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All of the cerebral cortex has some degree of laminar organization. These different layers are composed of neurons with distinct connectivity patterns, embryonic origins, and molecular profiles. There are little data on the laminar specificity of cognitive functions in the frontal cortex, however. We recorded neuronal spiking/local field potentials (LFPs) using laminar probes in the frontal cortex (PMd, 8A, 8B, SMA/ACC, DLPFC, and VLPFC) of monkeys performing working memory (WM) tasks. LFP power in the gamma band (50–250 Hz) was strongest in superficial layers, and LFP power in the alpha/beta band (4–22 Hz) was strongest in deep layers. Memory delay activity, including spiking and stimulus-specific gamma bursting, was predominately in superficial layers. LFPs from superficial and deep layers were synchronized in the alpha/beta bands. This was primarily unidirectional, with alpha/beta bands in deep layers driving superficial layer activity. The phase of deep layer alpha/beta modulated superficial gamma bursting associated with WM encoding. Thus, alpha/beta rhythms in deep layers may regulate the superficial layer gamma bands and hence maintenance of the contents of WM.

With the medial frontal cortex (MFC) centrally implicated in several major neuropsychiatric disorders, it is critical to understand the extent to which MFC organization is comparable between humans and animals commonly used in preclinical research (namely rodents and nonhuman primates). Although the cytoarchitectonic structure of the rodent MFC has mostly been conserved in humans, it is a long-standing question whether the structural analogies translate to functional analogies. Here, we probed this question using ultra high field fMRI data to compare rat, marmoset, and human MFC functional connectivity. First, we applied hierarchical clustering to intrinsically define the functional boundaries of the MFC in all three species, independent of cytoarchitectonic definitions. Then, we mapped the functional connectivity “fingerprints” of these regions with a number of different brain areas. Because rats do not share cytoarchitectonically defined regions of the lateral frontal cortex (LFC) with primates, the fingerprinting method also afforded the unique ability to compare the rat MFC and marmoset LFC, which have often been suggested to be functional analogs. The results demonstrated remarkably similar intrinsic functional organization of the MFC across the species, but clear differences between rodent and primate MFC whole-brain connectivity. Rat MFC patterns of connectivity showed greatest similarity with premotor regions in the marmoset, rather than dorsolateral prefrontal regions, which are often suggested to be functionally comparable. These results corroborate the viability of the marmoset as a preclinical model of human MFC dysfunction, and suggest divergence of functional connectivity between rats and primates in both the MFC and LFC.

In recent years, there has been substantial growth in neuroimaging studies investigating neural correlates of symbolic (e.g. Arabic numerals) and non-symbolic (e.g. dot arrays) number processing. At present it remains contested whether number is represented abstractly, or if number representations in the brain are format-dependent. In order to quantitatively evaluate the available neuroimaging evidence, we used activation likelihood estimation (ALE) to conduct quantitative meta-analyses of the results reported in 57 neuroimaging papers. Consistent with the existence of an abstract representation of number in the brain, conjunction analyses revealed overlapping activation for symbolic and nonsymbolic numbers in frontal and parietal lobes. Consistent with the notion of format-dependent activation, contrast analyses demonstrated anatomically distinct fronto-parietal activation for symbolic and non-symbolic processing. Therefore, symbolic and non-symbolic numbers are subserved by format-dependent and abstract neural systems. Moreover, the present results suggest that regions across the parietal cortex, not just the intraparietal sulcus, are engaged in both symbolic and non-symbolic number processing, challenging the notion that the intraparietal sulcus is the key region for number processing. Additionally, our analyses indicate that regions in the frontal cortex subserve magnitude representations rather than non-numerical cognitive processes associated with number tasks, thereby highlighting the importance of considering both frontal and parietal regions as important for number processing.

•Multimodal analysis of macaque motor and premotor cortex reveals novel parcellation•3D atlas with cyto- and multireceptor architectonic features of 16 (pre)motor areas•Primary motor area 4 is cyto- and receptor architectonically heterogeneous•(Pre)motor areas differ in their functional connectivity fingerprints In the present study we reevaluated the parcellation scheme of the macaque frontal agranular cortex by implementing quantitative cytoarchitectonic and multireceptor analyses, with the purpose to integrate and reconcile the discrepancies between previously published maps of this region. We applied an observer-independent and statistically testable approach to determine the position of cytoarchitectonic borders. Analysis of the regional and laminar distribution patterns of 13 different transmitter receptors confirmed the position of cytoarchitectonically identified borders. Receptor densities were extracted from each area and visualized as its “receptor fingerprint”. Hierarchical and principal components analyses were conducted to detect clusters of areas according to the degree of (dis)similarity of their fingerprints. Finally, functional connectivity pattern of each identified area was analyzed with areas of prefrontal, cingulate, somatosensory and lateral parietal cortex and the results were depicted as “connectivity fingerprints” and seed-to-vertex connectivity maps. We identified 16 cyto- and receptor architectonically distinct areas, including novel subdivisions of the primary motor area 4 (i.e. 4a, 4p, 4m) and of premotor areas F4 (i.e. F4s, F4d, F4v), F5 (i.e. F5s, F5d, F5v) and F7 (i.e. F7d, F7i, F7s). Multivariate analyses of receptor fingerprints revealed three clusters, which first segregated the subdivisions of area 4 with F4d and F4s from the remaining premotor areas, then separated ventrolateral from dorsolateral and medial premotor areas. The functional connectivity analysis revealed that medial and dorsolateral premotor and motor areas show stronger functional connectivity with areas involved in visual processing, whereas 4p and ventrolateral premotor areas presented a stronger functional connectivity with areas involved in somatomotor responses. For the first time, we provide a 3D atlas integrating cyto- and multi-receptor architectonic features of the macaque motor and premotor cortex. This atlas constitutes a valuable resource for the analysis of functional experiments carried out with non-human primates, for modeling approaches with realistic synaptic dynamics, as well as to provide insights into how brain functions have developed by changes in the underlying microstructure and encoding strategies during evolution. [Display omitted]

Covert endogenous (voluntary) attention improves visual performance. Human neuroimaging studies suggest that the putative human homolog of macaque frontal eye fields (FEF+) is critical for this improvement, whereas early visual areas are not. Yet, correlational MRI methods do not manipulate brain function. We investigated whether rFEF+ or V1/V2 plays a causal role in endogenous attention. We used transcranial magnetic stimulation (TMS) to alter activity in the visual cortex or rFEF+ when observers performed an orientation discrimination task while attention was manipulated. On every trial, they received double-pulse TMS at a predetermined site (stimulated region) around V1/V2 or rFEF+. Two cortically magnified gratings were presented, one in the stimulated region (contralateral to the stimulated area) and another in the symmetric (ipsilateral) nonstimulated region. Grating contrast was varied to measure contrast response functions (CRFs) for all attention and stimulation combinations. In experiment 1, the CRFs were similar at the stimulated and nonstimulated regions, indicating that early visual areas do not modulate endogenous attention during stimulus presentation. In contrast, occipital TMS eliminates exogenous (involuntary) attention effects on performance [A. Fernández, M. Carrasco,- Curr. Biol. 30, 4078–4084 (2020)]. In experiment 2, rFEF+ stimulation decreased the overall attentional effect; neither benefits at the attended location nor costs at the unattended location were significant. The frequency and directionality of microsaccades mimicked this pattern: Whereas occipital stimulation did not affect microsaccades, rFEF+ stimulation caused a higher microsaccade rate directed toward the stimulated hemifield. These results provide causal evidence of the role of this frontal region for endogenous attention.

Much of human learning emerges as a result of interaction with others. Yet, this interpersonal process has been poorly characterized from a neurophysiological perspective. This study investigated (i) whether Interpersonal Brain Synchronization (IBS) can reliably mark social interactive learning, and specifically (ii) during what kind of interactive behavior. We recorded brain activity from learner-instructor dyads using functional Near-Infrared Spectroscopy (fNIRS) during the acquisition of a music song. We made four fundamental observations. First, during the interactive learning task, brain activity recorded from the bilateral Inferior Frontal Cortex (IFC) synchronized across the learner and the instructor. Second, such IBS was observed in particular when the learner was observing the instructor's vocal behavior and when the learning experience entailed a turn-taking and more active mode of interaction. Third, this specific enhancement of IBS predicted learner's behavioral performance. Fourth, Granger causality analyses further disclosed that the signal recorded from the instructor's brain better predicted that recorded from the learner's brain than vice versa. Together, these results indicate that social interactive learning can be neurophysiologically characterized in terms of IBS. Furthermore, they suggest that the learner's involvement in the learning experience, alongside the instructor's modeling, are key factors driving the alignment of neural processes across learner and instructor. Such alignment impacts upon the real-time acquisition of new information and eventually upon the learning (behavioral) performance. Hence, besides providing a biological characterization of social interactive learning, our results hold relevance for clinical and pedagogical practices. •fNIRS hyperscanning is performed during social interactive learning.•Interpersonal brain synchronization (IBS) of IFC increases during learning.•IBS is affected by the learners' involvement.•IBS correlates with learners' performance.•IBS may function as a neural alignment across learner and instructor.

Attention Deficit Hyperactivity Disorder (ADHD) is associated with poor self-control, underpinned by inferior fronto-striatal deficits. We showed previously that 18 ADHD adolescents over 11 runs of 8.5 min of real-time functional magnetic resonance neurofeedback of the right inferior frontal cortex (rIFC) progressively increased activation in 2 regions of the rIFC which was associated with clinical symptom improvement. In this study, we used functional connectivity analyses to investigate whether fMRI-Neurofeedback of rIFC resulted in dynamic functional connectivity changes in underlying neural networks. Whole-brain seed-based functional connectivity analyses were conducted using the two clusters showing progressively increased activation in rIFC as seed regions to test for changes in functional connectivity before and after 11 fMRI-Neurofeedback runs. Furthermore, we tested whether the resulting functional connectivity changes were associated with clinical symptom improvements and whether they were specific to fMRI-Neurofeedback of rIFC when compared to a control group who had to self-regulate another region. rIFC showed increased positive functional connectivity after relative to before fMRI-Neurofeedback with dorsal caudate and anterior cingulate and increased negative functional connectivity with regions of the default mode network (DMN) such as posterior cingulate and precuneus. Furthermore, the functional connectivity changes were correlated with clinical improvements and the functional connectivity and correlation findings were specific to the rIFC-Neurofeedback group. The findings show for the first time that fMRI-Neurofeedback of a typically dysfunctional frontal region in ADHD adolescents leads to strengthening within fronto-cingulo-striatal networks and to weakening of functional connectivity with posterior DMN regions and that this may be underlying clinical improvement. [Display omitted]

We investigated afferent inputs from all areas in the frontal cortex (FC) to different subregions in the rostral anterior cingulate cortex (rACC). Using retrograde tracing in macaque monkeys, we quantified projection strength by counting retrogradely labeled cells in each FC area. The projection from different FC regions varied across injection sites in strength, following different spatial patterns. Importantly, a site at the rostral end of the cingulate sulcus stood out as having strong inputs from many areas in diverse FC regions. Moreover, it was at the integrative conjunction of three projection trends across sites. This site marks a connectional hub inside the rACC that integrates FC inputs across functional modalities. Tractography with monkey diffusion magnetic resonance imaging (dMRI) located a similar hub region comparable to the tracing result. Applying the same tractography method to human dMRI data, we demonstrated that a similar hub can be located in the human rACC.

Previous studies have shown that information held in visual working memory is represented in the occipital, parietal, and frontal cortices. However, less is known about whether the mnemonic information of multi-feature objects is modulated by task demand in the parietal and frontal regions. To address this question, we asked participants to remember either color or orientation of one of the two colored gratings for a delay. Using fMRI and an inverted encoding model, we reconstructed population-level, feature-selective responses in the occipital, parietal and frontal cortices during memory maintenance. We found that not only orientation but also color information can be maintained in higher-order parietal and frontal cortices as well as the early visual cortex when it was cued to be remembered. Conversely, neither the task-irrelevant feature of the cued object, nor any feature of the uncued object was maintained in the occipital, parietal, or frontal cortices. These results suggest a highly selective mechanism of visual working memory that maintains task-relevant features only. •Non-spatial features can be maintained in the parietal and frontal cortices.•Feature-specific representations in visual working memory are highly selective.•Only task-relevant features are maintained during visual working memory.

•We delineate fundamental neurophysiological mechanisms for how people can act intentionally.•We show that the ability to foresee the effects of action is a role of a theta activity.•There is bi-directional communication between temporo-frontal cortices is essential.•The results conceptually broaden the relevance of theta band activity. The ability to plan and carry out goal-directed behavior presupposes knowledge about the contingencies between movements and their effects. Ideomotor accounts of action control assume that agents integrate action-effect contingencies by creating action-effect bindings, which associate movement patterns with their sensory consequences. However, the neurophysiological underpinnings of action-effect binding are not yet well understood. Given that theta band activity has been linked to information integration, we thus studied action-effect integration in an electrophysiological study with N = 31 healthy individuals with a strong focus on theta band activity. We examined how information between functional neuroanatomical structures is exchanged to enable action planning. We show that theta band activity in a network encompassing the insular cortex (IC), the anterior temporal lobe (ATL), and the inferior frontal cortex (IFC) supports the establishment of action-effect bindings. All regions revealed bi-directional effective connectivities, indicating information transfer between these regions. The IC and ATL create a loop for information integration and the conceptual abstraction of it. The involvement of anterior regions of the IFC, particularly during the acquisition phase of the action-effect, likely reflects episodic control mechanisms in which a past event defines a “template” of what action-effect is to be expected. Taken together, the current findings connect well with major cognitive concepts. Our study suggests a functional relevance of theta band activity in an IC-ATL-IFC network, which in turn implies that basic ideomotor action-effect integration is implemented through theta band activity and effective connectivities between temporo-frontal structures.