Publications

Abstract

Spinocerebellar ataxia type 14 (SCA14) is a subtype of the autosomal dominant cerebellar ataxias that is characterized by slowly progressive cerebellar dysfunction and neurodegeneration. SCA14 is caused by mutations in the PRKCG gene, encoding protein kinase C gamma (PKCγ). Despite the identification of 40 distinct disease-causing mutations in PRKCG, the pathological mechanisms underlying SCA14 remain poorly understood. Here we report the molecular neuropathology of SCA14 in post-mortem cerebellum and in human patient-derived induced pluripotent stem cells (iPSCs) carrying two distinct SCA14 mutations in the C1 domain of PKCγ, H36R and H101Q. We show that endogenous expression of these mutations results in the cytoplasmic mislocalization and aggregation of PKCγ in both patient iPSCs and cerebellum. PKCγ aggregates were not efficiently targeted for degradation. Moreover, mutant PKCγ was found to be hyper-activated, resulting in increased substrate phosphorylation. Together, our findings demonstrate that a combination of both, loss-of-function and gain-of-function mechanisms are likely to underlie the pathogenesis of SCA14, caused by mutations in the C1 domain of PKCγ. Importantly, SCA14 patient iPSCs were found to accurately recapitulate pathological features observed in post-mortem SCA14 cerebellum, underscoring their potential as relevant disease models and their promise as future drug discovery tools.

Abstract

Previous studies suggest that bilinguals' lexical access is language non-selective, especially for orthographically identical translation equivalents across languages (i.e., identical cognates). The present study investigated how such words (e.g., meaning "school" in both Chinese and Japanese) are processed in the (late) Chinese - Japanese bilingual brain. Using an L2-Japanese lexical decision task, both behavioral and electrophysiological data were collected. Reaction times (RTs), as well as the N400 component, showed that cognates are more easily recognized than non-cognates. Additionally, an early component (i.e., the N250), potentially reflecting activation at the word-form level, was also found. Cognates elicited a more positive N250 than non-cognates in the frontal region, indicating that the cognate facilitation effect occurred at an early stage of word formation for languages with logographic scripts.

Abstract

Language users process utterances by segmenting them into many cognitive units, which vary in their sizes and linguistic levels. Although we can do such unitization/segmentation easily, its cognitive mechanism is still not clear. This paper proposes an unsupervised model, Less-is-Better (LiB), to simulate the human cognitive process with respect to language unitization/segmentation. LiB follows the principle of least effort and aims to build a lexicon which minimizes the number of unit tokens (alleviating the effort of analysis) and number of unit types (alleviating the effort of storage) at the same time on any given corpus. LiB’s workflow is inspired by empirical cognitive phenomena. The design makes the mechanism of LiB cognitively plausible and the computational requirement light-weight. The lexicon generated by LiB performs the best among different types of lexicons (e.g. ground-truth words) both from an information-theoretical view and a cognitive view, which suggests that the LiB lexicon may be a plausible proxy of the mental lexicon.

Abstract

A core question in language acquisition is whether children’s syntactic processing is experience-dependent and language-specific, or whether it is governed by abstract, universal syntactic machinery. We address this question by presenting corpus and on-line processing dat a from children learning Mandarin Chinese, a language that has been important in debates about the universality of parsing processes. The corpus data revealed that two different relative clause constructions in Mandarin are differentially used to modify syntactic subjects and objects. In the experiment, 4-year-old children’s eye-movements were recorded as they listened to the two RC construction types (e.g., Can you pick up the pig that pushed the sheep?). A permutation analysis showed that children’s ease of comprehension was closely aligned with the distributional frequencies, suggesting syntactic processing preferences are shaped by the input experience of these constructions.

Abstract

Chunking in language comprehension is a process that segments continuous linguistic input into smaller chunks that are in the reader’s mental lexicon. Effective chunking during reading facilitates disambiguation and enhances efficiency for comprehension. However, the chunking mechanisms remain elusive, especially in reading given that information arrives simultaneously yet the written systems may not have explicit cues for labeling boundaries such as Chinese. What are the mechanisms of chunking that mediates the reading of the text that contains hierarchical information? We investigated this question by manipulating the lexical status of the chunks at distinct levels in four-character Chinese strings, including the two-character local chunk and four-character global chunk. Male and female human participants were asked to make lexical decisions on these strings in a behavioral experiment, followed by a passive reading task when their electroencephalography (EEG) was recorded. The behavioral results showed that the lexical decision time of lexicalized two-character local chunks was influenced by the lexical status of the four-character global chunk, but not vice versa, which indicated the processing of global chunks possessed priority over the local chunks. The EEG results revealed that familiar lexical chunks were detected simultaneously at both levels and further processed in a different temporal order – the onset of lexical access for the global chunks was earlier than that of local chunks. These consistent results suggest a two-stage operation for chunking in reading–– the simultaneous detection of familiar lexical chunks at multiple levels around 100 ms followed by recognition of chunks with global precedence.

Abstract

Electroencephalography (EEG) provides high temporal resolution cognitive information from non-invasive recordings. However, one of the common practices-using a subset of sensors in ERP analysis is hard to provide a holistic and precise dynamic results. Selecting or grouping subsets of sensors may also be subject to selection bias, multiple comparison, and further complicated by individual differences in the group-level analysis. More importantly, changes in neural generators and variations in response magnitude from the same neural sources are difficult to separate, which limit the capacity of testing different aspects of cognitive hypotheses. We introduce EasyEEG, a toolbox that includes several multivariate analysis methods to directly test cognitive hypotheses based on topographic responses that include data from all sensors. These multivariate methods can investigate effects in the dimensions of response magnitude and topographic patterns separately using data in the sensor space, therefore enable assessing neural response dynamics. The concise workflow and the modular design provide user-friendly and programmer-friendly features. Users of all levels can benefit from the open-sourced, free EasyEEG to obtain a straightforward solution for efficient processing of EEG data and a complete pipeline from raw data to final results for publication.

Abstract

In a masked priming word-naming task, a facilitation due to the initial-segmental sound overlap for 2-character kanji prime-target pairs was affected by certain orthographic properties (Yoshihara, Nakayama, Verdonschot, & Hino, 2017). That is, the facilitation that was due to the initial mora overlap occurred only when the mora was the whole pronunciation of their initial kanji characters (i.e., match pairs; e.g., /ka-se.ki/-/ka-rjo.ku/). When the shared initial mora was only a part of the kanji characters' readings, however, there was no facilitation (i.e., mismatch pairs; e.g., /ha.tu-a.N/-/ha.ku-bu.tu/). In the present study, we used a masked priming picture-naming task to investigate whether the previous results were relevant only when the orthography of targets is visually presented. In Experiment 1. the main findings of our word-naming task were fully replicated in a picture-naming task. In Experiments 2 and 3. the absence of facilitation for the mismatch pairs were confirmed with a new set of stimuli. On the other hand, a significant facilitation was observed for the match pairs that shared the 2 initial morae (in Experiment 4), which was again consistent with the results of our word-naming study. These results suggest that the orthographic properties constrain the phonological expression of masked priming for kanji words across 2 tasks that are likely to differ in how phonology is retrieved. Specifically, we propose that orthography of a word is activated online and constrains the phonological encoding processes in these tasks.

Abstract

Word learning happens in everyday contexts with many words and many potential referents for those words in view at the same time. It is challenging for young learners to find the correct referent upon hearing an unknown word at the moment. This problem of referential uncertainty has been deemed as the crux of early word learning (Quine, 1960). Recent empirical and computational studies have found support for a statistical solution to the problem termed cross-situational learning. Cross-situational learning allows learners to acquire word meanings across multiple exposures, despite each individual exposure is referentially uncertain. Recent empirical research shows that infants, children and adults rely on cross-situational learning to learn new words (Smith & Yu, 2008; Suanda, Mugwanya, & Namy, 2014; Yu & Smith, 2007). However, researchers have found evidence supporting two very different theoretical accounts of learning mechanisms: Hypothesis Testing (Gleitman, Cassidy, Nappa, Papafragou, & Trueswell, 2005; Markman, 1992) and Associative Learning (Frank, Goodman, & Tenenbaum, 2009; Yu & Smith, 2007). Hypothesis Testing is generally characterized as a form of learning in which a coherent hypothesis regarding a specific word-object mapping is formed often in conceptually constrained ways. The hypothesis will then be either accepted or rejected with additional evidence. However, proponents of the Associative Learning framework often characterize learning as aggregating information over time through implicit associative mechanisms. A learner acquires the meaning of a word when the association between the word and the referent becomes relatively strong. In this chapter, we consider these two psychological theories in the context of cross-situational word-referent learning. By reviewing recent empirical and cognitive modeling studies, our goal is to deepen our understanding of the underlying word learning mechanisms by examining and comparing the two theoretical learning accounts.

Abstract

To acquire the meaning of a verb, language learners not only need to find the correct mapping between a specific verb and an action or event in the world, but also infer the underlying relational meaning that the verb encodes. Most verb naming instances in naturalistic contexts are highly ambiguous as many possible actions can be embedded in the same scenario and many possible verbs can be used to describe those actions. To understand whether learners can find the correct verb meaning from referentially ambiguous learning situations, we conducted three experiments using the Human Simulation Paradigm with adult learners. Our results suggest that although finding the right verb meaning from one learning instance is hard, there is a statistical solution to this problem. When provided with multiple verb learning instances all referring to the same verb, learners are able to aggregate information across situations and gradually converge to the correct semantic space. Even in cases where they may not guess the exact target verb, they can still discover the right meaning by guessing a similar verb that is semantically close to the ground truth.

Abstract

Bilinguals usually select the right language to speak for the particular context they are in, but sometimes the nontarget language intrudes. Despite a large body of research into language selection and language control, it remains unclear where intrusion errors originate from. These errors may be due to incorrect selection of the nontarget language at the conceptual level, or be a consequence of erroneous word selection (despite correct language selection) at the lexical level. We examined the former possibility in two language switching experiments using a manipulation that supposedly affects language selection on the conceptual level, namely whether the conversational language context was associated with the target language (congruent) or with the alternative language (incongruent) on a trial. Both experiments showed that language intrusion errors occurred more often in incongruent than in congruent contexts, providing converging evidence that language selection during concept preparation is one driving force behind language intrusion.

Abstract

Bilingual speakers have to control their languages to avoid interference, which may be achieved by enhancing the target language and/or inhibiting the nontarget language. Previous research suggests that bilinguals use inhibition (e.g., Jackson et al., 2001), which should be reflected in the N2 component of the event-related potential (ERP) in the EEG. In the current study, we investigated the dynamics of inhibitory control by measuring the N2 during language switching and repetition in bilingual picture naming. Participants had to name pictures in Dutch or English depending on the cue. A run of same-language trials could be short (two or three trials) or long (five or six trials). We assessed whether RTs and N2 changed over the course of same-language runs, and at a switch between languages. Results showed that speakers named pictures more quickly late as compared to early in a run of same-language trials. Moreover, they made a language switch more quickly after a long run than after a short run. This run-length effect was only present in the first language (L1), not in the second language (L2). In ERPs, we observed a widely distributed switch effect in the N2, which was larger after a short run than after a long run. This effect was only present in the L2, not in the L1, although the difference was not significant between languages. In contrast, the N2 was not modulated during a same-language run. Our results suggest that the nontarget language is inhibited at a switch, but not during the repeated use of the target language.

Abstract

Although bilingual speakers are very good at selectively using one language rather than another, sometimes language selection errors occur. To investigate how bilinguals monitor their speech errors and control their languages in use, we recorded event-related potentials (ERPs) in unbalanced Dutch-English bilingual speakers in a cued language-switching task. We tested the conflict-based monitoring model of Nozari and colleagues by investigating the error-related negativity (ERN) and comparing the effects of the two switching directions (i.e., to the first language, L1 vs. to the second language, L2). Results show that the speakers made more language selection errors when switching from their L2 to the L1 than vice versa. In the EEG, we observed a robust ERN effect following language selection errors compared to correct responses, reflecting monitoring of speech errors. Most interestingly, the ERN effect was enlarged when the speakers were switching to their L2 (less conflict) compared to switching to the L1 (more conflict). Our findings do not support the conflict-based monitoring model. We discuss an alternative account in terms of error prediction and reinforcement learning.

Abstract

Although bilingual speakers are very good at selectively using one language rather than another, sometimes language selection errors occur. We examined the relative contribution of top-down cognitive control and bottom-up language priming to these errors. Unbalanced Dutch-English bilinguals named pictures and were cued to switch between languages under time pressure. We also manipulated the number of same-language trials before a switch (long vs. short runs). Results show that speakers made more language selection errors when switching from their second language (L2) to the first language (L1) than vice versa. Furthermore, they made more errors when switching to the L1 after a short compared to a long run of L2 trials. In the reverse switching direction (L1 to L2), run length had no effect. These findings are most compatible with an account of language selection errors that assigns a strong role to top-down processes of cognitive control.

Abstract

In sentences like “the mouse that chased the cat was hungry”, the syntactically correct interpretation (the mouse chases the cat) is contradicted by semantic and pragmatic knowledge. Previous research has shown that L1 speakers sometimes base sentence interpretation on this type of knowledge (so-called “shallow” or “good-enough” processing). We made use of both behavioural and ERP measurements to investigate whether L2 learners differ from native speakers in the extent to which they engage in “shallow” syntactic processing. German learners of Dutch as well as Dutch native speakers read sentences containing relative clauses (as in the example above) for which the plausible thematic roles were or were not reversed, and made plausibility judgments. The results show that behaviourally, L2 learners had more difficulties than native speakers to discriminate plausible from implausible sentences. In the ERPs, we replicated the previously reported finding of a “semantic P600” for semantic reversal anomalies in native speakers, probably reflecting the effort to resolve the syntax-semantics conflict. In L2 learners, though, this P600 was largely attenuated and surfaced only in those trials that were judged correctly for plausibility. These results generally point at a more prevalent, but not exclusive occurrence of shallow syntactic processing in L2 learners.

Abstract

Semantic unification during sentence comprehension has been associated with amplitude change of the N400 in event-related potential (ERP) studies, and activation in the left inferior frontal gyrus (IFG) in functional magnetic resonance imaging (fMRI) studies. However, the specificity of this activation to semantic unification remains unknown. To more closely examine the brain processes involved in semantic unification, we employed simultaneous EEG-fMRI to time-lock the semantic unification related N400 change, and integrated trial-by-trial variation in both N400 and BOLD change beyond the condition-level BOLD change difference measured in traditional fMRI analyses. Participants read sentences in which semantic unification load was parametrically manipulated by varying cloze probability. Separately, ERP and fMRI results replicated previous findings, in that semantic unification load parametrically modulated the amplitude of N400 and cortical activation. Integrated EEG-fMRI analyses revealed a different pattern in which functional activity in the left IFG and bilateral supramarginal gyrus (SMG) was associated with N400 amplitude, with the left IFG activation and bilateral SMG activation being selective to the condition-level and trial-level of semantic unification load, respectively. By employing the EEG-fMRI integrated analyses, this study among the first sheds light on how to integrate trial-level variation in language comprehension.

Abstract

Genome-wide association studies using thousands to hundreds of thousands of single nucleotide polymorphism (SNP) markers and region-wide association studies using a dense panel of SNPs are already in use to identify disease susceptibility genes and to predict disease risk in individuals. Because these tasks become increasingly important, three different data sets were provided for the Genetic Analysis Workshop 15, thus allowing examination of various novel and existing data mining methods for both classification and identification of disease susceptibility genes, gene by gene or gene by environment interaction. The approach most often applied in this presentation group was random forests because of its simplicity, elegance, and robustness. It was used for prediction and for screening for interesting SNPs in a first step. The logistic tree with unbiased selection approach appeared to be an interesting alternative to efficiently select interesting SNPs. Machine learning, specifically ensemble methods, might be useful as pre-screening tools for large-scale association studies because they can be less prone to overfitting, can be less computer processor time intensive, can easily include pair-wise and higher-order interactions compared with standard statistical approaches and can also have a high capability for classification. However, improved implementations that are able to deal with hundreds of thousands of SNPs at a time are required.

Abstract

The purpose of this study is to shed light to the linguistic deficit in populations with dementia, and more specifically with Mild Cognitive Impairment and Alzheimer’s Disease; by examining the assignment of thematic roles (θ-roles) in sentences including psychological verbs.
The interest in types of dementia and its precursor is due to the relevance of the disease in present-day world society (Caloi, 2017). 47 millions of people worldwide were reported by the World Alzheimer Report in 2016 (Prince et al. 2016) as people with a type of dementia. This number surpasses the number of inhabitants in Spain, a whole country, and it is expected, according to the report, to triplicate until 2050 reaching the number of 131 million. The impact of this disease is observed not only at the social level but also in the economic one, because of their need for assistance in their everyday life. What is worrying, is the lack of total treatment once the disease has started. Despite the efforts of medicine, dementia is problematic in terms of its diagnosis, because a variety of cognitive abilities is assessed in combination with medical workup. Language is a crucial component in the procedure of diagnosis as linguistic deficits are among the first symptoms that accompany the onset of the disease. Therefore, further investigation of linguistic impairment is a necessity in order to enhance the diagnostic techniques used nowadays. Furthermore, the lack of efficient drugs for the treatment of the disease has necessitated the development of training programs for maintenance and increase of the cognitive abilities in people with either Mild Cognitive Impairment or a type of dementia …

Abstract

In informal interaction, speakers rarely thank a person who has complied with a request. Examining data from British English, German, Italian, Polish, and Telugu, we ask when speakers do thank after compliance. The results show that thanking treats the other’s assistance as going beyond what could be taken for granted in the circumstances. Coupled with the rareness of thanking after requests, this suggests that cooperation is to a great extent governed by expectations of helpfulness, which can be long-standing, or built over the course of a particular interaction. The higher frequency of thanking in some languages (such as English or Italian) suggests that cultures differ in the importance they place on recognizing the other’s agency in doing as requested.

Abstract

It is undisputed that presenting a rhythmic stimulus leads to a measurable brain response that follows the rhythmic structure of this stimulus. What is still debated, however, is the question whether this brain response exclusively reflects a regular repetition of evoked responses, or whether it also includes entrained oscillatory activity. Here we systematically present evidence in favor of an involvement of entrained neural oscillations in the processing of rhythmic input while critically pointing out which questions still need to be addressed before this evidence could be considered conclusive. In this context, we also explicitly discuss the potential functional role of such entrained oscillations, suggesting that these stimulus-aligned oscillations reflect, and serve as, predictive processes, an idea often only implicitly assumed in the literature.

Abstract

Change in linguistic prosody generates a mismatch negativity response (MMN), indicating neural representation of linguistic prosody, while change in affective prosody generates a positive response (P3a), reflecting its motivational salience. However, the neural response to concurrent affective and linguistic prosody is unknown. The present paper investigates the integration of these two prosodic features in the brain by examining the neural response to separate and concurrent processing by electroencephalography (EEG). A spoken pair of Swedish words—[ˈfɑ́ːsɛn] phase and [ˈfɑ̀ːsɛn] damn—that differed in emotional semantics due to linguistic prosody was presented to 16 subjects in an angry and neutral affective prosody using a passive auditory oddball paradigm. Acoustically matched pseudowords—[ˈvɑ́ːsɛm] and [ˈvɑ̀ːsɛm]—were used as controls. Following the constructionist concept of emotions, accentuating the conceptualization of emotions based on language, it was hypothesized that concurrent affective and linguistic prosody with the same valence—angry [ˈfɑ̀ːsɛn] damn—would elicit a unique late EEG signature, reflecting the temporal integration of affective voice with emotional semantics of prosodic origin. In accordance, linguistic prosody elicited an MMN at 300–350 ms, and affective prosody evoked a P3a at 350–400 ms, irrespective of semantics. Beyond these responses, concurrent affective and linguistic prosody evoked a late positive component (LPC) at 820–870 ms in frontal areas, indicating the conceptualization of affective prosody based on linguistic prosody. This study provides evidence that the brain does not only distinguish between these two functions of prosody but also integrates them based on language and experience.

Abstract

Swedish morphemes are classified as prosodically specified or prosodically unspecified, depending on lexical or phonological stress, respectively. Here, we investigate the allomorphy of the suffix -(i)sk, which indicates the distinction between lexical and phonological stress; if attached to a lexically stressed morpheme, it takes a non-syllabic form (-sk), whereas if attached to a phonologically stressed morpheme, an epenthetic vowel is inserted (-isk). Using mismatch negativity (MMN), we explored the neural processing of this allomorphy across lexically stressed and phonologically stressed morphemes. In an oddball paradigm, participants were occasionally presented with congruent and incongruent derivations, created by the suffix -(i)sk, within the repetitive presentation of their monomorphemic stems. The results indicated that the congruent derivation of the lexically stressed stem elicited a larger MMN than the incongruent sequences of the same stem and the derivational suffix, whereas after the phonologically stressed stem a non-significant tendency towards an opposite pattern was observed. We argue that the significant MMN response to the congruent derivation in the lexical stress condition is in line with lexical MMN, indicating a holistic processing of the sequence of lexically stressed stem and derivational suffix. The enhanced MMN response to the incongruent derivation in the phonological stress condition, on the other hand, is suggested to reflect combinatorial processing of the sequence of phonologically stressed stem and derivational suffix. These findings bring a new aspect to the dual-system approach to neural processing of morphologically complex words, namely the specification of word stress.

Abstract

Speakers remember their own utterances better than those of their interlocutors, suggesting that language production is beneficial to memory. This may be partly explained by a generation effect: The act of generating a word is known to lead to a memory advantage (Slamecka & Graf, 1978). In earlier work, we showed a generation effect for recognition of images (Zormpa, Brehm, Hoedemaker, & Meyer, 2019). Here, we tested whether the recognition of their names would also benefit from name generation. Testing whether picture naming improves memory for words was our primary aim, as it serves to clarify whether the representations affected by generation are visual or conceptual/lexical. A secondary aim was to assess the influence of processing time on memory. Fifty-one participants named pictures in three conditions: after hearing the picture name (identity condition), backward speech, or an unrelated word. A day later, recognition memory was tested in a yes/no task. Memory in the backward speech and unrelated conditions, which required generation, was superior to memory in the identity condition, which did not require generation. The time taken by participants for naming was a good predictor of memory, such that words that took longer to be retrieved were remembered better. Importantly, that was the case only when generation was required: In the no-generation (identity) condition, processing time was not related to recognition memory performance. This work has shown that generation affects conceptual/lexical representations, making an important contribution to the understanding of the relationship between memory and language.

Abstract

The production effect (better memory for words read aloud than words read silently) and the picture superiority effect (better memory for pictures than words) both improve item memory in a picture naming task (Fawcett, J. M., Quinlan, C. K., & Taylor, T. L. (2012). Interplay of the production and picture superiority effects: A signal detection analysis. Memory (Hove, England), 20(7), 655–666. doi:10.1080/09658211.2012.693510). Because picture naming requires coming up with an appropriate label, the generation effect (better memory for generated than read words) may contribute to the latter effect. In two forced-choice memory experiments, we tested the role of generation in a picture naming task on later recognition memory. In Experiment 1, participants named pictures silently or aloud with the correct name or an unreadable label superimposed. We observed a generation effect, a production effect, and an interaction between the two. In Experiment 2, unreliable labels were included to ensure full picture processing in all conditions. In this experiment, we observed a production and a generation effect but no interaction, implying the effects are dissociable. This research demonstrates the separable roles of generation and production in picture naming and their impact on memory. As such, it informs the link between memory and language production and has implications for memory asymmetries between language production and comprehension.

Abstract

There is a rich tradition of building computational models in cognitive science, but modeling, theoretical, and experimental research are not as tightly integrated as they could be. In this paper, we show that computational techniques—even simple ones that are straightforward to use—can greatly facilitate designing, implementing, and analyzing experiments, and generally help lift research to a new level. We focus on the domain of artificial grammar learning, and we give five concrete examples in this domain for (a) formalizing and clarifying theories, (b) generating stimuli, (c) visualization, (d) model selection, and (e) exploring the hypothesis space.