Cerebral Cortex Communications

Abstract Background: Long COVID frequently includes post-exertional malaise, neuromuscular fatigue, internal vibrations, tremor-like symptoms, orthostatic intolerance, sleep disturbance, cognitive dysfunction, and autonomic instability. Neuroimaging studies in Long COVID and post-infectious ME/CFS have identified abnormalities across limbic, thalamic, mesiotemporal, brainstem, cerebellar, and white-matter systems, but no single established circuit accounts for this clinical phenotype. We examined whether abnormalities cluster at the mammillary body-fornix-superior tuberal hypothalamic interface and within connected brainstem-cerebellar pathways. Methods: Structural MRI and diffusion tensor imaging were analysed in 88 participants with clinician-diagnosed Long COVID and 34 healthy controls (total N = 122). Thirty-three patients were classified as bedridden. Volumetric and diffusion analyses focused on the mammillary bodies, superior tuberal hypothalamic region, fornix, dorsal and median raphe, midbrain reticular formation, and cerebellar peduncles. Fractional anisotropy was used as an index of white-matter microstructural organisation. Exploratory clinical associations included motor impairment, proprioceptive dysfunction, autonomic symptoms, fatigue, internal vibrations, and tremor-like symptoms. Results: Three mammillary-body volume phenotypes were identified: reduced, enlarged, and control-range. Left and right mammillary-body volumes differed across groups, with large effect estimates. Segmentation showed narrowing or loss of a visible internal passage at the superior tuberal-mammillary interface, while the fornix showed altered diffusion measures and reduced tract coherence in the hypothesised gate region. Additional findings included lower superior cerebellar peduncle volume, lower middle cerebellar peduncle fractional anisotropy, and lower dorsal raphe and midbrain reticular formation volumes in Long COVID. Conclusions: The findings support an exploratory mammillary body-fornix gate model in which a vulnerable periventricular hypothalamic-limbic interface may contribute to network dysfunction in a subgroup of patients with severe Long COVID. The data do not establish direct viral invasion, a coronavirus entry route, axonal destruction, or a single causal pathway. Prospective replication with standardised acquisition, preregistered regions of interest, correction for multiple comparisons, objective movement and autonomic measures, and longitudinal follow-up is required. Keywords: Long COVID; post-COVID condition; ME/CFS; mammillary body; fornix; hypothalamus; diffusion tensor imaging; internal vibrations; tremor-like symptoms; neuromuscular fatigue; dysautonomia; brainstem

Moderate-to-severe traumatic brain injury (msTBI) is recognized as a chronic and evolving neurological condition characterized by progressive structural brain changes and persistent cognitive impairment. While prior studies have demonstrated widespread atrophy following msTBI, less is known regarding the longitudinal trajectory of gray matter (GM) changes during recovery and post-rehabilitation. The current study used longitudinal voxel-based morphometry (VBM) to characterize GM volume changes over a period of 9 months, in individuals with msTBI relative to healthy controls (HC). Associations between regional GM volume and neuropsychological functioning were examined. Twenty-eight participants (14 msTBI, 14 HC) completed MRI and neuropsychological assessments across three timepoints spanning outpatient rehabilitation and follow-up. Longitudinal VBM analyses revealed significant group and time interactions within subcortical and limbic regions. Relative to HC, individuals with msTBI showed lower GM volume in these regions at baseline, with trajectories that converged toward HC values (right hippocampus) or increased relative to HC over the rehabilitation period (bilateral pulvinar), whereas the right amygdala and inferior cerebellar vermis remained persistently reduced. Significant longitudinal improvements in memory and psychomotor speed during the rehabilitation period were demonstrated in msTBI. Greater (preserved) GM volume within the right hippocampus, thalamus, and bilateral pulvinar was associated with better performance across measures of verbal memory, processing speed, executive functioning, and cognitive flexibility. These findings suggest that msTBI is associated with dynamic structural brain changes involving subcortical, limbic, and cerebellar networks, and that the rehabilitation period was accompanied by relative volumetric stabilization in these regions and by meaningful cognitive improvement.

Voluntary movement requires the central nervous system to transform and integrate visual and somatosensory information into coordinated motor outputs. Although mirrorlike neuronal activity during both action execution and observation has been extensively described in premotor, motor, and parietal cortices, it remains unknown whether the primary somatosensory cortex (S1) also participates in the action observation network. Here, we recorded single-unit activity from cytoarchitectonically defined areas 3a, 3b, 1, and 2 in macaque S1 while monkeys either executed or observed grasping movements. Approximately one-third of neurons across S1 modulated their firing during action observation, with the proportion of responsive neurons increasing from area 3 to areas 1 and 2, consistent with the hierarchical organization of somatosensory processing. Most action observation neurons showed congruent activity during action execution and observation, suggesting that these responses may reflect top-down motor-related or integrated visuomotor signals and are unlikely to be explained by visual input alone. The higher prevalence of action observation neurons in areas 1 and 2 suggests that action observation-related signals preferentially influence later stages of somatosensory processing, potentially via cortico-cortical interactions with motor and parietal regions.

Optogenetic stimulation of the rostral pedunculotegmental nucleus (PTg) induces global motor arrest, but it remains unclear whether this is merely a suppression of motor activity or a broader disruption of brain processes required to guide action. We developed a visuospatial cue task for rats, to test if sensory information presented during PTg-induced arrest can guide later responses. Here, we show that optogenetic stimulation during cue presentation reduces accuracy to chance level. By moving stimulation to only before or only after the cue, we found that performance was only affected when stimulation and cue presentation overlapped, that rats recover cue-guided behavior almost immediately at the end of stimulation, and that stimulation does not appear to abolish responses based on cue information acquired before arrest. These findings indicate that stimulation of the rostral PTg does not only pause motor output but transiently disrupts the ability to process and use cue information effectively.

The ventral premotor cortex (PMv) has been implicated in both action selection and the perception of observed actions, but it remains unclear whether PMv activity during action observation reflects the observed action itself, or variables related to the observers own action when the observed action becomes behaviourally relevant. Here, we recorded neural activity in macaque PMv during a task that dissociated observed action, rule context, and the subsequently selected self-action. Population activity during observation was already biased toward the upcoming self-action and became increasingly aligned with it. At the level of single neurons, subsets showed modulation by rule and required self-action beyond the observed action. Task-related variables coexisted during observation, rather than being organized into distinct sequential stages. These findings indicate that PMv activity during action observation is not solely determined by the observed action, but instead reflects variables related to the selection of the agents own action, consistent with a transformation from observed action to self-action.

Cognitive functions encompass a large set of abstract constructs used for adapting our behavior to environmental constraints, each of them acting at specific timescales. For example, while working memory operates over several seconds or minutes, decision making occurs over much shorter periods. Solving a behavioral task thus relies on specific cognitive strategies that use several functions over time. Here, we investigated how two macaque monkeys coordinate working memory, selective attention, decision-making and executive control of eye movements during performance of a modified delay match-to-sample task. The economy of this task (including reward expected value and cost of errors) evolves on short timescales within trials. In addition, this task allowed us to manipulate engagement of cognitive resources at longer timescales. Using tools from signal detection theory, we closely analyze how monkeys performances evolve over time and infer their specific strategies in terms of control of cognitive functions. In addition, during covert attention, fixational eye movements and pupil size, provided reliable markers of slow variations in cognitive state across trials, although not of rapid within-trial changes in cognitive control. Together, these results show that each monkey adapted to the same task by implementing individual, dynamically evolving cognitive strategies across multiple timescales.

Object representations in the human ventral occipitotemporal cortex (VOTC) are organized along multiple dimensions, including shape (rectilinear vs. curvilinear), real-world size (large vs. small), and mobility (stationary vs. mobile). However, these dimensions are strongly correlated in naturalistic vision, making their separate contributions to VOTC organization unclear. For example, large objects (e.g., a wardrobe, a house) are typically rectilinear and stationary, while small objects (e.g., a ball, a cup) are more curvilinear and mobile. Here, we used fMRI, together with a new stimulus set that orthogonally manipulates shape, size, and mobility, to investigate the separate influences of these dimensions on VOTC organization. Example stimuli include air balloon (large, curvilinear, mobile), radar dish (large, curvilinear, stationary), and mailbox (small, rectilinear, stationary). Contrasts revealed that large (vs. small), rectilinear (vs. curvilinear), and stationary (vs. mobile) dimensions all independently evoked strong and overlapping activity in medio-anterior VOTC. This overlapping activity was at the intersection of the parahippocampal place area (PPA) and the ventral place-memory area (VPMA). Similar results were found at the intersection of the scene-selective occipital place area and the lateral place-memory area (LPMA). Finally, large (vs. small), but not rectilinear (vs. curvilinear) or stationary (vs. mobile) activity, was found in additional posterior ventral scene-selective regions, as well as in early visual cortex. Overall, these results indicate that object shape, real-world size, and mobility dimensions all independently activate scene-selective PPA and OPA, showing joint selectivity for distinct low- and high-level object properties that are highly correlated in naturalistic vision.

Cognitive flexibility is the ability to change the way of responding when the demands of the environment change. This study tested how cognitive flexibility develops across the lifespan. We used a new computerized task that gives a continuous score instead of just right or wrong answers. 221 healthy adults aged 18 to 71 completed the Continuous-score Probabilistic Reversal Learning Test (CPRLT). We calculated mean absolute error and adjusted error for rule-based learning, and fitted a Rescorla-Wagner model to estimate each persons learning rate (alpha) for reward-based learning. All three scores have one breakpoint, performance improved rapidly from childhood to young adulthood, then declined slowly. Rule-based learning peaked around age 20. Reward-based learning peaked earlier, around age 18. This suggests that reward-based learning matures before rule-based learning. The pattern fits with brain development: reward circuits mature earlier, while prefrontal regions for rule-based learning develop later. Our continuous measure captured this difference, which binary tasks would miss.

Memorizing a new phone number or address is hard at first, but becomes easier with repetition, as information shifts from working memory to long-term memory. Here we investigated how repetition affects the storage and transition of different aspects of mnemonic information by comparing univariate neural markers of active object storage with multivariate decoding of memory content. Thirty participants encoded lateralized stimuli from a continuous shape space into memory. Memory items were repeated six times in a row to induce learning. In line with earlier work, EEG recordings revealed that repetition led to a reduction in contralateral delay activity (CDA), a measure of active storage that has been taken to reflect a pointer-like representation of the individual object or its original source. In contrast, shape decoding during the retention and also after an impulse perturbation remained constant across repetitions. These results suggest that learning over repetitions reflects the abolishment of active and individuated object memory representations while passive, source-independent memory representations are retained.

Audiovisual semantic matching can be achieved using either written words or pictures, yet whether these formats engage shared semantic matching representations with similar temporal dynamics remains unclear. We recorded electroencephalography from 27 participants while they performed audiovisual semantic matching tasks in which spoken words were paired with either written words or pictures. Stimuli included both natural and man-made objects. Time-resolved multivariate pattern analyses (MVPA or decoding), cross-decoding, and temporal generalization analyses were used to characterize the temporal dynamics of semantic processing. Reliable decoding of matching versus mismatching judgments emerged in both word and picture conditions. Decoding onset that significant above chance level occurred earlier for written words than for pictures and cross-decoding analyses revealed successful generalization between word and picture formats. Temporal generalization analyses further demonstrated distinct representational dynamics across formats, with word processing characterized by predominantly time-specific neural representations and picture processing showing more sustained and temporally stable representations. In addition, matching-related discrimination emerged earlier for natural objects than for man-made objects across both formats. The results suggest that speech-word matching shows earlier neural evidence of audiovisual alignment than speech-picture matching, potentially reflecting differences in how auditory linguistic input is integrated with visual information across representational formats.

Audiovisual cues ("bells and whistles") are ubiquitous in commercial gambling products. Pairing wins with sound and light cues in laboratory-based gambling paradigms increases risky choice, but the neurocognitive basis of this effect is unclear. Here we compared patterns of neural activation using functional MRI in healthy volunteers (n = 31) while they performed a two-choice lottery task. Reward-paired cues were either present or absent in a mixed-block, event-related design. As predicted, participants made riskier choices on cued trials. Choice latencies were also longer when cues were present, particularly on trials following a win. Activity within the nucleus accumbens and orbitofrontal cortex was greater during the decision phase when participants made risky choices. Nucleus accumbens signal was also greater when participants were anticipating risky outcomes, and in response to risky wins. Contrary to our pre-registered hypotheses, cue condition did not alter patterns of activity across any task phase, in either of these a priori regions of interest. As such, cue-induced risky choice does not appear to be driven by altered representation of risk or value within this canonically reward-sensitive circuitry. Instead, exploratory analyses revealed that the anterior insula was selectively activated by cued, risky wins. Such activation may signal the saliency of these events, or their emotional impact, and may reflect one mechanism through which cue-induced craving develops in vulnerable individuals.

Our visual environment can be parsed into objects and scenes, a distinction that is reflected in the organization of the human visual cortex. Previous research has shown that object and scene perception nevertheless closely interact, such that scenes influence object perception and objects influence scene perception. It remains unclear, however, whether and how objects that are not inherently diagnostic of their surroundings aid the recognition of poorly visible scenes (e.g., a person standing in a dark living room). Here, in three behavioral experiments, we show that participants made more accurate indoor/outdoor judgments when degraded scene photographs were presented together with an object than when the scene or the object was shown alone, even though the same object categories appeared in indoor and outdoor scenes. This object-driven benefit vanished once scene structure was removed through phase scrambling and was reduced when objects appeared in physically inconsistent locations within the scenes. These results suggest that objects in consistent locations (e.g., a person standing on a floor) disambiguate scene layout. Finally, in a pre-registered transcranial magnetic stimulation (TMS) study (N = 48), we provide causal evidence that the object-selective lateral occipital cortex (LOC) supports scene categorization when scene layout is disambiguated by within-scene objects. Stimulation of the LOC, particularly at 260-300 ms after stimulus onset, selectively disrupted object-based scene recognition. Together, these findings demonstrate that objects facilitate the read-out of the surrounding space in service of efficient scene recognition. Significance StatementUnderstanding how scene and object processing interact for efficient recognition is a key question in natural vision. Research has long emphasized how surrounding scenes help us identify objects, yet the reverse - how objects shape the perception of scenes - has received little attention. In the dark, does a glimpse of a floating boat tell us we are looking at a lake? In this study we demonstrate that a single object helps people recognize hardly visible scenes. This benefit required intact scene structure and depended on where the object appeared in the scene. In addition, object selective visual cortex was causally related to this benefit. Together, these findings show that objects visual appearance can be used to better understand our surroundings.

Background Chemotherapy-related cognitive impairment is a well-documented concern among cancer survivors, yet the neural mechanisms underlying deficits in cognitive control remain poorly understood. This study examined group differences in brain activation during a flanker task using functional MRI (fMRI) between chemotherapy-exposed participants and healthy controls. Methods Participants (21 survivors (24.9 years old; 71.4 % female; 15 years from diagnosis) and 21 healthy controls (26.7 years old; 61.9 % female) completed a flanker task during fMRI, with congruent and incongruent conditions. Reaction time, accuracy, and Flanker scores were collected. Whole-brain group comparisons were performed for congruent, incongruent, and incongruent > congruent contrasts. Associations between the incongruent > congruent contrast and cognitive performance were examined. Results Compared to controls, the Chemo group had longer reaction times in both congruent and incongruent conditions (p < .001) and lower NIH Flanker scores (p = .01), with no differences in accuracy. They showed reduced activation in the bilateral inferior frontal gyri, supplementary motor area, and bilateral caudate, but greater activation in the right inferior temporal and cerebellar regions. The incongruent > congruent contrast correlated with increased activation in the orbitofrontal cortex, inferior temporal gyri, and fusiform gyrus with cognitive performance. Conclusions Chemotherapy-exposed participants showed cognitive control deficits and altered neural activation during a flanker task, indicating disrupted recruitment of frontoparietal and subcortical regions key for conflict processing. These findings improve understanding of neural causes of chemotherapy-related cognitive impairment and may help identify at-risk survivors and guide personalized rehabilitation.

Motor outputs vary even in response to identical sensory inputs, yet the origin of this variability within the sensorimotor pathway remains unresolved. Here, we evaluate how variability in sensory representations can explain behavioural fluctuations in a reflexive oculomotor task. We used Neuropixels probes to record neuronal activity in area MT of marmosets during ocular following responses and applied partial least squares regression to extract the shared variance between population activity and eye movements. Stimulus-evoked activity reliably predicted trial-by-trial variability in open-loop eye velocity. Additionally, closed-loop analyses revealed trial-by-trial correspondence between eye movements and subsequent neural responses. These results demonstrate that variability in sensory neural populations contributes to motor variability, supporting the claim that sensory noise is propagated through the sensorimotor pathway. Surprisingly, however, traditional models trained to only capture neural variability across trials poorly predicted behavioural variations, suggesting that only a subset of sensory representations is accessible to the motor system.

A long-standing debate surrounding the neural bases of social concepts concerns the role of anterior temporal lobe (ATL). One perspective suggests ATL subregions are dedicated specifically to social knowledge; another suggests the ATLs constitute a domain-general hub for conceptual knowledge, but with graded functional specialisation depending on connectivity to modality specific spokes. The positions have been difficult to adjudicate due to many confounding factors in tests of social and non-social knowledge. We address these challenges via three innovations in assessment of knowledge in frontotemporal dementia (FTD). First, we introduce a new task that controls for several potential confounds. Second, we apply mixed linear models to behavioural data analysis, allowing further control over confounding factors. Third, we extend the mixed-model approach to lesion-symptom mapping, identifying cortical regions where structural pathology yields a disproportionate impairment on social versus non-social knowledge when other factors are controlled. We used these techniques to probe social and non-social knowledge in FTD subtypes: semantic dementia (SD), associated with asymmetric-bilateral ATL atrophy (n=21), and behavioural-variant (bvFTD), characterised by frontoinsular atrophy (n=24). When confounding factors were controlled, people with SD showed an equal impairment for social and non-social concepts, whereas those with bvFTD were disproportionately impaired on social concepts. The differential impairment of social concepts was associated with atrophy in the insula, orbitofrontal and ventromedial prefrontal cortex and other regions implicated in social knowledge generally. The results suggest that the bilateral ATLs constitute a domain-general semantic hub, whereas ventral prefrontal and insula cortex contribute preferentially to knowledge about people.

In human brains, portions of the frontoparietal cortex respond when people reason about the physical world (e.g. track objects, make predictions), and when people engage in a variety of demanding tasks (e.g. working memory, motor inhibition). Here, we use functional neuroimaging to address an open question about whether the same neural resources support both mental functions. Twenty-eight human adults (Mage = 26.5y; 17 female; 26 right handed) were scanned using functional magnetic resonance imaging (fMRI) while they (i) tracked the dynamics of physical objects (vs social agents), and (ii) performed a hard (vs easy) spatial working memory task. For each participant, we identified functional regions of interest (fROIs) that were maximally engaged by each task (physical > social; hard > easy), and studied their engagement in the held out task. We report three pieces of evidence that physical > social fROIs are recruited during the spatial working memory task. First, fROIs maximally engaged during physical (vs social) processing responded just as strongly during hard (vs easy) spatial working memory. Second, people with strong hard > easy responses in one set of regions also tended to have strong physical > social responses in the other set. Third, people with strong hard > easy responses in physical > social fROIs, but not in hard > easy fROIs, performed more strongly at spatial working memory in the scanner. These findings suggest that portions of the frontoparietal cortex that preferentially respond to physical (vs social) stimuli are involved in functions beyond physical reasoning: either spatial processing specifically, or attentionally demanding tasks in general.

The consolidation of novel motor skills has traditionally been investigated over the timescale of hours-days following practice. However, a growing body of evidence has demonstrated that consolidation of skills occurs more rapidly across a scale of seconds - a process termed micro-consolidation. Micro-consolidation of explicitly cued motor sequences is supported by frontoparietal beta oscillations. Whether beta power modulation is a common mechanism supporting rapid consolidation of other forms of skill learning, such as implicit sequence learning, is yet to be elucidated. 72 healthy adults aged 18-35 (55% female) completed a serial reaction time task with concurrent electroencephalography recording. In line with our previous work, we show that the early fast learning on an implicit sequence task is primarily expressed as micro-offline gains during brief rest periods between periods of practice. Beta power was modulated as a function of active practice vs rest epochs (i.e., event-related synchronisation/desynchronisation), and here we demonstrate that micro-offline gains are associated with this modulation of beta power during the rest epochs. This relationship was specific to the beta frequency, and was not observed across either mu or gamma bands. Overall, in line with seminal work implicating beta in early explicit motor learning, our results indicate that beta is a shared neurophysiological signature of micro-consolidation of implicit sequences. Key points summary- Recent evidence demonstrates rapid consolidation of motor skills over seconds, termed micro-consolidation. - Seminal work has implicated beta oscillations in early explicit motor learning, though whether beta supports rapid consolidation of other forms of skill learning is unclear. - Using electroencephalography, we demonstrate that beta modulation is a neurophysiological signature of micro-consolidation of implicitly learnt motor sequences. - Lower beta at rest is associated with a higher degree of micro-consolidation. - Our findings shed critical new insight into the neurophysiological mechanisms mediating rapid consolidation of implicit motor skills during early fast learning.

The locus coeruleus-norepinephrine (LC-NE) system plays multiple roles in higher brain function that are thought to depend on its mode of activation, which reflects relationships between baseline and evoked activation levels. These relationships are evident in both single-unit LC activity and proposed physiological proxies of LC-NE activity, such as pupil size. Here we used measurements in awake monkeys to show that the baseline-evoked relationships evident within these two different measures are unreliably coupled between them: baseline-evoked relationships of the pupil are not predictive of those in the LC, and vice versa. These results imply that pupil modulations, which can reflect LC-NE activity, should be used with caution to make inferences about "phasic" (moderate baseline, high evoked) and "tonic" (high baseline, low evoked) LC-NE activity modes that are thought to support different forms of information processing in the brain.

The noradrenergic nucleus locus coeruleus (LC) is involved in numerous cognitive functions. Its activation often enhances sensory and motor performance and its activity correlates with arousal, which altogether suggest a general role in the mobilization of resources for cognition and action. We recently showed its strong and specific implication in physical effort and indirect evidence suggest that it might also be involved in cognitive effort. To address this question directly, we used a pharmacogenetic approach in rhesus macaques to selectively and reversibly inhibit LC neurons in a cognitively challenging task. Two monkeys were injected with viral vectors expressing inhibitory DREADDs (hM4Di) specifically in noradrenergic LC neurons, allowing reversible suppression of LC activity via systemic administration of deschloroclozapine (DCZ, 0.1 mg/kg). A third monkey served as a control and only received DCZ injections. Monkeys performed a simple hole-board task in which they searched for food rewards (raisins) hidden in a 5x5 grid of wells. In the transparent condition, rewards were visible, requiring minimal cognitive effort. In the opaque condition, rewards were hidden, such that monkey had to rely upon working memory to avoid revisiting empty wells. Thus, performance in opaque condition required more cognitive effort. Behavioral analysis showed that LC inhibition had no effect on performance in the transparent condition. However, in the opaque condition, it significantly impaired performance by increasing errors (revisits), without affecting the total number of rewards obtained, response times, or overall motivation. Monkeys compensated the decrease in success rate by performing more trials, indicating reduced efficiency rather than disengagement. These findings demonstrate that the LC plays a critical causal role in mobilizing cognitive resources for a demanding task. This is in line with the idea that the noradrenergic system contributes broadly to cognitive control and effort, extending previous findings on its involvement in physical effort. Overall, the study provides strong evidence linking LC activity to cognitive effort regulation, thereby complementing non-invasive studies in humans.

Human social perception relies on multiple visual signals, including object-directed actions and emotional facial expressions. It remains unclear whether responses to these signals converge within action-observation regions and whether apparent convergence depends on the spatial scale of analysis. We scanned 39 healthy human adults during two block-design fMRI tasks, hand action observations contrasted with paper-tube controls, and emotional expression observations contrasted with neutral faces. Using independently defined meta-analytic action observation network (AON) ROIs in the inferior frontal gyrus (IFG), inferior parietal lobule (IPL), and posterior superior temporal sulcus (pSTS), we analysed responses at whole-brain, ROI-average, and AON-constrained voxelwise scales. Whole-brain overlap between the two contrasts was localised primarily to the left lateral occipitotemporal extrastriate cortex, outside the AON. At the regional ROI-average scale, the IFG showed positive responses to both contrasts, whereas the IPL and pSTS showed opposing task preferences. Crucially, regional IFG co-engagement did not imply that the same voxels responded to both contrasts. Suprathreshold voxelwise overlap within the AON was virtually absent, and individual-level analysis revealed a small but reliable posterior displacement of hand action relative to emotional face within IFG. These findings show that spatial convergence between hand and emotion observation responses depends on the scale of analysis. The same IFG region can be engaged by both stimulus domains while retaining distinct local response topographies.