/child MTHFR C677T genotype and maternal folate intake during pregnancy contribute to common aetiological pathways that are shared between schizophrenia and non-clinical psychotic symptoms in adolescents, assuming that decreased folate-status increases schizophrenia risk.

Abstract

PURPOSE: Conduct problems and peer effects are among the strongest risk factors for adolescent substance use and problem use. However, it is unclear to what extent the effects of conduct problems and peer behavior interact, and whether adolescents' capacity to refuse the offer of substances may moderate such links. This study was conducted to examine relationships between conduct problems, close friends' substance use, and refusal assertiveness with adolescents' alcohol use problems, tobacco, and marijuana use. METHODS: We studied a population-based sample of 1,237 individuals from the Cardiff Study of All Wales and North West of England Twins aged 11-18 years. Adolescent and mother-reported information was obtained. Statistical analyses included cross-sectional and prospective logistic regression models and family-based permutations. RESULTS: Conduct problems and close friends' substance use were associated with increased adolescents' substance use, whereas refusal assertiveness was associated with lower use of cigarettes, alcohol, and marijuana. Peer substance use moderated the relationship between conduct problems and alcohol use problems, such that conduct problems were only related to increased risk for alcohol use problems in the presence of substance-using friends. This effect was found in both cross-sectional and prospective analyses and confirmed using the permutation approach. CONCLUSIONS: Reduced opportunities for interaction with alcohol-using peers may lower the risk of alcohol use problems in adolescents with conduct problems.

Abstract

Previous studies have described the existence of a phonotactic bias called the Labial–Coronal (LC) bias, corresponding to a tendency to produce more words beginning with a labial consonant followed by a coronal consonant (i.e. “bat”) than the opposite CL pattern (i.e. “tap”). This bias has initially been interpreted in terms of articulatory constraints of the human speech production system. However, more recently, it has been suggested that this presumably language-general LC bias in production might be accompanied by LC and CL biases in perception, acquired in infancy on the basis of the properties of the linguistic input. The present study investigates the origins of these perceptual biases, testing infants learning Japanese, a language that has been claimed to possess more CL than LC sequences, and comparing them with infants learning French, a language showing a clear LC bias in its lexicon. First, a corpus analysis of Japanese IDS and ADS revealed the existence of an overall LC bias, except for plosive sequences in ADS, which show a CL bias across counts. Second, speech preference experiments showed a perceptual preference for CL over LC plosive sequences (all recorded by a Japanese speaker) in 13- but not in 7- and 10-month-old Japanese-learning infants (Experiment 1), while revealing the emergence of an LC preference between 7 and 10 months in French-learning infants, using the exact same stimuli. These crosslinguistic behavioral differences, obtained with the same stimuli, thus reflect differences in processing in two populations of infants, which can be linked to differences in the properties of the lexicons of their respective native languages. These findings establish that the emergence of a CL/LC bias is related to exposure to a linguistic input.

Abstract

Humans appear to rely on spatial mappings to represent and describe concepts. The conceptual cuing effect describes the tendency for participants to orient attention to a spatial location following the presentation of an unrelated cue word (e.g., orienting attention upward after reading the word sky). To date, such effects have predominately been explained within the embodied cognition framework, according to which people’s attention is oriented on the basis of prior experience (e.g., sky → up via perceptual simulation). However, this does not provide a compelling explanation for how abstract words have the same ability to orient attention. Why, for example, does dream also orient attention upward? We report on an experiment that investigated the role of language use (specifically, collocation between concept words and spatial words for up and down dimensions) and found that it predicted the cuing effect. The results suggest that language usage patterns may be instrumental in explaining conceptual cuing.

Abstract

Our recent studies suggest that congenitally blind adults have severely impaired thresholds in an auditory spatial bisection task, pointing to the importance of vision in constructing complex auditory spatial maps (Gon etal., 2014). To explore strategies that may improve the auditory spatial sense in visually impaired people, we investigated the impact of tactile feedback on spatial auditory localization in 48 blindfolded sighted subjects. We measured auditory spatial bisection thresholds before and after training, either with tactile feedback, verbal feedback, or no feedback. Audio thresholds were first measured with a spatial bisection task: subjects judged whether the second sound of a three sound sequence was spatially closer to the first or the third sound. The tactile feedback group underwent two audio-tactile feedback sessions of 100 trials, where each auditory trial was followed by the same spatial sequence played on the subject's forearm; auditory spatial bisection thresholds were evaluated after each session. In the verbal feedback condition, the positions of the sounds were verbally reported to the subject after each feedback trial.The no feedback group did the same sequence of trials, with no feedback. Performance improved significantly only after audio-tactile feedback. The results suggest that direct tactile feedback interacts with the auditory spatial localization system, possibly by a process of cross-sensory recalibration. Control tests with the subject rotated suggested that this effect occurs only when the tactile and acoustic sequences are spatially congruent. Our results suggest that the tactile system can be used to recalibrate the auditory sense of space. These results encourage the possibility of designing rehabilitation programs to help blind persons establish a robust auditory sense of space, through training with the tactile modality.

Abstract

The involvement of neural motor and sensory systems in the processing of language has so far mainly been studied in native (L1) speakers. In an fMRI experiment, we investigated whether non-native (L2) semantic representations are rich enough to allow for activation in motor and somatosensory brain areas. German learners of Dutch and a control group of Dutch native speakers made lexical decisions about visually presented Dutch motor and non-motor verbs. Region-of-interest (ROI) and whole-brain analyses indicated that L2 speakers, like L1 speakers, showed significantly increased activation for simple motor compared to non-motor verbs in motor and somatosensory regions. This effect was not restricted to Dutch-German cognate verbs, but was also present for non-cognate verbs. These results indicate that L2 semantic representations are rich enough for motor-related activations to develop in motor and somatosensory areas.

Abstract

In this functional magnetic resonance imaging (fMRI) long-lag priming study, we investigated the processing of Dutch semantically transparent, derived prefix verbs. In such words, the meaning of the word as a whole can be deduced from the meanings of its parts, e.g., wegleggen “put aside.” Many behavioral and some fMRI studies suggest that native (L1) speakers decompose transparent derived words. The brain region usually implicated in morphological decomposition is the left inferior frontal gyrus (LIFG). In non-native (L2) speakers, the processing of transparent derived words has hardly been investigated, especially in fMRI studies, and results are contradictory: some studies find more reliance on holistic (i.e., non-decompositional) processing by L2 speakers; some find no difference between L1 and L2 speakers. In this study, we wanted to find out whether Dutch transparent derived prefix verbs are decomposed or processed holistically by German L2 speakers of Dutch. Half of the derived verbs (e.g., omvallen “fall down”) were preceded by their stem (e.g., vallen “fall”) with a lag of 4–6 words (“primed”); the other half (e.g., inslapen “fall asleep”) were not (“unprimed”). L1 and L2 speakers of Dutch made lexical decisions on these visually presented verbs. Both region of interest analyses and whole-brain analyses showed that there was a significant repetition suppression effect for primed compared to unprimed derived verbs in the LIFG. This was true both for the analyses over L2 speakers only and for the analyses over the two language groups together. The latter did not reveal any interaction with language group (L1 vs. L2) in the LIFG. Thus, L2 speakers show a clear priming effect in the LIFG, an area that has been associated with morphological decomposition. Our findings are consistent with the idea that L2 speakers engage in decomposition of transparent derived verbs rather than processing them holistically

Abstract

Language in high-functioning autism is characterized by pragmatic and semantic deficits, and people with autism have a reduced tendency to integrate information. Because the left and right inferior frontal (LIF and RIF) regions are implicated with integration of speaker information, world knowledge, and semantic knowledge, we hypothesized that abnormal functioning of the LIF and RIF regions might contribute to pragmatic and semantic language deficits in autism. Brain activation of sixteen 12- to 18-year-old, high-functioning autistic participants was measured with functional magnetic resonance imaging during sentence comprehension and compared with that of twenty-six matched controls. The content of the pragmatic sentence was congruent or incongruent with respect to the speaker characteristics (male/female, child/adult, and upper class/lower class). The semantic- and world-knowledge sentences were congruent or incongruent with respect to semantic expectancies and factual expectancies about the world, respectively. In the semanticknowledge and world-knowledge condition, activation of the LIF region did not differ between groups. In sentences that required integration of speaker information, the autism group showed abnormally reduced activation of the LIF region. The results suggest that people with autism may recruit the LIF region in a different manner in tasks that demand integration of social information.

Abstract

The left and right sides of the human brain are specialized for different kinds of information processing, and much of our cognition is lateralized to an extent towards one side or the other. Handedness is a reflection of nervous system lateralization. Roughly ten percent of people are mixed- or left-handed, and they show an elevated rate of reductions or reversals of some cerebral functional asymmetries compared to right-handers. Brain anatomical correlates of left-handedness have also been suggested. However, the relationships of left-handedness to brain structure and function remain far from clear. We carried out a comprehensive analysis of cortical surface area differences between 106 left-handed subjects and 1960 right-handed subjects, measured using an automated method of regional parcellation (FreeSurfer, Destrieux atlas). This is the largest study sample that has so far been used in relation to this issue. No individual cortical region showed an association with left-handedness that survived statistical correction for multiple testing, although there was a nominally significant association with the surface area of a previously implicated region: the left precentral sulcus. Identifying brain structural correlates of handedness may prove useful for genetic studies of cerebral asymmetries, as well as providing new avenues for the study of relations between handedness, cerebral lateralization and cognition.

Abstract

Functional and anatomical asymmetries are prevalent features of the human brain, linked to gender, handedness, and cognition. However, little is known about the neurodevelopmental processes involved. In zebrafish, asymmetries arise in the diencephalon before extending within the central nervous system. We aimed to identify genes involved in the development of subtle, left-right volumetric asymmetries of human subcortical structures using large datasets. We first tested the feasibility of measuring left-right volume differences in such large-scale samples, as assessed by two automated methods of subcortical segmentation (FSL|FIRST and FreeSurfer), using data from 235 subjects who had undergone MRI twice. We tested the agreement between the first and second scan, and the agreement between the segmentation methods, for measures of bilateral volumes of six subcortical structures and the hippocampus, and their volumetric asymmetries. We also tested whether there were biases introduced by left-right differences in the regional atlases used by the methods, by analyzing left-right flipped images. While many bilateral volumes were measured well (scan-rescan r = 0.6-0.8), most asymmetries, with the exception of the caudate nucleus, showed lower repeatabilites. We meta-analyzed genome-wide association scan results for caudate nucleus asymmetry in a combined sample of 3,028 adult subjects but did not detect associations at genome-wide significance (P < 5 × 10-8). There was no enrichment of genetic association in genes involved in left-right patterning of the viscera. Our results provide important information for researchers who are currently aiming to carry out large-scale genome-wide studies of subcortical and hippocampal volumes, and their asymmetries

Abstract

A large body of evidence has shown that visual context information can rapidly modulate language comprehension for concrete sentences and when it is mediated by a referential or a lexical-semantic link. What has not yet been examined is whether visual context can also modulate comprehension of abstract sentences incrementally when it is neither referenced by, nor lexically associated with, the sentence. Three eye-tracking reading experiments examined the effects of spatial distance between words (Experiment 1) and objects (Experiment 2 and 3) on participants’ reading times for sentences that convey similarity or difference between two abstract nouns (e.g., ‘Peace and war are certainly different...’). Before reading the sentence, participants inspected a visual context with two playing cards that moved either far apart or close together. In Experiment 1, the cards turned and showed the first two nouns of the sentence (e.g., ‘peace’, ‘war’). In Experiments 2 and 3, they turned but remained blank. Participants’ reading times at the adjective (Experiment 1: first-pass reading time; Experiment 2: total times) and at the second noun phrase (Experiment 3: first-pass times) were faster for sentences that expressed similarity when the preceding words/objects were close together (vs. far apart) and for sentences that expressed dissimilarity when the preceding words/objects were far apart (vs. close together). Thus, spatial distance between words or entirely unrelated objects can rapidly and incrementally modulate the semantic interpretation of abstract sentences.

Abstract

Despite the literature on the role of input in adult second-language (L2) acquisition and on artificial and statistical language learning, surprisingly little is known about how adults break into a new language in the wild. This article reports on a series of behavioral and neuroimaging studies that examine what linguistic information adults can extract from naturalistic but controlled audiovisual input in an unknown and typologically distant L2 after minimal exposure (7–14 min) without instruction or training. We tested the stepwise development of segmental, phonotactic, and lexical knowledge in Dutch adults after minimal exposure to Mandarin Chinese and the role of item frequency, speech-associated gestures, and word length at the earliest stages of learning. In an exploratory neural connectivity study we further examined the neural correlates of word recognition in a new language, identifying brain regions whose connectivity was related to performance both before and after learning. While emphasizing the complexity of the learning task, the results suggest that the adult learning mechanism is more powerful than is normally assumed when faced with small amounts of complex, continuous audiovisual language input.

Abstract

All accounts of communicative behaviour in general, and communicative strategies in particular, mention gesture1 in relation to language acquisition (cf. Faerch & Kasper 1983 for an overview). However, few attempts have been made to investigate how spoken language and spontaneous gesture combine to determine discourse referents. Referential gesture and referential discourse will be of particular interest, since communicative strategies in second language discourse often involve labelling problems.

This paper will focus on two issues:

1) Within a cognitive account of communicative strategies, gesture will be seen to be part of conceptual or analysis-based strategies, in that relational features in the referents are exploited;

2) It will be argued that communication strategies can be seen in terms of cue manipulation in the same sense as sentence processing has been analysed in terms of competing cues. Strategic behaviour, and indeed the process of referring in general, are seen in terms of cues, combining or competing to determine discourse referents. Gesture can then be regarded as being such a cue at the discourse level, and as a cue-based communicative strategy, in that gesture functions by exploiting physically based cues which can be recognised as being part of the referent. The question of iconicity and motivation vs. the arbitrary qualities of gesture as a strategic cue will be addressed in connection with this.

Abstract

Gestures, the symbolic movements speakers perform while they speak, form a closely inter-connected system with speech where gestures serve both addressee-directed (‘communicative’) and speaker-directed (’internal’) functions. This paper aims (1) to show that a combined analysis of gesture and speech offers new ways to address theoretical issues in SLA and bilingualism studies, probing SLA and bilingualism as product and process; and (2) to outline some methodological concerns and desiderata to facilitate the inclusion of gesture in SLA and bilingualism research.

Abstract

Placement verbs describe every-day events like putting a toy in a box. Dutch uses two semi-obligatory caused posture verbs (leggen ‘lay’ and zetten ‘set/stand’) to distinguish between events based on whether the located object is placed horizontally or vertically. Although prevalent in the input, these verbs cause Dutch children difficulties even at age five (Narasimhan & Gullberg, submitted). Children overextend leggen to all placement events and underextend use of zetten. This study examines what gestures can reveal about Dutch three- and five-year-olds’ semantic representations of such verbs. The results show that children gesture differently from adults in this domain. Three-year-olds express only the path of the caused motion, whereas five-year-olds, like adults, also incorporate the located object. Crucially, gesture patterns are tied to verb use: those children who over-use leggen 'lay' for all placement events only gesture about path. Conversely, children who use the two verbs differentially for horizontal and vertical placement also incorporate objects in gestures like adults. We argue that children's gestures reflect their current knowledge of verb semantics, and indicate a developmental transition from a system with a single semantic component – (caused) movement – to an (adult-like) focus on two semantic components – (caused) movement-and-object

Abstract

Lexical-semantic processing impairments in aphasic patients with left hemisphere lesions and non-aphasic patients with right hemisphere lesions were investigated by recording event-related brain potentials (ERPs) while subjects listened to auditorily presented word pairs. The word pairs consisted of unrelated words, or words that were related in meaning. The related words were either associatively related, e.g. 'bread-butter', or were members of the same semantic category without being associatively related, e.g. 'churchvilla '. The latter relationships are assumed to be more distant than the former ones. The most relevant ERP component in this study is the N400. In elderly control subjects, the N400 amplitude to associatively and semantically related word targets is reduced relative to the N400 elicited by unrelated targets. Compared with this normal N400 effect, the different patient groups showed the following pattern of results: aphasic patients with only minor comprehension deficits (high comprehenders) showed N400 effects of a similar size as the control subjects. In aphasic patients with more severe comprehension deficits (low comprehenders) a clear reduction in the N400 effects was obtained, both for the associative and the semantic word pairs. The patients with right hemisphere lesions showed a normal N400 effect for the associatively related targets, but a trend towards a reduced N400 effect for the semantically related word pairs. A dissociation between the N400 results in the word pair paradigm and P300 results in a classical tone oddball task indicated that the N400 effects were not an aspecific consequence of brain lesion, but were related to the nature of the language comprehension impairment. The conclusions drawn from the ERP results are that comprehension deficits in the aphasic patients are due to an impairment in integrating individual word meanings into an overall meaning representation. Right hemisphere patients are more specifically impaired in the processing of semantically more distant relationships, suggesting the involvement of the right hemisphere in semantically coarse coding.

Abstract

Current views on the neurobiological underpinnings of language are discussed that deviate in a number of ways from the classical Wernicke–Lichtheim–Geschwind model. More areas than Broca's and Wernicke's region are involved in language. Moreover, a division along the axis of language production and language comprehension does not seem to be warranted. Instead, for central aspects of language processing neural infrastructure is shared between production and comprehension. Three different accounts of the role of Broca's area in language are discussed. Arguments are presented in favor of a dynamic network view, in which the functionality of a region is co-determined by the network of regions in which it is embedded at particular moments in time. Finally, core regions of language processing need to interact with other networks (e.g. the attentional networks and the ToM network) to establish full functionality of language and communication.

Abstract

A hallmark of human language is that we combine lexical building blocks retrieved from memory in endless new ways. This combinatorial aspect of language is referred to as unification. Here we focus on the neurobiological infrastructure for syntactic and semantic unification. Unification is characterized by a high-speed temporal profile including both prediction and integration of retrieved lexical elements. A meta-analysis of numerous neuroimaging studies reveals a clear dorsal/ventral gradient in both left inferior frontal cortex and left posterior temporal cortex, with dorsal foci for syntactic processing and ventral foci for semantic processing. In addition to core areas for unification, further networks need to be recruited to realize language-driven communication to its full extent. One example is the theory of mind network, which allows listeners and readers to infer the intended message (speaker meaning) from the coded meaning of the linguistic utterance. This indicates that sensorimotor simulation cannot handle all of language processing.

Abstract

In the recent past the work on large-scale linguistic distributions across the globe has intensified considerably. Work on macro-areal relationships in Africa (Güldemann, 2010) suggests that the shape of convergence areas may be determined by climatic factors and geophysical features such as mountains, water bodies, coastlines, etc. Worldwide data is now available for geophysical features as well as linguistic features, including numeral systems and basic constituent order. We explore the possibility that the shape of areal aggregations of individual features in these two linguistic domains correlates with Köppen-Geiger climate zones. Furthermore, we test the hypothesis that the shape of such areal feature aggregations is determined by the contour of adjacent geophysical features like mountain ranges or coastlines. In these first basic tests, we do not find clear evidence that either Köppen-Geiger climate zones or the contours of geophysical features are major predictors for the linguistic data at hand

Abstract

While the notion of the ‘area’ or ‘Sprachbund’ has a long history in linguistics, with geographically-defined regions frequently cited as a useful means to explain typological distributions, the problem of delimiting areas has not been well addressed. Lists of general-purpose, largely independent ‘macro-areas’ (typically continent size) have been proposed as a step to rule out contact as an explanation for various large-scale linguistic phenomena. This squib points out some problems in some of the currently widely-used predetermined areas, those found in the World Atlas of Language Structures (Haspelmath et al., 2005). Instead, we propose a principled division of the world’s landmasses into six macro-areas that arguably have better geographical independence properties

Abstract

One attempt at explaining why some language families are large (while others are small) is the hypothesis that the families that are now large became large because their ancestral speakers had a technological advantage, most often agriculture. Variants of this idea are referred to as the Language Farming Dispersal Hypothesis. Previously, detailed language family studies have uncovered various supporting examples and counterexamples to this idea. In the present paper I weigh the evidence from ALL attested language families. For each family, I use the number of member languages as a measure of cardinal size, member language coordinates to measure geospatial size and ethnographic evidence to assess subsistence status. This data shows that, although agricultural families tend to be larger in cardinal size, their size is hardly due to the simple presence of farming. If farming were responsible for language family expansions, we would expect a greater east-west geospatial spread of large families than is actually observed. The data, however, is compatible with weaker versions of the farming dispersal hypothesis as well with models where large families acquire farming because of their size, rather than the other way around.

Abstract

This paper aims to list all known language families that are not yet extinct and all of whose member languages are very poorly documented, i.e., less than a sketch grammar’s worth of data has been collected. It explains what constitutes a valid family, what amount and kinds of documentary data are sufficient, when a language is considered extinct, and more. It is hoped that the survey will be useful in setting priorities for documentation fieldwork, in particular for those documentation efforts whose underlying goal is to understand linguistic diversity.

Abstract

Slovak (sometimes also called Slovakian) is an Indo-European language belonging to the West-Slavic branch, and is most closely related to Czech. Slovak is spoken as a native language by 4.6 million speakers in Slovakia (that is by roughly 85% of the population), and by over two million Slovaks living abroad, most of them in the USA, the Czech Republic, Hungary, Canada and Great Britain (Office for Slovaks Living Abroad 2009).

Abstract

The possible-word constraint (PWC; Norris, McQueen, Cutler, & Butterfield, 1997) has been proposed as a language-universal segmentation principle: Lexical candidates are disfavoured if the resulting segmentation of continuous speech leads to vowelless residues in the input—for example, single consonants. Three word-spotting experiments investigated segmentation in Slovak, a language with single-consonant words and fixed stress. In Experiment 1, Slovak listeners detected real words such as ruka “hand” embedded in prepositional-consonant contexts (e.g., /gruka/) faster than those in nonprepositional-consonant contexts (e.g., /truka/) and slowest in syllable contexts (e.g., /dugruka/). The second experiment controlled for effects of stress. Responses were still fastest in prepositional-consonant contexts, but were now slowest in nonprepositional-consonant contexts. In Experiment 3, the lexical and syllabic status of the contexts was manipulated. Responses were again slowest in nonprepositional-consonant contexts but equally fast in prepositional-consonant, prepositional-vowel, and nonprepositional-vowel contexts. These results suggest that Slovak listeners use fixed stress and the PWC to segment speech, but that single consonants that can be words have a special status in Slovak segmentation. Knowledge about what constitutes a phonologically acceptable word in a given language therefore determines whether vowelless stretches of speech are or are not treated as acceptable parts of the lexical parse.

Abstract

Drawings of objects were presented in series of 54 each to 14 German speaking subjects with the tasks to indicate by button presses a) whether the grammatical gender of an object name was masculine ("der") or feminine ("die") and b) whether the depicted object was man-made or nature-made. The magnetoencephalogram (MEG) was recorded with a whole-head neuromagnetometer and task-specific patterns of brain activity were determined in the source space (Minimum Norm Estimates, MNE). A left-temporal focus of activity 150-275 ms after stimulus onset in the gender decision compared to the semantic classification task was discussed as indicating the retrieval of syntactic information, while a more expanded left hemispheric activity in the gender relative to the semantic task 300-625 ms after stimulus onset was discussed as indicating phonological encoding. A predominance of activity in the semantic task was observed over right fronto-central region 150-225 ms after stimulus-onset, suggesting that semantic and syntactic processes are prominent in this stage of lexical selection.

Abstract

All primates learn things from conspecifics socially, but it is not clear whether they conform to the behavior of these conspecifics—if conformity is defined as overriding individually acquired behavioral tendencies in order to copy peers’ behavior. In the current study, chimpanzees, orangutans, and 2-year-old human children individually acquired a problem-solving strategy. They then watched several conspecific peers demonstrate an alternative strategy. The children switched to this new, socially demonstrated strategy in roughly half of all instances, whereas the other two great-ape species almost never adjusted their behavior to the majority’s. In a follow-up study, children switched much more when the peer demonstrators were still present than when they were absent, which suggests that their conformity arose at least in part from social motivations. These results demonstrate an important difference between the social learning of humans and great apes, a difference that might help to account for differences in human and nonhuman cultures

Abstract

Contemporary comparative cognition has a large repertoire of animal models and methods, with concurrent theoretical advances that are providing initial answers to crucial questions about human cognition. What cognitive traits are uniquely human? What are the species-typical inherited predispositions of the human mind? What is the human mind capable of without certain types of specific experiences with the surrounding environment? Here, we review recent findings from the domains of space, time and number cognition. These findings are produced using different comparative methodologies relying on different animal species, namely birds and non-human great apes. The study of these species not only reveals the range of cognitive abilities across vertebrates, but also increases our understanding of human cognition in crucial ways.

Abstract

We advance the hypothesis here that the higher-than-average vocal pitch (FO) found for speech of Broca's aphasics in experimental settings is due, in part, to increased psychological stress. Two experiments were conducted which manipulated conversational constraints and the sentence forms to be produced by aphasic patients. Our study revealed significant differences between changes in vocal pitch of agrammatic Broca's aphasics versus those of Wernicke's aphasics and normal controls. It is suggested that the greater psychological stress experienced by the Broca's aphasics, but not by the Wernicke's aphasics, accounts for these observed differences.

Abstract

OBJECTIVE: Plasma adiponectin is strongly associated with various components of metabolic syndrome, type 2 diabetes and cardiovascular outcomes. Concentrations are highly heritable and differ between men and women. We therefore aimed to investigate the genetics of plasma adiponectin in men and women. METHODS: We combined genome-wide association scans of three population-based studies including 4659 persons. For the replication stage in 13795 subjects, we selected the 20 top signals of the combined analysis, as well as the 10 top signals with p-values less than 1.0 x 10(-4) for each the men- and the women-specific analyses. We further selected 73 SNPs that were consistently associated with metabolic syndrome parameters in previous genome-wide association studies to check for their association with plasma adiponectin. RESULTS: The ADIPOQ locus showed genome-wide significant p-values in the combined (p=4.3 x 10(-24)) as well as in both women- and men-specific analyses (p=8.7 x 10(-17) and p=2.5 x 10(-11), respectively). None of the other 39 top signal SNPs showed evidence for association in the replication analysis. None of 73 SNPs from metabolic syndrome loci exhibited association with plasma adiponectin (p>0.01). CONCLUSIONS: We demonstrated the ADIPOQ gene as the only major gene for plasma adiponectin, which explains 6.7% of the phenotypic variance. We further found that neither this gene nor any of the metabolic syndrome loci explained the sex differences observed for plasma adiponectin. Larger studies are needed to identify more moderate genetic determinants of plasma adiponectin.

Abstract

This paper provides an overview of the question–response system of Danish, based on a collection of 350 questions (and responses) collected from video recordings of naturally occurring face-to-face interactions between native speakers of Danish. The paper identifies the lexico-grammatical options for formulating questions, the range of social actions that can be implemented through questions and the relationship between questions and responses. It further describes features where Danish questions differ from a range of other languages in terms of, for instance, distribution and the relationship between question format and social action. For instance, Danish has a high frequency of interrogatively formatted questions and questions that are negatively formulated, when compared to languages that have the same grammatical options. In terms of action, Danish shows a higher number of questions that are used for making suggestions, offers and requests and does not use repetition as a way of answering a question as often as other languages.

Abstract

Objective: This study investigates the relationship of ‘online commentary’(contemporaneous physician comments about physical examination [PE] findings) with (i) parent questioning of the treatment recommendation and (ii) inappropriate antibiotic prescribing. Methods: A nested cross-sectional study of 522 encounters motivated by upper respiratory symptoms in 27 California pediatric practices (38 pediatricians). Physicians completed a post-visit survey regarding physical examination findings, diagnosis, treatment, and whether they perceived the parent as expecting an antibiotic. Taped encounters were coded for ‘problem’ online commentary (PE findings discussed as significant or clearly abnormal) and ‘no problem’ online commentary (PE findings discussed reassuringly as normal or insignificant). Results: Online commentary during the PE occurred in 73% of visits with viral diagnoses (n = 261). Compared to similar cases with ‘no problem’ online commentary, ‘problem’ comments were associated with a 13% greater probability of parents uestioning a non-antibiotic treatment plan (95% CI 0-26%, p = .05,) and a 27% (95% CI: 2-52%, p < .05) greater probability of an inappropriate antibiotic prescription. Conclusion: With viral illnesses, problematic online comments are associated with more pediatrician-parent conflict over non-antibiotic treatment recommendations. This may increase inappropriate antibiotic prescribing. Practice implications: In viral cases, physicians should consider avoiding the use of problematic online commentary.

Abstract

Bilingual listeners comprehend speech-in-noise better in their native than non-native language. This native-language benefit is thought to arise from greater use of top-down linguistic information to assist degraded speech comprehension. Using functional magnetic resonance imaging, we recently showed that left angular gyrus activation is modulated when semantic context is used to assist native language speech-in-noise comprehension (Golestani, Hervais-Adelman, Obleser, & Scott, 2013). Here, we extend the previous work, by reanalyzing the previous data alongside the results obtained in the non-native language of the same late bilingual participants. We found a behavioral benefit of semantic context in processing speech-in-noise in the native language only, and the imaging results also revealed a native language context effect in the left angular gyrus. We also find a complementary role of lower-level auditory regions during stimulus-driven processing. Our findings help to elucidate the neural basis of the established native language behavioral benefit of speech-in-noise processing. (C) 2014 Elsevier Inc. All rights reserved.

Abstract

Superiority in visual search for individuals diagnosed with autism spectrum disorder (ASD) is a well-reported finding. We administered two visual search tasks to individuals with ASD and matched controls. One showed no difference between the groups, and one did show the expected superior performance for individuals with ASD. These results offer an explanation, formulated in terms of load theory. We suggest that there is a limit to the superiority in visual search for individuals with ASD, related to the perceptual load of the stimuli. When perceptual load becomes so high that no additional task-(ir)relevant information can be processed, performance will be based on single stimulus identification, in which no differences between individuals with ASD and controls have been demonstrated

Abstract

Theories of embodied language comprehension have proposed that language is understood through perceptual simulation of the sensorimotor characteristics of its meaning. Strong support for this claim requires demonstration of encoding-based activation of sensorimotor representations that is distinct from task-related or goal-driven processes. Participants in 3 eye-tracking experiments were presented with triplets of either numbers or object and animal names. In Experiment 1, participants indicated whether the size of the referent of the middle object or animal name was in between the size of the 2 outer items. In Experiment 2, the object and animal names were encoded for an immediate recognition memory task. In Experiment 3, participants completed the same comparison task of Experiment 1 for both words and numbers. During the comparison tasks, word and number decision times showed a symbolic distance effect, such that response time was inversely related to the size difference between the items. A symbolic distance effect was also observed for animal and object encoding times in cases where encoding time likely reflected some goal-driven processes as well. When semantic size was irrelevant to the task (Experiment 2), it had no effect on word encoding times. Number encoding times showed a numerical distance priming effect: Encoding time increased with numerical difference between items. Together these results suggest that while activation of numerical magnitude representations is encoding-based as well as goal-driven, activation of size information associated with words is goal-driven and does not occur automatically during encoding. This conclusion challenges strong theories of embodied cognition which claim that language comprehension consists of activation of analog sensorimotor representations irrespective of higher level processes related to context or task-specific goals

Abstract

Two eye-tracking experiments were conducted in which the manual response mode typically used in lexical decision tasks (LDTs) was replaced with an eye-movement response through a sequence of 3 words. This ocular LDT combines the explicit control of task goals found in LDTs with the highly practiced ocular response used in reading text. In Experiment 1, forward saccades indicated an affirmative lexical decision (LD) on each word in the triplet. In Experiment 2, LD responses were delayed until all 3 letter strings had been read. The goal of the study was to evaluate the contribution of task goals and response mode to semantic priming. Semantic priming is very robust in tasks that involve recognition of words in isolation, such as LDT, but limited during text reading, as measured using eye movements. Gaze durations in both experiments showed robust semantic priming even though ocular response times were much shorter than manual LDs for the same words in the English Lexicon Project. Ex-Gaussian distribution fits revealed that the priming effect was concentrated in estimates of tau (τ), meaning that priming was most pronounced in the slow tail of the distribution. This pattern shows differential use of the prime information, which may be more heavily recruited in cases in which the LD is difficult, as indicated by longer response times. Compared with the manual LD responses, ocular LDs provide a more sensitive measure of this task-related influence on word recognition as measured by the LDT.

Abstract

When a sentence such as The model embraced the designer and the photographer laughed is read, the noun phrase the photographer is temporarily ambiguous: It can be either one of the objects of embraced (NP-coordination) or the subject of a new, conjoined sentence (S-coordination). It has been shown for a number of languages, including Dutch (the language used in this study), that readers prefer NP-coordination over S-coordination, at least in isolated sentences. In the present paper, it will be suggested that NP-coordination is preferred because it is the simpler of the two options in terms of topic-structure; in NP-coordinations there is only one topic, whereas S-coordinations contain two. Results from off-line (sentence completion) and online studies (a self-paced reading and an eye tracking experiment) support this topic-structure explanation. The processing difficulty associated with S-coordinated sentences disappeared when these sentences followed contexts favoring a two-topic continuation. This finding establishes topic-structure as an important factor in online sentence processing.

Abstract

Participants in interaction routinely orient to gaze, bodily comportment, and nonlexical vocalizations as salient for developing an analysis of the unfolding course of action. In this article, I address the respiratory phenomenon of sighing, the aim being to describe sighing as a situated practice that contributes to the achievement of particular actions in interaction. I report on the various actions sighs implement or construct and how their positioning and delivery informs participants’ understandings of their significance for interaction. Data are in American English

Abstract

Background: Individuals with autism spectrum disorder (ASD) and their parents demonstrate impaired performance in rapid automatized naming (RAN), a task that recruits a variety of linguistic and executive processes. Though the basic processes that contribute to RAN differences remain unclear, eye-voice relationships, as measured through eye tracking, can provide insight into cognitive and perceptual processes contributing to RAN performance. For example, in RAN, eye-voice span (EVS), the distance ahead the eyes are when articulation of a target item's label begins, is an indirect measure of automaticity of the processes underlying RAN. The primary objective of this study was to investigate automaticity in naming processes, as indexed by EVS during RAN. The secondary objective was to characterize RAN difficulties in individuals with ASD and their siblings. Methods: Participants (aged 15 – 33 years) included 21 individuals with ASD, 23 siblings of individuals with ASD, and 24 control subjects, group-matched on chronological age. Naming time, frequency of errors, and EVS were measured during a RAN task and compared across groups. Results: A stepwise pattern of RAN performance was observed, with individuals with ASD demonstrating the slowest naming across all RAN conditions, controls demonstrating the fastest naming, and siblings demonstrating intermediate performance. Individuals with ASD exhibited smaller EVSs than controls on all RAN conditions, and siblings exhibited smaller EVSs during number naming (the most highly automatized type of naming). EVSs were correlated with naming times in controls only, and only in the more automatized conditions. Conclusions: These results suggest that reduced automaticity in the component processes of RAN may underpin differences in individuals with ASD and their siblings. These findings also provide further support that RAN abilities are impacted by genetic liability to ASD. This study has important implications for understanding the underlying skills contributing to language-related deficits in ASD.

Abstract

In human face-to-face communication, language comprehension is a multi-modal, situated activity. However, little is known about how we combine information from different modalities during comprehension, and how perceived communicative intentions, often signaled through visual signals, influence this process. We explored this question by simulating a multi-party communication context in which a speaker alternated her gaze between two recipients. Participants viewed speech-only or speech + gesture object-related messages when being addressed (direct gaze) or unaddressed (gaze averted to other participant). They were then asked to choose which of two object images matched the speaker’s preceding message. Unaddressed recipients responded significantly more slowly than addressees for speech-only utterances. However, perceiving the same speech accompanied by gestures sped unaddressed recipients up to a level identical to that of addressees. That is, when unaddressed recipients’ speech processing suffers, gestures can enhance the comprehension of a speaker’s message. We discuss our findings with respect to two hypotheses attempting to account for how social eye gaze may modulate multi-modal language comprehension.

Abstract

The FOXP2 transcription factor is one of the most well-known genes to have been implicated in developmental speech and language disorders. Rare mutations disrupting the function of this gene have been described in different families and cases. In a large three-generation family carrying a missense mutation, neuroimaging studies revealed significant effects on brain structure and function, most notably in the inferior frontal gyrus, caudate nucleus and cerebellum. After the identification of rare disruptive FOXP2 variants impacting on brain structure, several reports proposed that common variants at this locus may also have detectable effects on the brain, extending beyond disorder into normal phenotypic variation. These neuroimaging genetics studies used groups of between 14 and 96 participants. The current study assessed effects of common FOXP2 variants on neuroanatomy using voxel-based morphometry and volumetric techniques in a sample of >1300 people from the general population. In a first targeted stage we analyzed single nucleotide polymorphisms (SNPs) claimed to have effects in prior smaller studies (rs2253478, rs12533005, rs2396753, rs6980093, rs7784315, rs17137124, rs10230558, rs7782412, rs1456031), beginning with regions proposed in the relevant papers, then assessing impact across the entire brain. In the second gene-wide stage, we tested all common FOXP2 variation, focusing on volumetry of those regions most strongly implicated from analyses of rare disruptive mutations. Despite using a sample that is more than ten times that used for prior studies of common FOXP2 variation, we found no evidence for effects of SNPs on variability in neuroanatomy in the general population. Thus, the impact of this gene on brain structure may be largely limited to extreme cases of rare disruptive alleles. Alternatively, effects of common variants at this gene exist but are too subtle to be detected with standard volumetric techniques

Abstract

Very little research has attempted to describe normal human variation in female genitalia, and no studies have compared the visual images that women might use in constructing their ideas of average and acceptable genital morphology to see if there are any systematic differences. Our objective was to determine if visual depictions of the vulva differed according to their source so as to alert medical professionals and their patients to how these depictions might capture variation and thus influence perceptions of "normality". We conducted a comparative analysis by measuring (a) published visual materials from human anatomy textbooks in a university library, (b) feminist publications (both print and online) depicting vulval morphology, and (c) online pornography, focusing on the most visited and freely accessible sites in the UK. Post-hoc tests showed that labial protuberance was significantly less (p < .001, equivalent to approximately 7 mm) in images from online pornography compared to feminist publications. All five measures taken of vulval features were significantly correlated (p < .001) in the online pornography sample, indicating a less varied range of differences in organ proportions than the other sources where not all measures were correlated. Women and health professionals should be aware that specific sources of imagery may depict different types of genital morphology and may not accurately reflect true variation in the population, and consultations for genital surgeries should include discussion about the actual and perceived range of variation in female genital morphology.

Abstract

This paper examines ╪Ākhoe Hai||om, a Khoe language of the Khoisan family spoken in Northern Namibia. I document the way questions are posed in natural conversation, the actions the questions are used for and the manner in which they are responded to. I show that in this language speakers rely most heavily on content questions. I also find that speakers of ╪Ākhoe Hai||om address fewer questions to a specific individual than would be expected from prior research on Indo European languages. Finally, I discuss some possible explanations for these findings.

Abstract

In two eye-tracking studies we investigated the influence of Mandarin numeral classifiers - a grammatical category in the language - on online overt attention. Mandarin speakers were presented with simple sentences through headphones while their eye-movements to objects presented on a computer screen were monitored. The crucial question is what participants look at while listening to a pre-specified target noun. If classifier categories influence Mandarin speakers' general conceptual processing, then on hearing the target noun they should look at objects that are members of the same classifier category - even when the classifier is not explicitly present (cf. Huettig & Altmann, 2005). The data show that when participants heard a classifier (e.g., ba3, Experiment 1) they shifted overt attention significantly more to classifier-match objects (e.g., chair) than to distractor objects. But when the classifier was not explicitly presented in speech, overt attention to classifier-match objects and distractor objects did not differ (Experiment 2). This suggests that although classifier distinctions do influence eye-gaze behavior, they do so only during linguistic processing of that distinction and not in moment-to-moment general conceptual processing.

Abstract

At present, more than one-fifth of humanity is unable to read and write. We critically examine experimental evidence and theories of how (il)literacy affects the human mind. In our discussion we show that literacy has significant cognitive consequences that go beyond the processing of written words and sentences. Thus, cultural inventions such as reading shape general cognitive processing in non-trivial ways. We suggest that this has important implications for educational policy and guidance as well as research into cognitive processing and brain functioning.

Abstract

Theories of verbal self-monitoring generally assume an internal (pre-articulatory) monitoring channel, but there is debate about whether this channel relies on speech perception or on production-internal mechanisms. Perception-based theories predict that listening to one's own inner speech has similar behavioral consequences as listening to someone else's speech. Our experiment therefore registered eye-movements while speakers named objects accompanied by phonologically related or unrelated written words. The data showed that listening to one's own speech drives eye-movements to phonologically related words, just as listening to someone else's speech does in perception experiments. The time-course of these eye-movements was very similar to that in other-perception (starting 300 ms post-articulation), which demonstrates that these eye-movements were driven by the perception of overt speech, not inner speech. We conclude that external, but not internal monitoring, is based on speech perception.

Abstract

The acquisition and maintenance of new language information, such as picking up new words, is a critical human ability that is needed throughout the life span. Most likely you learned the word “blog” quite recently as an adult, whereas the word “kipe,” which in the 1970s denoted stealing, now seems unfamiliar. Brain mechanisms underlying the long-term maintenance of new words have remained unknown, albeit they could provide important clues to the considerable individual differences in the ability to remember words. After successful training of a set of novel object names we tracked, over a period of 10 months, the maintenance of this new vocabulary in 10 human participants by repeated behavioral tests and magnetoencephalography measurements of overt picture naming. When namingrelated activation in the left frontal and temporal cortex was enhanced 1 week after training, compared with the level at the end of training, the individual retained a good command of the new vocabulary at 10 months; vice versa, individuals with reduced activation at 1 week posttraining were less successful in recalling the names at 10 months. This finding suggests an individual neural marker for memory, in the context of language. Learning is not over when the acquisition phase has been successfully completed: neural events during the access to recently established word representations appear to be important for the long-term outcome of learning.

Abstract

The articles in this volume are the result of an invited conference entitled "The Earliest Stages of Language Learning" held at the Max Planck Institute for Psycholinguistics in Nijmegen, The Netherlands, in October 2009.

Abstract

Hickok's enterprise to unify psycholinguistic and motor control models is highly stimulating. Nonetheless, there are problems of the model with respect to the time course of neural activation in word production, the flexibility for continuous speech, and the need for non-motor feedback.

Abstract

The Abelson helper integration site 1 (AHI1) gene locus on chromosome 6q23 is among a group of candidate loci for schizophrenia susceptibility that were initially identified by linkage followed by linkage disequilibrium mapping, and subsequent replication of the association in an independent sample. Here, we present results of a replication study of AHI1 locus markers, previously implicated in schizophrenia, in a large European sample (in total 3907 affected and 7429 controls). Furthermore, we perform a meta-analysis of the implicated markers in 4496 affected and 18,920 controls. Both the replication study of new samples and the meta-analysis show evidence for significant overrepresentation of all tested alleles in patients compared with controls (meta-analysis; P = 8.2 x 10(-5)-1.7 x 10(-3), common OR = 1.09-1.11). The region contains two genes, AHI1 and C6orf217, and both genes-as well as the neighbouring phosphodiesterase 7B (PDE7B)-may be considered candidates for involvement in the genetic aetiology of schizophrenia.

Abstract

The results of a self-paced reading study with German second language (L2) learners of Dutch showed that noun animacy affected the learners' on-line commitments when comprehending relative clauses in their L2. Earlier research has found that German L2 learners of Dutch do not show an on-line preference for subject–object word order in temporarily ambiguous relative clauses when no disambiguating material is available prior to the auxiliary verb. We investigated whether manipulating the animacy of the ambiguous noun phrases would push the learners to make an on-line commitment to either a subject- or object-first analysis. Results showed they performed like Dutch native speakers in that their reading times reflected an interaction between topichood and animacy in the on-line assignment of grammatical roles

Abstract

Initial lexical activation in typical populations is a direct reflection of the goodness of fit between the presented stimulus and the intended target. In this study, lexical activation was investigated upon presentation of polysyllabic pseudowords (such as procodile for crocodile) for the atypical population of dyslexic adults to see to what extent mismatching phonemic information affects lexical activation in the face of overwhelming support for one specific lexical candidate. Results of an auditory lexical decision task showed that sensitivity to phonemic mismatch was less in the dyslexic population, compared to the respective control group. However, the dyslexic participants were outperformed by their controls only for word-initial mismatches. It is argued that a subtle speech decoding deficit affects lexical activation levels and makes spoken word processing less robust against distortion.

Abstract

Background: There is evidence that, unlike in typical populations, initial lexical activation upon hearing spoken words in aphasic patients is not a direct reflection of the goodness of fit between the presented stimulus and the intended target. Earlier studies have mainly used short monosyllabic target words. Short words are relatively difficult to recognise because they are not highly redundant: changing one phoneme will often result in a (similar-sounding) different word. Aims: The present study aimed to investigate sensitivity of the lexical recognition system in aphasia. The focus was on longer words that contain more redundancy, to investigate whether aphasic adults might be impaired in deactivation of strongly activated lexical candidates. This was done by studying lexical activation upon presentation of spoken polysyllabic pseudowords (such as procodile) to see to what extent mismatching phonemic information leads to deactivation in the face of overwhelming support for one specific lexical candidate. Methods & Procedures: Speeded auditory lexical decision was used to investigate response time and accuracy to pseudowords with a word-initial or word-final phonemic mismatch in 21 aphasic patients and in an age-matched control group. Outcomes & Results: Results of an auditory lexical decision task showed that aphasic participants were less sensitive to phonemic mismatch if there was strong evidence for one particular lexical candidate, compared to the control group. Classifications of patients as Broca's vs Wernicke's or as fluent vs non-fluent did not reveal differences in sensitivity to mismatch between aphasia types. There was no reliable relationship between measures of auditory verbal short-term memory and lexical decision performance. Conclusions: It is argued that the aphasic results can best be viewed as lexical “overactivation” and that a verbal short-term memory account is less appropriate.

Abstract

Many older listeners report difficulties in understanding speech in noisy situations. Working memory and other cognitive skills may modulate, however, older listeners’ ability to use context information to alleviate the effects of noise on spoken-word recognition. In the present study, we investigated whether working memory predicts older adults’ ability to immediately use context information in the recognition of words embedded in sentences, presented in different listening conditions. In a phoneme-monitoring task, older adults were asked to detect as fast and as accurately as possible target phonemes in sentences spoken by a target speaker. Target speech was presented without noise, with fluctuating speech-shaped noise, or with competing speech from a single distractor speaker. The gradient measure of contextual probability (derived from a separate offline rating study) mainly affected the speed of recognition, with only a marginal effect on detection accuracy. Contextual facilitation was modulated by older listeners’ working memory and age across listening conditions. Working memory and age, as well as hearing loss, were also the most consistent predictors of overall listening performance. Older listeners’ immediate benefit from context in spoken-word recognition thus relates to their ability to keep and update a semantic representation of the sentence content in working memory.

Abstract

The authors report six implicit priming experiments that examined the production of inflected forms. Participants produced words out of small sets in response to prompts. The words differed in form or shared word-initial segments, which allowed for preparation. In constant inflectional sets, the words had the same number of inflectional suffixes, whereas in variable sets the number of suffixes differed. In the experiments, preparation effects were obtained, which were larger in the constant than in the variable sets. Control experiments showed that this difference in effect was not due to syntactic class or phonological form per se. The results are interpreted in terms of a slot-and-filler model of word production, in which inflectional frames, on the one hand, and stems and affixes, on the other hand, are independently spelled out on the basis of an abstract morpho-syntactic specification of the word, which is followed by morpheme-to-frame association.

Abstract

Discrete phonological phenomena form our conscious experience of language: continuous changes in pitch appear as distinct tones to the speakers of tone languages, whereas the speakers of quantity languages experience duration categorically. The categorical nature of our linguistic experience is directly reflected in the traditionally clear-cut linguistic classification of languages into tonal or non-tonal. However, some evidence suggests that duration and pitch are fundamentally interconnected and co-vary in signaling word meaning in non-tonal languages as well. We show that pitch information affects real-time language processing in a (non-tonal) quantity language. The results suggest that there is no unidirectional causal link from a genetically-based perceptual sensitivity towards pitch information to the appearance of a tone language. They further suggest that the contrastive categories tone and quantity may be based on simultaneously co-varying properties of the speech signal and the processing system, even though the conscious experience of the speakers may highlight only one discrete variable at a time.

Abstract

In 7 experiments the authors investigated the locus of word frequency effects in speech production. Experiment 1 demonstrated a frequency effect in picture naming that was robust over repetitions. Experiments 2, 3, and 7 excluded contributions from object identification and initiation of articulation. Experiments 4 and 5 investigated whether the effect arises in accessing the syntactic word (lemma) by using a grammatical gender decision task. Although a frequency effect was found, it dissipated under repeated access to word's gender. Experiment 6 tested whether the robust frequency effect arises in accessing the phonological form (lexeme) by having Ss translate words that produced homophones. Low-frequent homophones behaved like high-frequent controls, inheriting the accessing speed of their high-frequent homophone twins. Because homophones share the lexeme, not the lemma, this suggests a lexeme-level origin of the robust effect.

Abstract

Visual cues to the individual segments of speech and to sentence prosody guide speech recognition. The present study tested whether visual suprasegmental cues to the stress patterns of words can also constrain recognition. Dutch listeners use acoustic suprasegmental cues to lexical stress (changes in duration, amplitude, and pitch) in spoken-word recognition. We asked here whether they can also use visual suprasegmental cues. In two categorization experiments, Dutch participants saw a speaker say fragments of word pairs that were segmentally identical but differed in their stress realization (e.g., 'ca-vi from cavia "guinea pig" vs. 'ka-vi from kaviaar "caviar"). Participants were able to distinguish between these pairs from seeing a speaker alone. Only the presence of primary stress in the fragment, not its absence, was informative. Participants were able to distinguish visually primary from secondary stress on first syllables, but only when the fragment-bearing target word carried phrase-level emphasis. Furthermore, participants distinguished fragments with primary stress on their second syllable from those with secondary stress on their first syllable (e.g., pro-'jec from projector "projector" vs. 'pro-jec from projectiel "projectile"), independently of phrase-level emphasis. Seeing a speaker thus contributes to spoken-word recognition by providing suprasegmental information about the presence of primary lexical stress.

Abstract

When listening to speech, we often benefit when also seeing the speaker talk. If this benefit is not domain-specific for speech, then the recognition of sung lyrics should likewise benefit from seeing the singer. Nevertheless, previous research failed to obtain a substantial improvement in that domain. Our study shows that this failure was not due to inherent differences between singing and speaking but rather to less informative visual presentations. By presenting a professional singer, we found a substantial audiovisual benefit of about 35% improvement for lyrics recognition. This benefit was further robust across participants, phrases, and repetition of the test materials. Our results provide the first evidence that lyrics recognition just like speech and music perception is a multimodal process.

Abstract

In the present study, we examined the distribution and processing of information over time in auditory and visual speech as it is used in unimodal and bimodal word recognition. English consonant—vowel—consonant words representing all possible initial consonants were presented as auditory, visual, or audiovisual speech in a gating task. The distribution of information over time varied across and within features. Visual speech information was generally fully available early during the phoneme, whereas auditory information was still accumulated. An audiovisual benefit was therefore already found early during the phoneme. The nature of the audiovisual recognition benefit changed, however, as more of the phoneme was presented. More features benefited at short gates rather than at longer ones. Visual speech information plays, therefore, a more important role early during the phoneme rather than later. The results of the study showed the complex interplay of information across modalities and time, since this is essential in determining the time course of audiovisual spoken-word recognition.

Abstract

Past research has demonstrated that infants can rapidly extract syllable distribution information from an artificial language and use this knowledge to infer likely word boundaries in speech. However, artificial languages are extremely simplified with respect to natural language. In this study, we ask whether infants’ ability to track transitional probabilities between syllables in an artificial language can scale up to the challenge of natural language. We do so by testing both 5.5- and 8-month-olds’ ability to segment an artificial language containing four words of uniform length (all CVCV) or four words of varying length (two CVCV, two CVCVCV). The transitional probability cues to word boundaries were held equal across the two languages. Both age groups segmented the language containing words of uniform length, demonstrating that even 5.5-month-olds are extremely sensitive to the conditional probabilities in their environment. However, either age group succeeded in segmenting the language containing words of varying length, despite the fact that the transitional probability cues defining word boundaries were equally strong in the two languages. We conclude that infants’ statistical learning abilities may not be as robust as earlier studies have suggested.

Abstract

In this study, we use the association between various measures of the morphological family and decision latencies to reveal the way in which the components of Dutch and English compounds are processed. The results show that for constituents of concatenated compounds in both languages, a position-related token count of the morphological family plays a role, whereas English open compounds show an effect of a type count, similar to the effect of family size for simplex words. When Dutch compounds are written with an artificial space, they reveal no effect of type count, which shows that the differential effect for the English open compounds is not superficial. The final experiment provides converging evidence for the lexical consequences of the space in English compounds. Decision latencies for English simplex words are better predicted from counts of the morphological family that include concatenated and hyphenated but not open family members.

Abstract

Explanations in the domain of kinship can be sought on several different levels: Jones addresses online processing, as well as issues of origins and innateness. We argue that his framework can more usefully be applied at the levels of developmental and historical change, the latter especially. A phylogenetic approach to the diversity of kinship terminologies is most urgently required.

Abstract

Recent behavioral and electrophysiological evidence has highlighted the long-term importance for language skills of an early ability to recognize words in continuous speech. We here present further tests of this long-term link in the form of follow-up studies conducted with two (separate) groups of infants who had earlier participated in speech segmentation tasks. Each study extends prior follow-up tests: Study 1 by using a novel follow-up measure that taps into online processing, Study 2 by assessing language performance relationships over a longer time span than previously tested. Results of Study 1 show that brain correlates of speech segmentation ability at 10 months are positively related to 16-month-olds’ target fixations in a looking-while-listening task. Results of Study 2 show that infant speech segmentation ability no longer directly predicts language profiles at the age of five. However, a meta-analysis across our results and those of similar studies (Study 3) reveals that age at follow-up does not moderate effect size. Together, the results suggest that infants’ ability to recognize words in speech certainly benefits early vocabulary development; further observed relationships of later language skills to early word recognition may be consequent upon this vocabulary size effect.

Abstract

Most words that infants hear occur within fluent speech. To compile a vocabulary, infants therefore need to segment words from speech contexts. This study is the first to investigate whether infants (here: 10-month-olds) can recognize words when both initial exposure and test presentation are in continuous speech. Electrophysiological evidence attests that this indeed occurs: An increased extended negativity (word recognition effect) appears for familiarized target words relative to control words. This response proved constant at the individual level: Only infants who showed this negativity at test had shown such a response, within six repetitions after first occurrence, during familiarization.

Abstract

The relocation of animals can induce stress when animals are placed in novel environmental conditions. The movement of captive animals among facilities is common, especially for non-human primates used in research. The stress response begins with the activation of the hypothalamic-pituitary-adrenal (HPA) axis which results in the release of glucocorticoid hormones (GC), which at chronic levels could lead to deleterious physiological effects. There is a substantial body of data concerning GC levels affecting reproduction, and rank and aggression in primates. However, the effect of transport has received much less attention. Fecal samples from eight (four male and four female) captive-bred cotton-top tamarins (Saguinus oedipus) were collected at four different time points (two pre-transport and two post-transport). The fecal samples were analyzed using an immunoassay to determine GC levels. A repeated measures analysis of variance (ANOVA) demonstrated that GC levels differed among transport times (p = 0.009), but not between sexes (p = 0.963). Five of the eight tamarins exhibited an increase in GC levels after transport. Seven of the eight tamarins exhibited a decrease in GC levels from three to six days post-transport to three weeks post-transport. Most values returned to pre-transport levels after three weeks. The results indicate that these tamarins experienced elevated GC levels following transport, but these increases were of short duration. This outcome would suggest that the negative effects of elevated GC levels were also of short duration.

Abstract

One of the major challenges in acquiring a language is being able to use morphology as an adult would, and thus, a considerable amount of acquisition research has focused on morphological production and comprehension. Most of this research, however, has focused on the acquisition of morphology in isolating languages, or languages (such as English) with limited inflectional morphology. The nature of the learning task is different, and potentially more challenging, when the child is learning a polysynthetic language – a language in which words are highly morphologically complex, expressing in a single word what in English takes a multi-word clause. To date, there has been no cross-linguistic survey of how children approach this puzzle and learn polysynthetic languages. This paper aims to provide such a survey, including a discussion of some of the general findings in the literature regarding the acquisition of polysynthetic systems

Abstract

Gesture and speech are assumed to form an integrated system during language production. Based on this view, we propose the integrated‐systems hypothesis, which explains two ways in which gesture and speech are integrated—through mutual and obligatory interactions—in language comprehension. Experiment 1 presented participants with action primes (e.g., someone chopping vegetables) and bimodal speech and gesture targets. Participants related primes to targets more quickly and accurately when they contained congruent information (speech: “chop”; gesture: chop) than when they contained incongruent information (speech: “chop”; gesture: twist). Moreover, the strength of the incongruence affected processing, with fewer errors for weak incongruities (speech: “chop”; gesture: cut) than for strong incongruities (speech: “chop”; gesture: twist). Crucial for the integrated‐systems hypothesis, this influence was bidirectional. Experiment 2 demonstrated that gesture’s influence on speech was obligatory. The results confirm the integrated‐systems hypothesis and demonstrate that gesture and speech form an integrated system in language comprehension.

Abstract

We hypothesized that bone resorption acts to increase bone strength through stimulation of periosteal expansion. Hence, we examined whether bone resorption, as reflected by serum β-C-telopeptides of type I collagen (CTX), is positively associated with periosteal circumference (PC), in contrast to inverse associations with parameters related to bone remodeling such as cortical bone mineral density (BMDC ). CTX and mid-tibial peripheral quantitative computed tomography (pQCT) scans were available in 1130 adolescents (mean age 15.5 years) from the Avon Longitudinal Study of Parents and Children (ALSPAC). Analyses were adjusted for age, gender, time of sampling, tanner stage, lean mass, fat mass, and height. CTX was positively related to PC (β=0.19 [0.13, 0.24]) (coefficient=SD change per SD increase in CTX, 95% confidence interval)] but inversely associated with BMDC (β=-0.46 [-0.52,-0.40]) and cortical thickness [β=-0.11 (-0.18, -0.03)]. CTX was positively related to bone strength as reflected by the strength-strain index (SSI) (β=0.09 [0.03, 0.14]). To examine the causal nature of this relationship, we then analyzed whether single-nucleotide polymorphisms (SNPs) within key osteoclast regulatory genes, known to reduce areal/cortical BMD, conversely increase PC. Fifteen such genetic variants within or proximal to genes encoding receptor activator of NF-κB (RANK), RANK ligand (RANKL), and osteoprotegerin (OPG) were identified by literature search. Six of the 15 alleles that were inversely related to BMD were positively related to CTX (p<}0.05 cut-off) (n=2379). Subsequently, we performed a meta-analysis of associations between these SNPs and PC in ALSPAC (n=3382), Gothenburg Osteoporosis and Obesity Determinants (GOOD) (n=938), and the Young Finns Study (YFS) (n=1558). Five of the 15 alleles that were inversely related to BMD were positively related to PC (p{<0.05 cut-off). We conclude that despite having lower BMD, individuals with a genetic predisposition to higher bone resorption have greater bone size, suggesting that higher bone resorption is permissive for greater periosteal expansion.

Abstract

Heritability of bone mineral density (BMD) varies across skeletal sites, reflecting different relative contributions of genetic and environmental influences. To quantify the degree to which common genetic variants tag and environmental factors influence BMD, at different sites, we estimated the genetic (rg) and residual (re) correlations between BMD measured at the upper limbs (UL-BMD), lower limbs (LL-BMD) and skull (SK-BMD), using total-body DXA scans of ∼ 4,890 participants recruited by the Avon Longitudinal Study of Parents and their Children (ALSPAC). Point estimates of rg indicated that appendicular sites have a greater proportion of shared genetic architecture (LL-/UL-BMD rg = 0.78) between them, than with the skull (UL-/SK-BMD rg = 0.58 and LL-/SK-BMD rg = 0.43). Likewise, the residual correlation between BMD at appendicular sites (r(e) = 0.55) was higher than the residual correlation between SK-BMD and BMD at appendicular sites (r(e) = 0.20-0.24). To explore the basis for the observed differences in rg and re, genome-wide association meta-analyses were performed (n ∼ 9,395), combining data from ALSPAC and the Generation R Study identifying 15 independent signals from 13 loci associated at genome-wide significant level across different skeletal regions. Results suggested that previously identified BMD-associated variants may exert site-specific effects (i.e. differ in the strength of their association and magnitude of effect across different skeletal sites). In particular, variants at CPED1 exerted a larger influence on SK-BMD and UL-BMD when compared to LL-BMD (P = 2.01 × 10(-37)), whilst variants at WNT16 influenced UL-BMD to a greater degree when compared to SK- and LL-BMD (P = 2.31 × 10(-14)). In addition, we report a novel association between RIN3 (previously associated with Paget's disease) and LL-BMD (rs754388: β = 0.13, SE = 0.02, P = 1.4 × 10(-10)). Our results suggest that BMD at different skeletal sites is under a mixture of shared and specific genetic and environmental influences. Allowing for these differences by performing genome-wide association at different skeletal sites may help uncover new genetic influences on BMD.