Ultralow-frequency neural entrainment to pain.

Nervous systems exploit regularities in the sensory environment to predict sensory input, adjust behavior, and thereby maximize fitness. Entrainment of neural oscillations allows retaining temporal regularities of sensory information, a prerequisite for prediction. Entrainment has been extensively described at the frequencies of periodic inputs most commonly present in visual and auditory landscapes (e.g., >0.5 Hz). An open question is whether neural entrainment also occurs for regularities at much longer timescales. Here, we exploited the fact that the temporal dynamics of thermal stimuli in natural environment can unfold very slowly. We show that ultralow-frequency neural oscillations preserved a long-lasting trace of sensory information through neural entrainment to periodic thermo-nociceptive input as low as 0.1 Hz. Importantly, revealing the functional significance of this phenomenon, both power and phase of the entrainment predicted individual pain sensitivity. In contrast, periodic auditory input at the same ultralow frequency did not entrain ultralow-frequency oscillations. These results demonstrate that a functionally significant neural entrainment can occur at temporal scales far longer than those commonly explored. The non-supramodal nature of our results suggests that ultralow-frequency entrainment might be tuned to the temporal scale of the statistical regularities characteristic of different sensory modalities.

The effect of salient stimuli on neural oscillations, isometric force, and their coupling.

Survival in a suddenly-changing environment requires animals not only to detect salient stimuli, but also to promptly respond to them by initiating or revising ongoing motor processes. We recently discovered that the large vertex brain potentials elicited by sudden supramodal stimuli are strongly coupled with a multiphasic modulation of isometric force, a phenomenon that we named cortico-muscular resonance (CMR). Here, we extend our investigation of the CMR to the time-frequency domain. We show that (i) both somatosensory and auditory stimuli evoke a number of phase-locked and non-phase-locked modulations of EEG spectral power. Remarkably, (ii) some of these phase-locked and non-phase-locked modulations are also present in the Force spectral power. Finally, (iii) EEG and Force time-frequency responses are correlated in two distinct regions of the power spectrum. An early, low-frequency region (∼4 Hz) reflects the previously-described coupling between the phase-locked EEG vertex potential and force modulations. A late, higher-frequency region (beta-band, ∼20 Hz) reflects a second coupling between the non-phase-locked increase of power observed in both EEG and Force. In both time-frequency regions, coupling was maximal over the sensorimotor cortex contralateral to the hand exerting the force, suggesting an effect of the stimuli on the tonic corticospinal drive. Thus, stimulus-induced CMR occurs across at least two different types of cortical activities, whose functional significance in relation to the motor system should be investigated further. We propose that these different types of corticomuscular coupling are important to alter motor behaviour in response to salient environmental events.

Saliency Detection as a Reactive Process: Unexpected Sensory Events Evoke Corticomuscular Coupling.

Survival in a fast-changing environment requires animals not only to detect unexpected sensory events, but also to react. In humans, these salient sensory events generate large electrocortical responses, which have been traditionally interpreted within the sensory domain. Here we describe a basic physiological mechanism coupling saliency-related cortical responses with motor output. In four experiments conducted on 70 healthy participants, we show that salient substartle sensory stimuli modulate isometric force exertion by human participants, and that this modulation is tightly coupled with electrocortical activity elicited by the same stimuli. We obtained four main results. First, the force modulation follows a complex triphasic pattern consisting of alternating decreases and increases of force, time-locked to stimulus onset. Second, this modulation occurs regardless of the sensory modality of the eliciting stimulus. Third, the magnitude of the force modulation is predicted by the amplitude of the electrocortical activity elicited by the same stimuli. Fourth, both neural and motor effects are not reflexive but depend on contextual factors. Together, these results indicate that sudden environmental stimuli have an immediate effect on motor processing, through a tight corticomuscular coupling. These observations suggest that saliency detection is not merely perceptive but reactive, preparing the animal for subsequent appropriate actions.SIGNIFICANCE STATEMENT Salient events occurring in the environment, regardless of their modalities, elicit large electrical brain responses, dominated by a widespread "vertex" negative-positive potential. This response is the largest synchronization of neural activity that can be recorded from a healthy human being. Current interpretations assume that this vertex potential reflects sensory processes. Contrary to this general assumption, we show that the vertex potential is strongly coupled with a modulation of muscular activity that follows the same pattern. Both the vertex potential and its motor effects are not reflexive but strongly depend on contextual factors. These results reconceptualize the significance of these evoked electrocortical responses, suggesting that saliency detection is not merely perceptive but reactive, preparing the animal for subsequent appropriate actions.

The Role of the Primary Sensory Cortices in Early Language Processing.

The results of this magnetoencephalography study challenge two long-standing assumptions regarding the brain mechanisms of language processing: First, that linguistic processing proper follows sensory feature processing effected by bilateral activation of the primary sensory cortices that lasts about 100 msec from stimulus onset. Second, that subsequent linguistic processing is effected by left hemisphere networks outside the primary sensory areas, including Broca's and Wernicke's association cortices. Here we present evidence that linguistic analysis begins almost synchronously with sensory, prelinguistic verbal input analysis and that the primary cortices are also engaged in these linguistic analyses and become, consequently, part of the left hemisphere language network during language tasks. These findings call for extensive revision of our conception of linguistic processing in the brain.

Brain activation profiles during kinesthetic and visual imagery: An fMRI study.

The aim of this study was to identify brain regions involved in motor imagery and differentiate two alternative strategies in its implementation: imagining a motor act using kinesthetic or visual imagery. Fourteen adults were precisely instructed and trained on how to imagine themselves or others perform a movement sequence, with the aim of promoting kinesthetic and visual imagery, respectively, in the context of an fMRI experiment using block design. We found that neither modality of motor imagery elicits activation of the primary motor cortex and that each of the two modalities involves activation of the premotor area which is also activated during action execution and action observation conditions, as well as of the supplementary motor area. Interestingly, the visual and the posterior cingulate cortices show reduced BOLD signal during both imagery conditions. Our results indicate that the networks of regions activated in kinesthetic and visual imagery of motor sequences show a substantial, while not complete overlap, and that the two forms of motor imagery lead to a differential suppression of visual areas.

Involvement of the superior temporal cortex in action execution and action observation.

The role of the superior temporal sulcus (STs) in action execution and action observation remains unsettled. In an attempt to shed more light on the matter, we used the quantitative method of (14)C-deoxyglucose to reveal changes in activity, in the cortex of STs and adjacent inferior and superior temporal convexities of monkeys, elicited by reaching-to-grasp in the light or in the dark and by observation of the same action executed by an external agent. We found that observation of reaching-to-grasp activated the components of the superior temporal polysensory area [STP; including temporo-parieto-occipital association area (TPO), PGa, and IPa], the motion complex [including medial superior temporal area (MST), fundus of superior temporal area (FST), and dorsal and ventral parts of the middle temporal area (MTd and MTv, respectively)], and area TS2. A significant part of most of these activations was associated with observation of the goal-directed action, and a smaller part with the perception of arm-motion. Execution of reaching-to-grasp in the light-activated areas TS2, STP partially and marginally, and MT compared with the fixation but not to the arm-motion control. Consequently, MT-activation is associated with the arm-motion and not with the purposeful action. Finally, reaching-to-grasp in complete darkness activated all components of the motion complex. Conclusively, lack of visibility of our own actions involves the motion complex, whereas observation of others' actions engages area STP and the motion complex. Our previous and present findings together suggest that sensory effects are interweaved with motor commands in integrated action codes, and observation of an action or its execution in complete darkness triggers the retrieval of the visual representation of the action.

Viewing a forelimb induces widespread cortical activations.

Given that prerequisite of activating the mirror neuron system is the preshaping of the hand and its interaction with the object during observation of a reaching-to-grasp-an-object action, the effects of viewing the object, the reaching forelimb and the static hand may obscure the effects of observing the grasping action per se. To disentangle these effects, we employed the (14)C-deoxyglucose quantitative autoradiographic method to map the functional activity in the entire cortex of monkeys (Macaca mulatta) which observed the experimenter performing non-goal-directed (purposeless) forelimb movements towards an object that was previously presented but no longer visible. Thus, our monkeys were exposed to the view of an object, a moving arm and a static hand with extended wrist and fingers. The distribution of metabolic activity was analyzed in 20µm thick brain sections, and two dimensional maps were reconstructed in the occipital operculum, the temporal, the lateral and medial parietal, the lateral and medial frontal, the lateral prefrontal and orbitofrontal cortices, including the cortex within the lunate, superior temporal, lateral, parietoccipital, intraparietal, central, arcuate and principal sulci. Increased metabolic activity, as compared to fixation-control monkeys, was measured in the forelimb representation of the primary motor and somatosensory cortices, the premotor cortices F2 and F5, cingulate motor areas, the secondary somatosensory cortex SII, the posterior intraparietal area 5 and areas TPOc and FST, in the hemisphere contralateral to the moving arm. Moreover, bilateral activations were elicited in areas pre-SMA, 8m, SSA and the somatorecipient area VS, the retroinsula, the auditory belt area CM, motion areas MT, MST, LOP/CIP, area 31, visual areas TEO, V6, V6Av and the parafoveal and peripheral visual representations of areas V1 and V2, respectively. Few parietal, auditory and visual areas were bilaterally depressed. In brief, a surprisingly wide cortical network is recruited even by mere observation of an arm executing goalless movements, which partially overlaps with the cortical network supporting the execution and observation of goal-directed forelimb actions. Interestingly, this overlap concerns mainly lower order sensory-motor rather than higher order association prefrontal and parietal cortices. Our results demonstrate that in order to reveal the net effects specifically induced by observation of a purposeful reaching-to-grasp action, the use of an appropriate control taking into account the effects of viewing the object to be grasped, the reaching arm and the static hand is crucial.

Grasping in the dark activates early visual cortices.

We have previously demonstrated that the primary motor and somatosensory cortices of monkeys are somatotopically activated for action-observation as are for action-generation, indicating that the recruitment of learned somatosensory-motor representations underlies the perception of others' actions. Here we examined the effects of seen and unseen actions on the early visual cortices, to determine whether stored visual representations are employed in addition to the somatosensory-motor ones. We used the quantitative (14)C-deoxyglucose method to map the activity throughout the cortex of the occipital operculum, lunate, and inferior occipital sulci of "rhesus monkeys" who reached to grasp a 3D object either in the light or in the dark or who observed the same action executed by another subject. In all cases, the extrastriate areas V3d and V3A displayed marked activation. We suggest that these activations reflect processing of visuospatial information useful for the reaching component of action, and 3D object-related information useful for the grasping part. We suggest that a memorized visual representation of the action supports action-recognition, as well as action-execution in complete darkness when the object and its environment are invisible. Accordingly, the internal representation that serves action-cognition is not purely somatosensory-motor but also includes a visual component.

Evaluation of image quality metrics for the prediction of subjective best focus.

PURPOSE: Seven existing and three new image quality metrics were evaluated in terms of their effectiveness in predicting subjective cycloplegic refraction. METHODS: Monochromatic wavefront aberrations (WA) were measured in 70 eyes using a Shack-Hartmann based device (Complete Ophthalmic Analysis System; Wavefront Sciences). Subjective cycloplegic spherocylindrical correction was obtained using a standard manifest refraction procedure. The dioptric amount required to optimize each metric was calculated and compared with the subjective refraction result. Metrics included monochromatic and polychromatic variants, as well as variants taking into consideration the Stiles and Crawford effect (SCE). WA measurements were performed using infrared light and converted to visible before all calculations. RESULTS: The mean difference between subjective cycloplegic and WA-derived spherical refraction ranged from 0.17 to 0.36 diopters (D), while paraxial curvature resulted in a difference of 0.68 D. Monochromatic metrics exhibited smaller mean differences between subjective cycloplegic and objective refraction. Consideration of the SCE reduced the standard deviation (SD) of the difference between subjective and objective refraction. CONCLUSIONS: All metrics exhibited similar performance in terms of accuracy and precision. We hypothesize that errors pertaining to the conversion between infrared and visible wavelengths rather than calculation method may be the limiting factor in determining objective best focus from near infrared WA measurements.