Brain and Language Lab


The lab investigates the neurocognition of language, memory, and other aspects of cognition in healthy populations and disorders. To find out more (e.g., on Research or Publications), select an option from the drop-down menu (three bars at the top left). Listed below are some recent findings and other developments that may be of interest.


Developmental language disorder (DLD) is a common neurodevelopmental disorder with adverse impacts that continue into adulthood. However, its neural bases remain unclear. Here we address this gap by systematically identifying and quantitatively synthesizing neuroanatomical studies of DLD using co-localization likelihood estimation, a recently developed neuroanatomical meta-analytic technique. Analyses of structural brain data (22 peer-reviewed papers, 577 participants) revealed highly consistent anomalies only in the basal ganglia (100% of participant groups in which this structure was examined, weighted by group sample sizes; 99.8% permutation-based likelihood the anomaly clustering was not due to chance). These anomalies were localized specifically to the anterior neostriatum (again 100% weighted proportion and 99.8% likelihood). As expected given the task dependence of activation, functional neuroimaging data (11 peer-reviewed papers, 414 participants) yielded less consistency, though anomalies again occurred primarily in the basal ganglia (79.0% and 95.1%). Multiple sensitivity analyses indicated that the patterns were robust. The meta-analyses elucidate the neuroanatomical signature of DLD, and implicate the basal ganglia in particular. The findings support the procedural circuit deficit hypothesis of DLD, have basic research and translational implications for the disorder, and advance our understanding of the neuroanatomy of language.

Figure from Janascek et al. 2022


Cognitive neuroscience has highlighted the cerebral cortex while often overlooking subcortical structures. This cortical proclivity is found in basic and translational research on many aspects of cognition, especially higher cognitive domains such as language, reading, music, and math. We suggest that, for both anatomical and evolutionary reasons, multiple subcortical structures play substantial roles across higher and lower cognition. We present a comprehensive review of existing evidence, which indeed reveals extensive subcortical contributions in multiple cognitive domains. We argue that the findings are overall both real and important. Next, we advance a theoretical framework to capture the nature of (sub)cortical contributions to cognition. Finally, we propose how new subcortical cognitive roles can be identified by leveraging anatomical and evolutionary principles, and we describe specific methods that can be used to reveal subcortical cognition. Altogether, this review aims to advance cognitive neuroscience by highlighting subcortical cognition and facilitating its future investigation. 


Many but not all cognitive abilities decline during ageing. Some even improve due to lifelong experience. The critical capacities of attention and executive functions have been widely posited to decline. However, these capacities are composed of multiple components, so multifaceted ageing outcomes might be expected. Indeed, prior findings suggest that whereas certain attention/executive functions clearly decline, others do not, with hints that some might even improve. We tested ageing effects on the alerting, orienting and executive (inhibitory) networks posited by Posner and Petersen’s influential theory of attention, in a cross-sectional study of a large sample (N= 702) of participants aged 58–98. Linear and nonlinear analyses revealed that whereas the efficiency of the alerting network decreased with age, orienting and executive inhibitory efficiency increased, at least until the mid-to-late 70s. Sensitivity analyses indicated that the patterns were robust. The results suggest variability in age-related changes across attention/executive functions, with some declining while others improve.


We  investigated  whether  learning  and  retaining  vocabulary  in  a  second  language  (L2)  can  be  improved  by  leveraging  a  combination  of  memory  enhancement  techniques.  Specifically,  we  tested whether combining retrieval practice, spacing, and related manipulations in a ‘multidomain’ pedagogical approach enhances vocabulary acquisition as compared to a typical learning approach. In a classroom-laboratory design, 48 Turkish university students studying L2 English were trained on  64  English  words  over  17  days.  They  were  assigned  to  either  a  ‘typical’  study  regimen  of  (re)studying the words on the first day (initial study) and last day (cramming) of training, or an ‘optimized’ regimen of retrieval practice (retrieving the words), moreover with feedback, spaced throughout  the  period,  moreover  with  expanding  gaps.  The  target  words  were  tested  before  training (pre-test) and one and 11 days afterwards (post-tests). Mixed-effects modeling revealed a training-group by test-session interaction, due to greater improvements from optimized training (a striking 18 percentage-point accuracy increase from pre-test to both post-tests) than typical training  (an  8  percentage-point  increase).  Further  analyses  showed  that  the  optimized  training  advantages were mainly driven by high (rather than low) frequency words. Overall, the results suggest  that  a  multidomain  approach  of  combining  different  memory  enhancement  techniques  can lead to substantial gains in both the learning and retention of L2 words, as compared to a typical study pattern. The findings have implications for L2 learning and pedagogy.


Lexical-processing declines are a hallmark of aging. However, the extent of these declines may vary as a function of different factors. Motivated by findings from neurodegenerative diseases and healthy aging, we tested whether ‘motor-relatedness’ (the degree to which words are associated with particular human body movements) might moderate such declines. We investigated this question by examining data from three experiments. The experiments were carried out in different languages (Dutch, German, English) using different tasks (lexical decision, picture naming), and probed verbs and nouns, in all cases controlling for potentially confounding variables (e.g., frequency, age-of-acquisition, imageability). Whereas ‘non-motor words’ (e.g., steak) showed age-related performance decreases in all three experiments, ‘motor words’ (e.g., knife) yielded either smaller decreases (in one experiment) or no decreases (in two experiments). The findings suggest that motor-relatedness can attenuate or even prevent age-related lexical declines, perhaps due to the relative sparing of neural circuitry underlying such words.  


Education appears to protect older adults, especially women, against memory loss, according to a new study. The results suggest that children—especially girls—who attend school for longer will have better declarative memory abilities in old age. Participants were shown drawings of objects, and then were tested several minutes later on their memory of these objects. Memory performance became progressively worse with aging. However, more years of early-life education countered these losses, especially in women.  In men, the memory gains associated with each year of education were two times larger than the losses experienced during each year of aging. However, in women, the gains were five times larger.  For example, the declarative memory abilities of an 80-year-old woman with a bachelor’s degree would be as good as those of a 60-year-old woman with a high school education. So, four extra years of education make up for the memory losses from 20 years of aging. The study may have  implications for education, as well as for memory loss in Alzheimer’s disease and other dementias. 

Michael Ullman gave an updated talk on the contributions of the declarative and procedural learning and memory brain circuits to typical and atypical language, at the University of Potsdam in Germany in November 2020. The talk may be useful in providing an up-to-date overview of the Brain and Language Lab's research program examining the role of the two learning and memory systems in language (that is, the declarative/procedural model of language). The talk first lays out the evolutionary and biological motivations for the research program. Then it summarizes the characteristics and neurobiology of the two learning systems, together with their interactions. Next it lays out the range of predictions for language, based on this independent understanding of the learning systems. It then examines the relevant evidence for roles of the learning systems in typical first and second language and in developmental disorders, with a focus on developmental language disorder. Finally, it summarizes newer lines of work, including the investigation of the roles of the systems in speech-sound representations; in language in aging; in enhancing second language learning and in therapeutic approaches for language disorders; and in learning to read and learning math, including contributions of the systems to developmental dyslexia and math disability.

Figure from Reifegerste et al 2020


This study examines language in Parkinson's disease (PD). The results suggest that the production of complex grammatical forms (e.g., producing walk + -ed) is impaired in PD, but only in males, and that this impairment is due to left basal ganglia degeneration in the disorder. In contrast to male patients, female patients appear to compensate for the basal ganglia degeneration by storing such forms as whole words in declarative memory (e.g., memorizing "walked"), thanks to female advantages at this memory system. Thus, the study shows that declarative memory can be used to compensate for grammatical deficits in PD, but that female PD patients are significantly more successful at such compensation than male patients. The study has clinical implications, since it suggests that enhancing declarative memory could enhance such compensation, in both female and male patients. 


Bilingualism affects the structure of the brain in adults, as evidenced by experience-dependent grey and white matter changes in brain structures implicated in language learning, processing, or control. However, limited evidence exists on how bilingualism may influence brain development. We examined the developmental patterns of both grey and white matter structures in a cross-sectional study of a large sample (n=711 for grey matter, n=637 for white matter) of bilingual and monolingual participants, aged 3-21 years. Metrics of grey matter (thickness, volume, surface area) and white matter (fractional anisotropy, mean diffusivity) were examined across 41 cortical and subcortical brain structures and 20 tracts, respectively. We used generalized additive modelling to analyze whether, how, and where the developmental trajectories of bilinguals and monolinguals might differ. Bilingual and monolingual participants manifested distinct developmental trajectories in both grey and white matter structures. As compared to monolinguals, bilinguals showed: a) more grey matter (less developmental loss) starting during late childhood and adolescence, mainly in frontal and parietal regions (particularly in the inferior frontal gyrus pars opercularis, superior frontal cortex, inferior and superior parietal cortex, and precuneus); and b) higher white matter integrity (greater developmental increase) starting during mid-late adolescence, specifically in striatal-inferior frontal fibers. The data suggest that there may be a developmental basis to the well-documented structural differences in the brain between bilingual and monolingual adults.  

Figure 1 Earle et al. 2020 paper


This study tested whether learning to read depends on declarative and procedural memory, two general-purpose learning and memory systems in the brain.  We investigated this hypothesis in a longitudinal study of 140 children, who were tested four times annually between 1st and 4th grade.  The study found that declarative learning abilities predicted reading abilities in first grade, while procedural learning abilities predicted reading ability in second grade. The findings suggest that reading acquisition may indeed depend on these well-studied learning and memory systems. The study has educational and clinical implications.  

Paper title and authors from Ullman et al, 2020


Developmental disorders of language include developmental language disorder, dyslexia, and motor-speech disorders such as articulation disorder and stuttering. These disorders have generally been explained by accounts that focus on their behavioral rather than neural characteristics; their processing rather than learning impairments; and each disorder separately rather than together, despite their commonalities and comorbidities. Here we update and review a unifying neurocognitive account—the Procedural circuit Deficit Hypothesis (PDH). The PDH posits that abnormalities of brain structures underlying procedural memory (learning and memory that rely on the basal ganglia and associated circuitry) can explain numerous brain and behavioral characteristics across learning and processing, in multiple disorders, including both commonalities and differences. We describe procedural memory, examine its role in various aspects of language, and then present the PDH and relevant evidence across language-related disorders. The PDH has substantial explanatory power, and both basic research and translational implications.

Title for Ullman 2020 Book Chapter


This chapter is an updated in-depth exposition of the declarative/procedural model, particularly as it applies to second language and bilingualism. After a primer on the brain, the chapter lays out background on the declarative and procedural memory systems, then predictions for both first and second language based on this independent knowledge of the memory systems, followed by the types of evidence that can test these predictions. Further sections discuss common misunderstandings about the declarative/procedural model, including comparisons between this and other models; a description of an exemplary study that tests the model in second language; and a discussion of the model with respect to the explicit/implicit debate.

Image for Koch et al. 2020

KOCH ET AL. 2020

Using a novel eye-tracking version of the serial reaction time (SRT) task, this study reveals that 9-month old infants can learn sequences. Moreover, direct comparisons with adults showed that infants and adults do not differ in how well they learn the sequences. This study is the first to report learning in an SRT task in infants as young as 9 months. The results suggest that procedural memory, that is, basal ganglia-based learning, is already highly functional as early as 9 month. The study also suggests that this eye-tracking SRT protocol is a promising procedure for measuring procedural memory in infants.  


In a neuroanatomical meta-analysis of the serial reaction time task, we systematically examined the neural bases of sequence learning, which underlies numerous motor, cognitive, and social skills. Controlling for visual, motor, and other factors (in sequence-random block contrasts), sequence learning yielded consistent activation only in the basal ganglia, across the striatum (anterior/mid caudate nucleus and putamen) and the globus pallidus. In contrast, when visual, motor, and other factors were not controlled for, premotor cortical and cerebellar activation were additionally observed. The study provides solid evidence that, at least as tested with the serial reaction time task, sequence learning in humans relies on the basal ganglia, whereas cerebellar and premotor regions appear to contribute to aspects of the task not related to sequence learning itself.

graphic for Tagarelli et al, article 2019


Grammar learning activates anterior caudate/putamen (procedural memory) structures, while word learning activates ventral stream occipito-temporal (declarative memory) structures. Moreover, grammar learning predicted to rely especially on declarative memory (e.g., with explicit training) shows hippocampal involvement, while grammar learning predicted to rely particularly on procedural memory (e.g., with implicit training) shows anterior caudate/putamen involvement.


Lower reliability in procedural learning tasks leads to larger rather than smaller effect sizes for procedural learning deficits in children with specific language impairment (developmental language disorder).


Female patients with Parkinson's disease are less impaired on aspects of grammar than male patients, likely due to stronger declarative memory-based compensation in females.


Working memory in older adults declines with age, but is modulated by sex and education: evidence from a study of 754 older adults in Taiwan.


Evidence that procedural memory is enhanced in children with Tourette syndrome.


Child first language and adult second language are both tied to general-purpose learning systems in the brain that are evolutionarily ancient, and are also found in other vertebrates. 

Title and authors for Ullman and Lovelett 2018


The declarative/procedural (DP) model posits that the learning, storage, and use of language critically depend on two learning and memory systems in the brain: declarative memory and procedural memory. Thus, on the basis of independent research on the memory systems, the model can generate specific and often novel predictions for language. Till now most such predictions and ensuing empirical work have been motivated by research on the neurocognition of the two memory systems. However, there is also a large literature on techniques that enhance learning and memory. The DP model provides a theoretical framework for predicting which techniques should extend to language learning, and in what circumstances they should apply. In order to lay the neurocognitive groundwork for these predictions, here we first summarize the neurocognitive fundamentals of the two memory systems and briefly lay out the resulting claims of the DP model for both first and second language. We then provide an overview of learning and memory enhancement techniques before focusing on two techniques – spaced repetition and retrieval practice – that have been linked to the memory systems. Next, we present specific predictions for how these techniques should enhance language learning, and review existing evidence, which suggests that they do indeed improve the learning of both first and second language. Finally, we discuss areas of future research and implications for second language pedagogy.

GREY ET AL. 2017

Does being bilingual help your brain learn additional languages? The answer seems to be yes: if you've learned two languages early in life, it may help your brain learn another language later on. 


The neural bases of the learning and generalization of affixal morphology: an fMRI study. 


Multiple micronutrient supplementation of pregnant women benefits the cognition of their children at ages 9-12 years, as shown in a study that examined 2879 children in Lombok, Indonesia. 


Evidence that consolidation in declarative memory may be enhanced in specific language impairment (SLI; that is, developmental language disorder). 


Abnormalities of brain structures underlying procedural memory may help explain developmental math disability. 

Chart image from Dye et al, 2016

DYE ET AL. 2016

Children with Tourette syndrome show evidence for speeded grammatical combination in phonology (in a nonword repetition task), complementing previous evidence for speeded combination in morphology. 

chart image from Ullman and Pullman, 2015


Declarative memory compensates for multiple deficits across neurodevelopmental disorders, including deficits of social skills in autism, reading in dyslexia, and grammar in specific language impairment (SLI; that is, developmental language disorder). 

graphic from Lum et al, article 2015

LUM ET AL. 2015

Declarative memory is normal in children with specific language impairment (SLI; that is, developmental language disorder) unless they also have working memory problems.

graphic from Clark et al, article 2015


Procedural learning is impaired in Parkinson’s disease (PD): Evidence from a meta-analysis of serial reaction time studies.

graphic from Walenski et al, article 2014


Children with autism show evidence for speeded grammatical combination in morphology. 

graphic from Lum et al, article 2014

LUM ET AL. 2014

Procedural learning is impaired in specific language impairment (SLI; that is, developmental language disorder): Evidence from a meta-analysis of serial reaction time studies.

graphic from Bowden et al, article 2013


Native-like brain processing of syntax can be attained by university foreign language learners.

graphic from Dye et al, article 2013

DYE ET AL. 2013

Imageability and frequency effects of regular and irregular past tense forms in children: Evidence that all children store irregulars, but only girls store regulars.

graphic from Lum et al, article 2013

LUM ET AL. 2013

Procedural learning is impaired in dyslexia: Evidence from a meta-analysis of serial reaction time studies.

graphic from Hedenius et al, article 2013


Children with developmental dyslexia (DD) are better than typically developing (TD) children at learning and retaining non-verbal information in declarative memory.

graphic from Prado et al, article 2012A


Multiple micronutrient supplementation benefits the cognition of pregnant women, in a study examining 640 women in Lombok, Indonesia.

graphic from Prado et al, article 2012B


Multiple micronutrient supplementation of pregnant women who were undernourished benefits the cognition of their children at ages 3-4 years.

graphic from Phillips et al, article 2012


In patients with early Parkinson’s disease, subthalamic nucleus deep brain stimulation affects grammatical (but not lexical) abilities, and naming manipulated (but not non-manipulated) objects.

graphic from Newman et al, article 2012


Both level of proficiency and native vs. late learning of a language independently affect brain processing of lexical/semantics.

graphic from Nemeth et al, article 2012


Patients with pre-symptomatic Huntington’s disease (pre-HD) show evidence of over-active grammatical rule use: Evidence from native speakers of Hungarian.

graphic from Morgan-Short et al, article 2012A


Immersion-like second language training leads to more native-like brain patterns for grammar than classroom-like training.

graphic for Morgan-Short et al, article 2012B


Second language syntactic processing shows increased native-like neural responses after months of no exposure to the language.

graphic for Lum et al, article 2012

LUM ET AL. 2012

Evidence that grammar relies on procedural memory in typically developing (TD) children, but on declarative memory in children with specific language impairment (SLI; that is, developmental language disorder); both rely on declarative memory for lexical abilities.

graphic for Babcock et al, article 2012


Complex linguistic forms can be stored or composed, as a function of multiple interacting factors, including regularity, sex, first vs. second language, and both length of residence and age of arrival in second language.

graphic for Hedenius et al, article 2011


Children with grammar impairments show consolidation problems at implicit sequence learning.