Is working memory domain-general or domain-specific?

Given the fundamental role of working memory (WM) in all domains of cognition, a central question has been whether WM is domain-general. However, the term 'domain-general' has been used in different, and sometimes misleading, ways. By reviewing recent evidence and biologically plausible models of WM, we show that the level of domain-generality varies substantially between three facets of WM: in terms of computations, WM is largely domain-general. In terms of neural correlates, it contains both domain-general and domain-specific elements. Finally, in terms of application, it is mostly domain-specific. This variance encourages a shift of focus towards uncovering domain-general computational principles and away from domain-general approaches to the analysis of individual differences and WM training, favoring newer perspectives, such as training-as-skill-learning.

The role of input vs. output phonological working memory in narrative production: Evidence from case series and case study approaches.

Separable input and output phonological working memory (WM) capacities have been proposed, with the input capacity supporting speech recognition and the output capacity supporting production. We examined the role of input vs. output phonological WM in narrative production, examining speech rate and pronoun ratio - two measures with prior evidence of a relation to phonological WM. For speech rate, a case series approach with individuals with aphasia found no significant independent contribution of input or output phonological WM capacity after controlling for single-word production. For pronoun ratio, there was some suggestion of a role for input phonological WM. Thus, neither finding supported a specific role for an output phonological buffer in speech production. In contrast, two cases demonstrating dissociations between input and output phonological WM capacities provided suggestive evidence of predicted differences in narrative production, though follow-up research is needed. Implications for case series vs. case study approaches are discussed.

Correction: Horne et al. White Matter Correlates of Domain-Specific Working Memory. Brain Sci. 2023, 13, 19.

In the original publication [...].

Corrigendum: Distinct neural substrates support phonological and orthographic working memory: implications for theories of working memory.

[This corrects the article DOI: 10.3389/fneur.2021.681141.].

Components of language processing and their long-term and working memory storage in the brain.

There is a consensus that the temporal lobes are involved in representing various types of information critical for language processing, including phonological (i.e., speech sound), semantic (meaning), and orthographic (spelling) representations. An important question is whether the same regions that represent our long-term knowledge of phonology, semantics, and orthography are used to support the maintenance of these types of information in working memory (WM) (for instance, maintaining semantic information during sentence comprehension), or whether regions outside the temporal lobes provide the neural basis for WM maintenance in these domains. This review focuses on the issue of whether temporal lobe regions support WM for phonological information, with a brief discussion of related findings in the semantic and orthographic domains. Across all three domains, evidence from lesion-symptom mapping and functional neuroimaging indicates that parietal or frontal regions are critical for supporting WM, with different regions supporting WM in the three domains. The distinct regions in different domains argue against these regions as playing a general attentional role. The findings imply an interaction between the temporal lobe regions housing the long-term memory representations in these domains and the frontal and parietal regions needed to maintain these representations over time.

Phonological Working Memory Representations in the Left Inferior Parietal Lobe in the Face of Distraction and Neural Stimulation.

The neural basis of phonological working memory (WM) was investigated through an examination of the effects of irrelevant speech distractors and disruptive neural stimulation from transcranial magnetic stimulation (TMS). Embedded processes models argue that the same regions involved in speech perception are used to support phonological WM whereas buffer models assume that a region separate from speech perception regions is used to support WM. Thus, according to the embedded processes approach but not the buffer approach, irrelevant speech and TMS to the speech perception region should disrupt the decoding of phonological WM representations. According to the buffer account, decoding of WM items should be possible in the buffer region despite distraction and should be disrupted with TMS to this region. Experiment 1 used fMRI and representational similarity analyses (RSA) with a delayed recognition memory paradigm using nonword stimuli. Results showed that decoding of memory items in the speech perception regions (superior temporal gyrus, STG) was possible in the absence of distractors. However, the decoding evidence in the left STG was susceptible to interference from distractors presented during the delay period whereas decoding in the proposed buffer region (supramarginal gyrus, SMG) persisted. Experiment 2 examined the causal roles of the speech processing region and the buffer region in phonological WM performance using TMS. TMS to the SMG during the early delay period caused a disruption in recognition performance for the memory nonwords, whereas stimulations at the STG and an occipital control region did not affect WM performance. Taken together, results from the two experiments are consistent with predictions of a buffer model of phonological WM, pointing to a critical role of the left SMG in maintaining phonological representations.

Semantic Working Memory Predicts Sentence Comprehension Performance: A Case Series Approach.

Sentence comprehension involves maintaining and continuously integrating linguistic information and, thus, makes demands on working memory (WM). Past research has demonstrated that semantic WM, but not phonological WM, is critical for integrating word meanings across some distance and resolving semantic interference in sentence comprehension. Here, we examined the relation between phonological and semantic WM and the comprehension of center-embedded relative clause sentences, often argued to make heavy demands on WM. Additionally, we examined the relation between phonological and semantic WM and the comprehension of transitive and dative active and passive sentences, which may also draw on WM resources depending on the number of propositions that must be maintained and the difficulty of processing passive clauses. In a large sample of individuals with aphasia (N = 56), we assessed whether comprehension performance on more complex vs. simpler active-passive or embedded relative clause sentences would be predicted by semantic but not phonological WM when controlling for single word comprehension. For performance on the active-passive comprehension task, we found that semantic WM, but not phonological WM, predicted comprehension of dative sentences when controlling for comprehension of transitive sentences. We also found that phonological WM, but not semantic WM, predicted mean comprehension for reversible active-passive sentences when controlling for trials with lexical distractors. On the relative clause comprehension task, consistent with prior results, we found that semantic WM, but not phonological WM, predicted comprehension of object relative clause sentences and relative clause sentences with a passive construction. However, both phonological WM and semantic WM predicted mean comprehension across all relative clause types for reversible trials when controlling for trials with lexical distractors. While we found evidence of semantic WM's role in comprehension, we also observed unpredicted relations between phonological WM and comprehension in some conditions. Post-hoc analyses provided preliminary evidence that phonological WM maintains a backup phonological representation of the sentence that may be accessed when sentence comprehension processing is less efficient. Future work should investigate possible roles that phonological WM may play across sentence types.

Contributions of semantic and phonological working memory to narrative language independent of single word production: Evidence from acute stroke.

ABSTRACTNeuropsychological case studies have provided evidence that individuals with semantic, but not phonological, working memory (WM) deficits have difficulty producing phrases containing several content words. These findings supported the claim of a phrasal scope of planning at the grammatical formulation stage of production, where semantic WM supports the maintenance of lexical-semantic representations as they are inserted into slots in phrasal constituents. Recent narrative production results for individuals at the acute stage of stroke supported the role for semantic WM in phrasal elaboration while suggesting a role for phonological WM at a subsequent phonological encoding stage in supporting fluent, rapid speech. In the present study, we employed a larger participant sample while controlling for single word production abilities at semantic and phonological levels. Results confirmed the relations between semantic WM and phrasal elaboration whereas the relation between phonological WM and speech rate was eliminated. There was, however, evidence that both impaired phonological retrieval and restricted phonological WM were related to the tendency to produce short, highly frequent words such as pronouns rather than longer, less frequent words. Future research is needed to address whether the results relating to phonological WM depend on damage specific to an output rather than an input phonological capacity.

Effects of semantic diversity and word frequency on single word processing.

Some research suggests that semantic diversity (SemD), a measure of the variability of contexts in which a word appears, plays an important role in language processing, determining the availability of word representations (e.g., Adelman et al., 2006) and causing task-specific benefits or detriments to performance (e.g., Hoffman & Woollams, 2015). Some researchers have claimed that word frequency has no effect once such diversity measures are taken into account (Adelman et al., 2006). Taking advantage of the power of five large-scale databases, we investigated the effects of SemD, word frequency, and their interaction in five tasks, including word reading, lexical and concreteness decision, object picture naming, and word repetition. We found: (a) word frequency and SemD effects were consistently distinct; (b) effects of SemD were facilitatory in nearly all tasks, but inhibitory effects were also found; contrary to existing claims, we conclude that inhibitory SemD effects do not necessarily imply semantic selection requirements; (c) the presence of SemD effects minimally influenced the size of frequency effects when SemD was left uncontrolled, suggesting that SemD does not explain absent frequency effects in the patient literature; and (d) word frequency and SemD only interact in the largest data sets. Results are discussed in the context of rational models of memory (Anderson & Milson, 1989; Anderson & Schooler, 1991) and the Controlled Semantic Cognition framework (Lambon Ralph et al., 2017). (PsycInfo Database Record (c) 2022 APA, all rights reserved).

White Matter Correlates of Domain-Specific Working Memory.

Prior evidence suggests domain-specific working memory (WM) buffers for maintaining phonological (i.e., speech sound) and semantic (i.e., meaning) information. The phonological WM buffer's proposed location is in the left supramarginal gyrus (SMG), whereas semantic WM has been related to the left inferior frontal gyrus (IFG), the middle frontal gyrus (MFG), and the angular gyrus (AG). However, less is known about the white matter correlates of phonological and semantic WM. We tested 45 individuals with left hemisphere brain damage on single word processing, phonological WM, and semantic WM tasks and obtained T1 and diffusion weighted neuroimaging. Virtual dissections were performed for each participants' arcuate fasciculus (AF), inferior fronto-occipital fasciculus (IFOF), inferior longitudinal fasciculus (ILF), middle longitudinal fasciculus (MLF), and uncinate fasciculus (UF), which connect the proposed domain-specific WM buffers with perceptual or processing regions. The results showed that the left ILF, MLF, IFOF, and the direct segment of the AF were related to semantic WM performance. Phonological WM was related to both the left ILF and the MLF. This work informs our understanding of the white matter correlates of WM, especially semantic WM, which has not previously been investigated. In addition, this work helps to adjudicate between theories of verbal WM, providing some evidence for separate pathways supporting phonological and semantic WM.

The Critical Role of Semantic Working Memory in Language Comprehension and Production.

Although research on the role of verbal working memory (WM) in language processing has focused on phonological maintenance, considerable evidence indicates that maintenance of semantic information plays a more critical role. This paper reviews studies of brain damaged and healthy individuals, demonstrating the contribution of semantic WM to language processing. On the sentence comprehension side, semantic WM supports the retention of individual word meanings prior to their integration. It also serves to maintain semantic information in an activated state such that semantic interference between constituents in a sentence can be resolved. Phonological WM does not appear to contribute to either of these functions, though it contributes to verbatim sentence recall. On the production side, evidence points to the phrase as the minimal scope of advance planning in sentence formulation, with semantic WM supporting the representation of the meanings of content words within a phrase. Planning at the phonological level appears to have a very limited scope, making few demands on phonological WM. These findings imply that treatment of semantic but not phonological WM deficits should lead to improved sentence comprehension and production and preliminary findings (Harris, Olson, & Humphreys, 2014) support that view.

Distinct Neural Substrates Support Phonological and Orthographic Working Memory: Implications for Theories of Working Memory.

Prior behavioral and neuroimaging evidence supports a separation between working memory capacities in the phonological and orthographic domains. Although these data indicate distinct buffers for orthographic and phonological information, prior neural evidence does indicate that nearby left inferior parietal regions support both of these working memory capacities. Given that no study has directly compared their neural substrates based on data from the same individuals, it is possible that there is a common left inferior parietal region shared by both working memory capacities. In fact, those endorsing an embedded processes account of working memory might suggest that parietal involvement reflects a domain-general attentional system that directs attention to long-term memory representations in the two domains, implying that the same neural region supports the two capacities. Thus, in this work, a multivariate lesion-symptom mapping approach was used to assess the neural basis of phonological and orthographic working memory using behavioral and lesion data from the same set of 37 individuals. The results showed a separation of the neural substrates, with regions in the angular gyrus supporting orthographic working memory and with regions primarily in the supramarginal gyrus supporting phonological working memory. The results thus argue against the parietal involvement as supporting a domain-general attentional mechanism and support a domain-specific buffer account of working memory.

Working Memory Capacities Neurally Dissociate: Evidence from Acute Stroke.

Substantial behavioral evidence implies the existence of separable working memory (WM) components for maintaining phonological and semantic information. In contrast, only a few studies have addressed the neural basis of phonological versus semantic WM using functional neuroimaging and none has used a lesion-symptom mapping (LSM) approach. Here, we address this gap, reporting a multivariate LSM study of phonological and semantic WM for 94 individuals at the acute stage of left hemisphere stroke. Testing at the acute stage avoids issues of brain reorganization and the adoption of patient strategies for task performance. The LSM analyses for each WM component controlled for the other WM component and semantic and phonological knowledge at the single word level. For phonological WM, the regions uncovered included the supramarginal gyrus, argued to be the site of phonological storage, and several cortical and subcortical regions plausibly related to inner rehearsal. For semantic WM, inferior frontal regions and the angular gyrus were uncovered. The findings thus provide converging evidence for separable systems for phonological and semantic WM that are distinguished from the systems supporting long-term knowledge representations in those domains.

Maintaining verbal short-term memory representations in non-perceptual parietal regions.

Buffer accounts of verbal short-term memory (STM) assume dedicated buffers for maintaining different types of information (e.g., phonological, visual) whereas embedded processes accounts argue against the existence of buffers and claim that STM consists of the activated portion of long-term memory (LTM). We addressed this debate by determining whether STM recruits the same neural substrate as LTM, or whether additional regions are involved in short-term storage. Using fMRI with representational similarity analysis (RSA), we examined the representational correspondence of multi-voxel neural activation patterns with the theoretical predictions for the maintenance of both phonological and semantic codes in STM. We found that during the delay period of a phonological STM task, phonological representations could be decoded in the left supramarginal gyrus (SMG) but not the superior temporal gyrus (STG), a speech processing region, for word stimuli. Whereas the pattern in the SMG was specific to phonology, a different region in the left angular gyrus showed RSA decoding evidence for the retention of either phonological or semantic codes, depending on the task context. Taken together, the results provide clear support for a dedicated buffer account of phonological STM, although evidence for a semantic buffer is equivocal.

The role of left vs. right superior temporal gyrus in speech perception: An fMRI-guided TMS study.

Debate continues regarding the necessary role of right superior temporal gyrus (STG) regions in sublexical speech perception given the bilateral STG activation often observed in fMRI studies. To evaluate the causal roles, TMS pulses were delivered to inhibit and disrupt neuronal activity at the left and right STG regions during a nonword discrimination task based on peak activations from a blocked fMRI paradigm assessing speech vs. nonspeech perception (N = 20). Relative to a control region located in the posterior occipital lobe, TMS to the left anterior STG (laSTG) led to significantly worse accuracy, whereas TMS to the left posterior STG (lpSTG) and right anterior STG (raSTG) did not. Although the disruption from TMS was significantly greater for the laSTG than for raSTG, the difference in accuracy between the laSTG and lpSTG did not reach significance. The results argue for a causal role of the laSTG but not raSTG in speech perception. Further research is needed to establish the source of the differences between the laSTG and lpSTG.

Dissociation between frontal and temporal-parietal contributions to connected speech in acute stroke.

Humans are uniquely able to retrieve and combine words into syntactic structure to produce connected speech. Previous identification of focal brain regions necessary for production focused primarily on associations with the content produced by speakers with chronic stroke, where function may have shifted to other regions after reorganization occurred. Here, we relate patterns of brain damage with deficits to the content and structure of spontaneous connected speech in 52 speakers during the acute stage of a left hemisphere stroke. Multivariate lesion behaviour mapping demonstrated that damage to temporal-parietal regions impacted the ability to retrieve words and produce them within increasingly complex combinations. Damage primarily to inferior frontal cortex affected the production of syntactically accurate structure. In contrast to previous work, functional-anatomical dissociations did not depend on lesion size likely because acute lesions were smaller than typically found in chronic stroke. These results are consistent with predictions from theoretical models based primarily on evidence from language comprehension and highlight the importance of investigating individual differences in brain-language relationships in speakers with acute stroke.

Evaluating the distinction between semantic knowledge and semantic access: Evidence from semantic dementia and comprehension-impaired stroke aphasia.

Theories of semantic memory based on neuropsychological findings have posited a distinction between stored semantic representations and the mechanisms used to access and manipulate them (e.g., Lambon Ralph, Jefferies, Patterson, & Rogers, 2017; Warrington & Cipolotti, 1996). The most recent instantiation of this view, the controlled semantic cognition theory (Lambon Ralph et al., 2017), is supported by findings suggesting that multimodal (i.e., both verbal and nonverbal) semantic deficits may result from qualitatively different impairments: on the one hand, damage to a semantic access mechanism related to executive control, which is observed in semantic aphasia (SA), and on the other, damage to semantic representations, which is observed in semantic dementia (SD) (Jefferies & Lambon Ralph, 2006). In this study we compared SA and SD patients on several phenomena previously used to support these distinctions. Contrary to the prior results, we found that (1) overall, cross-task consistency was equivalent for the two groups; (2) neither patient group showed consistency driven by item identity across different semantic tasks; (3) correlations among task performance were not obviously driven by the semantic control demands of different tasks; (4) both groups showed executive function deficits; and (5) both groups showed strong effects of distractor interference in a synonym judgment task. Furthermore, we investigated the components of executive ability that could underlie semantic control deficits by correlating performance on updating, shifting, and inhibition tasks with performance on tasks testing semantic abilities. We found that updating was related to semantic processing generally, whereas shifting and inhibition were not. These results also suggest that complex executive function tasks relate to semantic tasks through their shared relationship with language abilities. Overall, evidence from SA and SD patients does not differentiate representations and access mechanisms in the semantic system, as has previously been suggested. Implications for the storage-access distinction are discussed.

Lexical processing depends on sublexical processing: Evidence from the visual world paradigm and aphasia.

Some early studies of people with aphasia reported strikingly better performance on lexical than on sublexical speech perception tasks. These findings challenged the claim that lexical processing depends on sublexical processing and suggested that acoustic information could be mapped directly to lexical representations. However, Dial and Martin (Neuropsychologia 96: 192-212, 2017) argued that these studies failed to match the discriminability of targets and distractors for the sublexical and lexical stimuli and showed that when using closely matched tasks with natural speech tokens, no patient performed substantially better at the lexical than at the sublexical processing task. In the current study, we sought to provide converging evidence for the dependence of lexical on sublexical processing by examining the perception of synthetic speech stimuli varied on a voice-onset time continuum using eye-tracking methodology, which is sensitive to online speech perception processes. Eight individuals with aphasia and ten age-matched controls completed two visual world paradigm tasks: phoneme (sublexical) and word (lexical) identification. For both identification and eye-movement data, strong correlations were observed between the sublexical and lexical tasks. Critically, no patient within the control range on the lexical task was impaired on the sublexical task. Overall, the current study supports the claim that lexical processing depends on sublexical processing. Implications for inferring deficits in people with aphasia and the use of sublexical tasks to assess sublexical processing are also discussed.

Non-perceptual Regions in the Left Inferior Parietal Lobe Support Phonological Short-term Memory: Evidence for a Buffer Account?

Buffer versus embedded processes accounts of short-term memory (STM) for phonological information were addressed by testing subjects' perception and memory for speech and non-speech auditory stimuli. Univariate and multivariate (MVPA) approaches were used to assess whether brain regions recruited in recognizing speech were involved in maintaining speech representations over a delay. As expected, a left superior temporal region was found to support speech perception. However, contrary to the embedded processes approach, this region failed to show a load effect, or any sustained activation, during a maintenance delay. Moreover, MVPA decoding during the maintenance stage was unsuccessful in this region by a perception classifier or an encoding classifier. In contrast, the left supramarginal gyrus showed both sustained activation and a load effect. Using MVPA, stimulus decoding was successful during the delay period. In addition, a functional connectivity analysis showed that, as memory load increased, the left temporal lobe involved in perception became more strongly connected with the parietal region involved in maintenance. Taken together, the findings provide greater support for a buffer than embedded processes account of phonological STM.

Independent contributions of semantic and phonological working memory to spontaneous speech in acute stroke.

Patients with left hemisphere stroke often have language deficits which impair their ability to produce phrases and sentences. One possible source of these speech impairments is the disruption of verbal working memory (WM). Single-case studies of chronic stroke have suggested the existence of a WM capacity specific to maintaining semantic information that is critical for preparing multiple words in phrases prior to speech onset (Freedman, Martin, & Biegler, 2004; Martin & Freedman, 2001; Martin & He, 2004; Martin, Miller, & Vu, 2004). The current study tested this hypothesis by examining spontaneous narrative language production and working memory capacities in a large sample of individuals at the acute stage of stroke (N = 36), prior to the reorganization of function or strategy development. Here we show using a multiple regression approach that patients' semantic but not phonological WM capacity had an independent contribution in predicting phrasal elaboration and increasing utterance length whereas patients' phonological but not semantic WM capacity had an independent contribution in predicting a more rapid speech rate. Importantly, neither WM capacity independently predicted grammatical abilities in speech, implying that the other relations did not result from overall severity. These results indicate that separable semantic and phonological WM components exist that support different aspects of narrative speech. To our knowledge, this is the first study to examine spontaneous speech in a large group of acute stroke patients demonstrating a critical relationship between working memory and the ability to produce more words in phrases and longer utterances.