Individual differences in human gaze behavior generalize from faces to objects.

Individuals differ in where they fixate on a face, with some looking closer to the eyes while others prefer the mouth region. These individual biases are highly robust, generalize from the lab to the outside world, and have been associated with social cognition and associated disorders. However, it is unclear, whether these biases are specific to faces or influenced by domain-general mechanisms of vision. Here, we juxtaposed these hypotheses by testing whether individual face fixation biases generalize to inanimate objects. We analyzed >1.8 million fixations toward faces and objects in complex natural scenes from 405 participants tested in multiple labs. Consistent interindividual differences in fixation positions were highly inter-correlated across faces and objects in all samples. Observers who fixated closer to the eye region also fixated higher on inanimate objects and vice versa. Furthermore, the inter-individual spread of fixation positions scaled with target size in precisely the same, non-linear manner for faces and objects. These findings contradict a purely domain-specific account of individual face gaze. Instead, they suggest significant domain-general contributions to the individual way we look at faces, a finding with potential relevance for basic vision, face perception, social cognition, and associated clinical conditions.

Task learning is subserved by a domain-general brain network.

One of the most important human faculties is the ability to acquire not just new memories but the capacity to perform entirely new tasks. However, little is known about the brain mechanisms underlying the learning of novel tasks. Specifically, it is unclear to what extent learning of different tasks depends on domain-general and/or domain-specific brain mechanisms. Here human subjects (n = 45) learned to perform 6 new tasks while undergoing functional MRI. The different tasks required the engagement of perceptual, motor, and various cognitive processes related to attention, expectation, speed-accuracy tradeoff, and metacognition. We found that a bilateral frontoparietal network was more active during the initial compared with the later stages of task learning, and that this effect was stronger for task variants requiring more new learning. Critically, the same frontoparietal network was engaged by all 6 tasks, demonstrating its domain generality. Finally, although task learning decreased the overall activity in the frontoparietal network, it increased the connectivity strength between the different nodes of that network. These results demonstrate the existence of a domain-general brain network whose activity and connectivity reflect learning for a variety of new tasks, and thus may underlie the human capacity for acquiring new abilities.

Distinct and common mechanisms of cross-model semantic conflict and response conflict in an auditory relevant task.

The mechanisms of semantic conflict and response conflict in the Stroop task have mainly been investigated in the visual modality. However, the understanding of these mechanisms in cross-modal modalities remains limited. In this electroencephalography (EEG) study, an audiovisual 2-1 mapping Stroop task was utilized to investigate whether distinct and/or common neural mechanisms underlie cross-modal semantic conflict and response conflict. The response time data showed significant effects on both cross-modal semantic and response conflicts. Interestingly, the magnitude of semantic conflict was found to be smaller in the fast response time bins than in the slow response time bins, whereas no such difference was observed for response conflict. The EEG data demonstrated that cross-modal semantic conflict specifically increased the N450 amplitude. However, cross-modal response conflict specifically enhanced theta band power and theta phase synchronization between the medial frontal cortex (MFC) and lateral prefrontal electrodes as well as between the MFC and motor electrodes. In addition, both cross-modal semantic conflict and response conflict led to a decrease in P3 amplitude. Taken together, these findings provide cross-modal evidence for domain-specific mechanism in conflict detection and suggest both domain-specific and domain-general mechanisms exist in conflict resolution.

Comprehensive investigation of predictive processing: A cross- and within-cognitive domains fMRI meta-analytic approach.

Predictive processing (PP) stands as a predominant theoretical framework in neuroscience. While some efforts have been made to frame PP within a cognitive domain-general network perspective, suggesting the existence of a "prediction network," these studies have primarily focused on specific cognitive domains or functions. The question of whether a domain-general predictive network that encompasses all well-established cognitive domains exists remains unanswered. The present meta-analysis aims to address this gap by testing the hypothesis that PP relies on a large-scale network spanning across cognitive domains, supporting PP as a unified account toward a more integrated approach to neuroscience. The Activation Likelihood Estimation meta-analytic approach was employed, along with Meta-Analytic Connectivity Mapping, conjunction analysis, and behavioral decoding techniques. The analyses focused on prediction incongruency and prediction congruency, two conditions likely reflective of core phenomena of PP. Additionally, the analysis focused on a prediction phenomena-independent dimension, regardless of prediction incongruency and congruency. These analyses were first applied to each cognitive domain considered (cognitive control, attention, motor, language, social cognition). Then, all cognitive domains were collapsed into a single, cross-domain dimension, encompassing a total of 252 experiments. Results pertaining to prediction incongruency rely on a defined network across cognitive domains, while prediction congruency results exhibited less overall activation and slightly more variability across cognitive domains. The converging patterns of activation across prediction phenomena and cognitive domains highlight the role of several brain hubs unfolding within an organized large-scale network (Dynamic Prediction Network), mainly encompassing bilateral insula, frontal gyri, claustrum, parietal lobules, and temporal gyri. Additionally, the crucial role played at a cross-domain, multimodal level by the anterior insula, as evidenced by the conjunction and Meta-Analytic Connectivity Mapping analyses, places it as the major hub of the Dynamic Prediction Network. Results support the hypothesis that PP relies on a domain-general, large-scale network within whose regions PP units are likely to operate, depending on the context and environmental demands. The wide array of regions within the Dynamic Prediction Network seamlessly integrate context- and stimulus-dependent predictive computations, thereby contributing to the adaptive updating of the brain's models of the inner and external world.

Semantic-specific and domain-general mechanisms for integration and update of contextual information.

Recent research has highlighted the importance of domain-general processes and brain regions for language and semantic cognition. Yet, this has been mainly observed in executively demanding tasks, leaving open the question of the contribution of domain-general processes to natural language and semantic cognition. Using fMRI, we investigated whether neural processes reflecting context integration and context update-two key aspects of naturalistic language and semantic processing-are domain-specific versus domain-general. Thus, we compared neural responses during the integration of contextual information across semantic and non-semantic tasks. Whole-brain results revealed both shared (left posterior-dorsal inferior frontal gyrus, left posterior inferior temporal gyrus, and left dorsal angular gyrus/intraparietal sulcus) and distinct (left anterior-ventral inferior frontal gyrus, left anterior ventral angular gyrus, left posterior middle temporal gyrus for semantic control only) regions involved in context integration and update. Furthermore, data-driven functional connectivity analysis clustered domain-specific versus domain-general brain regions into distinct but interacting functional neural networks. These results provide a first characterisation of the neural processes required for context-dependent integration during language processing along the domain-specificity dimension, and at the same time, they bring new insights into the role of left posterior lateral temporal cortex and left angular gyrus for semantic cognition.

Recent developments in parrot cognition: a quadrennial update.

Psittacines, along with corvids, are commonly referred to as 'feathered apes' due to their advanced cognitive abilities. Until rather recently, the research effort on parrot cognition was lagging behind that on corvids, however current developments show that the number of parrot studies is steadily increasing. In 2018, M. L. Lambert et al. provided a comprehensive review on the status of the most important work done so far in parrot and corvid cognition. Nevertheless, only a little more than 4 years after this publication, more than 50 new parrot studies have been published, some of them chartering completely new territory. On the 25th anniversary of Animal Cognition we think this warrants a detailed review of parrot cognition research over the last 4 years. We aim to capture recent developments and current trends in this rapidly expanding and diversifying field.

Individual consistency in the learning abilities of honey bees: cognitive specialization within sensory and reinforcement modalities.

The question of whether individuals perform consistently across a variety of cognitive tasks is relevant for studies of comparative cognition. The honey bee (Apis mellifera) is an appropriate model to study cognitive consistency as its learning can be studied in multiple elemental and non-elemental learning tasks. We took advantage of this possibility and studied if the ability of honey bees to learn a simple discrimination correlates with their ability to solve two tasks of higher complexity, reversal learning and negative patterning. We performed four experiments in which we varied the sensory modality of the stimuli (visual or olfactory) and the type (Pavlovian or operant) and complexity (elemental or non-elemental) of conditioning to examine if stable correlated performances could be observed across experiments. Across all experiments, an individual's proficiency to learn the simple discrimination task was positively and significantly correlated with performance in both reversal learning and negative patterning, while the performances in reversal learning and negative patterning were positively, yet not significantly correlated. These results suggest that correlated performances across learning paradigms represent a distinct cognitive characteristic of bees. Further research is necessary to examine if individual cognitive consistency can be found in other insect species as a common characteristic of insect brains.

Global connectivity fingerprints predict the domain generality of multiple-demand regions.

A set of distributed cognitive control networks are known to contribute to diverse cognitive demands, yet it is unclear how these networks gain this domain-general capacity. We hypothesized that this capacity is largely due to the particular organization of the human brain's intrinsic network architecture. Specifically, we tested the possibility that each brain region's domain generality is reflected in its level of global (hub-like) intrinsic connectivity as well as its particular global connectivity pattern ("connectivity fingerprint"). Consistent with prior work, we found that cognitive control networks exhibited domain generality as they represented diverse task context information covering sensory, motor response, and logic rule domains. Supporting our hypothesis, we found that the level of global intrinsic connectivity (estimated with resting-state functional magnetic resonance imaging [fMRI]) was correlated with domain generality during tasks. Further, using a novel information fingerprint mapping approach, we found that each cognitive control region's unique rule response profile("information fingerprint") could be predicted based on its unique intrinsic connectivity fingerprint and the information content in regions outside cognitive control networks. Together, these results suggest that the human brain's intrinsic network architecture supports its ability to represent diverse cognitive task information largely via the location of multiple-demand regions within the brain's global network organization.

Precise Topology of Adjacent Domain-General and Sensory-Biased Regions in the Human Brain.

Recent functional MRI studies identified sensory-biased regions across much of the association cortices and cerebellum. However, their anatomical relationship to multiple-demand (MD) regions, characterized as domain-general due to their coactivation during multiple cognitive demands, remains unclear. For a better anatomical delineation, we used multimodal MRI techniques of the Human Connectome Project to scan subjects performing visual and auditory versions of a working memory (WM) task. The contrast between hard and easy WM showed strong domain generality, with essentially identical patterns of cortical, subcortical, and cerebellar MD activity for visual and auditory materials. In contrast, modality preferences were shown by contrasting easy WM with baseline; most MD regions showed visual preference while immediately adjacent to cortical MD regions, there were interleaved regions of both visual and auditory preference. The results may exemplify a general motif whereby domain-specific regions feed information into and out of an adjacent, integrative MD core.

Environmental susceptibility for all: A data-driven approach suggests individual differences in domain-general and domain-specific patterns of environmental susceptibility.

How we are influenced by our environment is a fundamental question in developmental science. Theories and empirical research have claimed that some individuals are susceptible to environmental influences and others are much less susceptible. The present study addressed four questions: (1) Is environmental susceptibility a continuous or categorical construct? (2) Is environmental susceptibility unidimensional (i.e., domain general) or multidimensional (i.e., domain specific)? (3) Are there genetic contributions to individual differences in environmental susceptibility? (4) What are the temperamental characteristics of different environmental susceptibility patterns? We used child- and mother-report data from a sample of 11-year-old twins (N = 1,507) and applied a novel data-driven approach to assess an environmental susceptibility space, based on simultaneous associations between multiple environmental exposures (18 measures relating to parenting, parent, peer, and twin relationships) and developmental outcomes (10 measures relating to empathy, prosocial behavior, aggression, and self-esteem). The results suggest that the environmental susceptibility space we assessed is better conceptualized as continuous and multidimensional. Different children showed susceptibility to different contexts and variation in domain-general versus domain-specific patterns. A comparison of distances between monozygotic and dizygotic twins within the space demonstrated genetic contributions. Finally, susceptibility patterns could not be differentiated based on a specific temperament trait, but rather related to temperament profiles.

Beyond orchids and dandelions: Susceptibility to environmental influences is not bimodal.

This study focused on generality versus specificity of susceptibility of effects of eight family and child-care exposures measured between 3 and 54 months of age (e.g., sensitive parenting, child-care quality) on five child development outcomes assessed at age 4.5 years (e.g. behavior problems, preacademic skill), using data from The National Institute of Child Health and Human Development (NICHD) Study of Early Child Care and Youth Development (n = 1,364, boys = 705; White = 1,097, Black = 176, other = 91), while applying a novel influence-statistics method. Results indicated that susceptibility across the environment-predictor:child-outcome associations is normally rather than bimodally (i.e., orchid-dandelion) distributed. Analysis of susceptibility documents both domain generality and specificity of developmental plasticity, with effect sizes proving small in the former case. As predicted, children who as infants had difficult temperaments or who scored higher on a polygenic-plasticity score (serotonin-transporter-linked promoter region [5-HTTLPR], dopamine receptor D4 [DRD4], brain-derived neurotrophic factor [BDNF]) proved somewhat more susceptible to some of the environmental effects investigated. Results lead to the recommendation that two-types-of-individuals vis-a-vis susceptibility to environmental influences be questioned and general-trait conceptions of susceptibility be further investigated.

Working memory training improves children's syntactic ability but not vice versa: A randomized control trial.

We tested several hypotheses about the relation between syntax and working memory (WM). In a pretest/posttest randomized control trial, 104 native Cuban Spanish-speaking children (Mage = 7 years 2 months; 54 girls) took part in syntax training in their first language, syntax training in their second language, WM training, or no training (control). Compared with the control, children in the training conditions showed cognitive transfer from WM to syntax but not from syntax to WM. The result was most striking in the case of their first language, where WM training was as effective as language training in boosting syntactic performance. As well as establishing cognitive transfer at the group level, we also found that individual differences in WM performance, both at baseline and in training, predicted the extent to which children's syntax improved. The directionality of transfer, the group-level and individual-level results, established in the context of a randomized control design, all point to a strong causal role for domain-general cognition in the processes of language acquisition.

Differential susceptibility 2.0: Are the same children affected by different experiences and exposures?

Differential susceptibility theory stipulates that some children are more susceptible than others to both supportive and adverse developmental experiences/exposures. What remains unclear is whether the same individuals are most affected by different exposures (i.e., domain general vs. specific). We address this issue empirically for the first time using, for illustrative and proof-of-principle purposes, a novel influence-statistics' method with data from the National Institute of Child Health and Human Development (NICHD) Study of Early Child Care. Results indicated that previously documented effects of greater quality of care on enhanced pre-academic skills and greater quantity of care on more behavior problems apply mostly to different children. Analyses validating the new method indicated, as predicted, that (a) the quantity-of-care effect applied principally to children from more socioeconomically advantaged families and that (b) being highly susceptible to both, one or neither childcare effect varied as a function of a three-gene, polygenic-plasticity score (serotonin transporter linked polymorphic region [5-HTTLPR], dopamine receptor D4 [DRD4], brain-derived neurotrophic factor [BDNF]) in a dose-response manner (i.e., 2>1>0). While domain-specific findings involving child-care effects cannot be generalized to other environmental influences, the influence-statistics' approach appears well suited for investigating the generality-specificity of environment effects, that is, of "differential, differential susceptibility."

The Domain-General Multiple Demand (MD) Network Does Not Support Core Aspects of Language Comprehension: A Large-Scale fMRI Investigation.

Aside from the language-selective left-lateralized frontotemporal network, language comprehension sometimes recruits a domain-general bilateral frontoparietal network implicated in executive functions: the multiple demand (MD) network. However, the nature of the MD network's contributions to language comprehension remains debated. To illuminate the role of this network in language processing in humans, we conducted a large-scale fMRI investigation using data from 30 diverse word and sentence comprehension experiments (481 unique participants [female and male], 678 scanning sessions). In line with prior findings, the MD network was active during many language tasks. Moreover, similar to the language-selective network, which is robustly lateralized to the left hemisphere, these responses were stronger in the left-hemisphere MD regions. However, in contrast with the language-selective network, the MD network responded more strongly (1) to lists of unconnected words than to sentences, and (2) in paradigms with an explicit task compared with passive comprehension paradigms. Indeed, many passive comprehension tasks failed to elicit a response above the fixation baseline in the MD network, in contrast to strong responses in the language-selective network. Together, these results argue against a role for the MD network in core aspects of sentence comprehension, such as inhibiting irrelevant meanings or parses, keeping intermediate representations active in working memory, or predicting upcoming words or structures. These results align with recent evidence of relatively poor tracking of the linguistic signal by the MD regions during naturalistic comprehension, and instead suggest that the MD network's engagement during language processing reflects effort associated with extraneous task demands.SIGNIFICANCE STATEMENT Domain-general executive processes, such as working memory and cognitive control, have long been implicated in language comprehension, including in neuroimaging studies that have reported activation in domain-general multiple demand (MD) regions for linguistic manipulations. However, much prior evidence has come from paradigms where language interpretation is accompanied by extraneous tasks. Using a large fMRI dataset (30 experiments/481 participants/678 sessions), we demonstrate that MD regions are engaged during language comprehension in the presence of task demands, but not during passive reading/listening, conditions that strongly activate the frontotemporal language network. These results present a fundamental challenge to proposals whereby linguistic computations, such as inhibiting irrelevant meanings, keeping representations active in working memory, or predicting upcoming elements, draw on domain-general executive resources.

Segregating domain-general from emotional context-specific inhibitory control systems - ventral striatum and orbitofrontal cortex serve as emotion-cognition integration hubs.

Inhibitory control hierarchically regulates cognitive and emotional systems in the service of adaptive goal-directed behavior across changing task demands and environments. While previous studies convergently determined the contribution of prefrontal-striatal systems to general inhibitory control, findings on the specific circuits that mediate emotional context-specific impact on inhibitory control remained inconclusive. Against this background we combined an evaluated emotional Go/No Go task with fMRI in a large cohort of subjects (N=250) to segregate brain systems and circuits that mediate domain-general from emotion-specific inhibitory control. Particularly during a positive emotional context, behavioral results showed a lower accuracy for No Go trials and a faster response time for Go trials. While the dorsal striatum and lateral frontal regions were involved in inhibitory control irrespective of emotional context, activity in the ventral striatum (VS) and medial orbitofrontal cortex (mOFC) varied as a function of emotional context. On the voxel-wise whole-brain network level, limbic and striatal systems generally exhibited highest changes in global brain connectivity during inhibitory control, while global brain connectivity of the left mOFC was less decreased during emotional contexts. Functional connectivity analyses moreover revealed that negative coupling between the VS with inferior frontal gyrus (IFG)/insula and mOFC varied as a function of emotional context. Together these findings indicate separable domain- general as well as emotional context-specific inhibitory brain systems which specifically encompass the VS and its connections with frontal regions.

Frontoparietal pattern similarity analyses of cognitive control in monozygotic twins.

The ability to flexibly adapt thoughts and actions in a goal-directed manner appears to rely on cognitive control mechanisms that are strongly impacted by individual differences. A powerful research strategy for investigating the nature of individual variation is to study monozygotic (identical) twins. Evidence of twin effects have been observed in prior behavioral and neuroimaging studies, yet within the domain of cognitive control, it remains to be demonstrated that the neural underpinnings of such effects are specific and reliable. Here, we utilize a multi-task, within-subjects event-related neuroimaging design with functional magnetic resonance imaging to investigate twin effects through multivariate pattern similarity analyses. We focus on fronto-parietal brain regions exhibiting consistently increased activation associated with cognitive control demands across four task domains: selective attention, context processing, multi-tasking, and working memory. Healthy young adult monozygotic twin pairs exhibited increased similarity of within- and cross-task activation patterns in these fronto-parietal regions, relative to unrelated pairs. Twin activation pattern similarity effects were clearest under high control demands, were not present in a set of task-unrelated parcels or due to anatomic similarity, and were primarily observed during the within-trial timepoints in which the control demands peaked. Together, these results indicate that twin similarity in the neural representation of cognitive control may be domain-general but also functionally and temporally specific in relation to the level of control demand. The findings suggest a genetic and/or environmental basis for individual variation in cognitive control function, and highlight the potential of twin-based neuroimaging designs for exploring heritability questions within this domain.

A case-study of language-specific executive disorder.

Executive control is recruited for language processing, particularly in complex linguistic tasks. Although the issue of the existence of an executive control specific to language is still an open issue, there is much evidence that executively-demanding language tasks rely on domain-general rather than language-specific executive resources. Here, we addressed this issue by assessing verbal and non-verbal executive capacities in LG, an aphasic patient after a stroke. First, we showed that LG's performance was spared in all non-verbal tasks regardless of the executive demands. Second, by contrasting conditions of high and low executive demand in verbal tasks, we showed that LG was only impaired in verbal task with high executive demand. The performance dissociation between low and high executive demand conditions in the verbal domain, not observed in the non-verbal domain, shows that verbal executive control partly dissociates from non-verbal executive control. This language-specific executive disorder suggests that some executive processes might be language-specific.

Neural substrates of the executive function construct, age-related changes, and task materials in adolescents and adults: ALE meta-analyses of 408 fMRI studies.

To explore the neural substrates of executive function (EF), we conducted an activation likelihood estimation meta-analysis of 408 functional magnetic resonance imaging studies (9639 participants, 7587 activation foci, 518 experimental contrasts) covering three fundamental EF subcomponents: inhibition, switching, and working memory. Our results found that activation common to all three EF subcomponents converged in the multiple-demand network across adolescence and adulthood. The function of EF with the multiple-demand network involved, especially for the prefrontal cortex and the parietal regions, could not be mature until adulthood. In adolescents, only working memory could be separable from common EF, whereas in adults, the three EF subcomponents could be separable from common EF. However, findings of switching in adolescents should be treated with substantial caution and may be exploratory due to limited data available on switching tasks. For task materials, inhibition and working memory showed both domain generality and domain specificity, undergirded by the multiple-demand network, as well as different brain regions in response to verbal and nonverbal task materials, respectively. In contrast, switching showed only domain generality with no activation specialized for either verbal or nonverbal task materials. These findings, taken together, support and contribute to the unitary-diverse nature of EF such that EF should be interpreted in an integrative model that relies on the integration of the EF construct, development, and task materials.

Dynamics of language reorganization after left temporo-parietal and frontal stroke.

The loss and recovery of language functions are still incompletely understood. This longitudinal functional MRI study investigated the neural mechanisms underlying language recovery in patients with post-stroke aphasia putting particular emphasis on the impact of lesion site. To identify patterns of language-related activation, an auditory functional MRI sentence comprehension paradigm was administered to patients with circumscribed lesions of either left frontal (n = 17) or temporo-parietal (n = 17) cortex. Patients were examined repeatedly during the acute (≤1 week, t1), subacute (1-2 weeks, t2) and chronic phase (>6 months, t3) post-stroke; healthy age-matched control subjects (n = 17) were tested once. The separation into two patient groups with circumscribed lesions allowed for a direct comparison of the contributions of distinct lesion-dependent network components to language reorganization between both groups. We hypothesized that activation of left hemisphere spared and perilesional cortex as well as lesion-homologue cortex in the right hemisphere varies between patient groups and across time. In addition, we expected that domain-general networks serving cognitive control independently contribute to language recovery. First, we found a global network disturbance in the acute phase that is characterized by reduced functional MRI language activation including areas distant to the lesion (i.e. diaschisis) and subsequent subacute network reactivation (i.e. resolution of diaschisis). These phenomena were driven by temporo-parietal lesions. Second, we identified a lesion-independent sequential activation pattern with increased activity of perilesional cortex and bilateral domain-general networks in the subacute phase followed by reorganization of left temporal language areas in the chronic phase. Third, we observed involvement of lesion-homologue cortex only in patients with frontal but not temporo-parietal lesions. Fourth, irrespective of lesion location, language reorganization predominantly occurred in pre-existing networks showing comparable activation in healthy controls. Finally, we detected different relationships of performance and activation in language and domain-general networks demonstrating the functional relevance for language recovery. Our findings highlight that the dynamics of language reorganization clearly depend on lesion location and hence open new perspectives for neurobiologically motivated strategies of language rehabilitation, such as individually-tailored targeted application of neuro-stimulation.

A Domain-General Cognitive Core Defined in Multimodally Parcellated Human Cortex.

Numerous brain imaging studies identified a domain-general or "multiple-demand" (MD) activation pattern accompanying many tasks and may play a core role in cognitive control. Though this finding is well established, the limited spatial localization provided by traditional imaging methods precluded a consensus regarding the precise anatomy, functional differentiation, and connectivity of the MD system. To address these limitations, we used data from 449 subjects from the Human Connectome Project, with the cortex of each individual parcellated using neurobiologically grounded multimodal MRI features. The conjunction of three cognitive contrasts reveals a core of 10 widely distributed MD parcels per hemisphere that are most strongly activated and functionally interconnected, surrounded by a penumbra of 17 additional areas. Outside cerebral cortex, MD activation is most prominent in the caudate and cerebellum. Comparison with canonical resting-state networks shows MD regions concentrated in the fronto-parietal network but also engaging three other networks. MD activations show modest relative task preferences accompanying strong co-recruitment. With distributed anatomical organization, mosaic functional preferences, and strong interconnectivity, we suggest MD regions are well positioned to integrate and assemble the diverse components of cognitive operations. Our precise delineation of MD regions provides a basis for refined analyses of their functions.