Ada Le

Spatial frequency-specific effects on the attentional bias: Evidence for two attentional systems

Using a gratingscales task as a sensitive measure of the attentional bias, we have recently observed a new form of frequency-specific cross-over; people showed left-biased preferences when comparing the high spatial frequency (HiSF) components of the task and rightward biases when comparing low spatial frequencies (LoSFs). Here we investigated which mechanisms underlie the cross-over. (1) We found that leftward and rightward biases were positively correlated, suggesting that the same set of mechanisms are involved in both versions of the task. (2) When we cued attention to the left or right side we found transient effects on gratingscales biases that were symmetrical for the LoSF condition but asymmetrical for the HiSF condition. This indicates that the HiSF condition itself biased stimulus-driven attention more to the left side than the LoSF condition. (3) When we lowered the contrast of the HiSF or the LoSF stimulus components, specifically the latter case made HiSF and LoSF conditions more different. This suggests that HiSF and LoSF conditions differ because HiSF components are more salient and more likely stir stimulus-driven attention. Our data are consistent with the idea that the attentional bias results from right-dominant control mechanisms of stimulus-driven attention potentially interacting with voluntary control mechanisms.

A right hemisphere dominance for bimanual grasps

To find points on the surface of an object that ensure a stable grasp, it would be most effective to employ one area in one cortical hemisphere. But grasping the object with both hands requires control through both hemispheres. To better understand the control mechanisms underlying this “bimanual grasping”, here we examined how the two hemispheres coordinate their control processes for bimanual grasping depending on visual field. We asked if bimanual grasping involves both visual fields equally or one more than the other. To test this, participants fixated either to the left or right of an object and then grasped or pushed it off a pedestal. We found that when participants grasped the object in the right visual field, maximum grip aperture (MGA) was larger and more variable, and participants were slower to react and to show MGA compared to when they grasped the object in the left visual field. In contrast, when participants pushed the object we observed no comparable visual field effects. These results suggest that grasping with both hands, specifically the computation of grasp points on the object, predominantly involves the right hemisphere. Our study provides new insights into the interactions of the two hemispheres for grasping.

The Right Anterior Intraparietal Sulcus Is Critical for Bimanual Grasping: A TMS Study

Grasping with 2 limbs in opposition to one another is older than the hand, yet the neural mechanisms for bimanual grasps remain unclear. Similar to unimanual grasping, bimanual grasping may require regions in the parietal cortex that use visual object-feature information to find matching stable grasp points on the object. The localization of matching points is computationally expensive, so it might make sense for the signals to converge in a single cortical area. To examine this, we use transcranial magnetic stimulation (TMS) to probe the contribution of cortical areas known to be associated with unimanual grasping, while participants performed bimanual grasps. We applied TMS to the anterior and caudal portion of the intra-parietal sulcus (aIPS and cIPS) in each hemisphere during a size-perturbation task using the index fingers of both hands to grasp an object whose orientation might or might not change. We found significant interaction effects between TMS and perturbation of the grasp-relevant object dimension that increased grip aperture only for the right aIPS. These results indicate that the aIPS is involved not only in unimanual, but also bimanual grasping, and the right aIPS is critically involved in bimanual grasps. This suggests that information from both hemispheres converges in the right hemisphere to achieve bimanual grasps.

A toggle switch of visual awareness

Major clues to the human brain mechanisms of spatial attention and visual awareness have come from the syndrome of neglect, where patients ignore one half of space. A longstanding puzzle, though, is that neglect almost always comes from right-hemisphere damage, which suggests that the two sides of the brain play distinct roles. But tests of attention in healthy people have revealed only slight differences between the hemispheres. Here we show that major differences emerge if we look at the timing of brain activity in a task optimized to identify attentional functions. Using EEG to map cortical activity on a millisecond timescale, we found transient (20-30 ms) periods of interhemispheric competition, followed by short phases of marked right-sided activity in the ventral attentional network. Our data are the first to show interhemispheric interactions that, much like a toggle switch, quickly allocate neural resources to one or the other hemisphere.

Left visual field preference for a bimanual grasping task with ecologically valid object sizes

Grasping using two forelimbs in opposition to one another is evolutionary older than the hand with an opposable thumb (Whishaw and Coles in Behav Brain Res 77:135–148, 1996); yet, the mechanisms for bimanual grasps remain unclear. Similar to unimanual grasping, the localization of matching stable grasp points on an object is computationally expensive and so it makes sense for the signals to converge in a single cortical hemisphere. Indeed, bimanual grasps are faster and more accurate in the left visual field, and are disrupted if there is transcranial stimulation of the right hemisphere (Le and Niemeier in Exp Brain Res 224:263–273, 2013; Le et al. in Cereb Cortex. doi:10.1093/cercor/bht115, 2013). However, research so far has tested the right hemisphere dominance based on small objects only, which are usually grasped with one hand, whereas bimanual grasping is more commonly used for objects that are too big for a single hand. Because grasping large objects might involve different neural circuits than grasping small objects (Grol et al. in J Neurosci 27:11877–11887, 2007), here we tested whether a left visual field/right hemisphere dominance for bimanual grasping exists with large and thus more ecologically valid objects or whether the right hemisphere dominance is a function of object size. We asked participants to fixate to the left or right of an object and to grasp the object with the index and middle fingers of both hands. Consistent with previous observations, we found that for objects in the left visual field, the maximum grip apertures were scaled closer to the object width and were smaller and less variable, than for objects in the right visual field. Our results demonstrate that bimanual grasping is predominantly controlled by the right hemisphere, even in the context of grasping larger objects.

Visual field preferences of object analysis for grasping with one hand.

When we grasp an object using one hand, the opposite hemisphere predominantly guides the motor control of grasp movements (Davare et al., 2007; Rice et al., 2007). However, it is unclear whether visual object analysis for grasp control relies more on inputs (a) from the contralateral than the ipsilateral visual field, (b) from one dominant visual field regardless of the grasping hand, or (c) from both visual fields equally. For bimanual grasping of a single object we have recently demonstrated a visual field preference for the left visual field (Le and Niemeier, 2013a,b), consistent with a general right-hemisphere dominance for sensorimotor control of bimanual grasps (Le et al., 2014). But visual field differences have never been tested for unimanual grasping. Therefore, here we asked right-handed participants to fixate to the left or right of an object and then grasp the object either with their right or left hand using a precision grip. We found that participants grasping with their right hand performed better with objects in the right visual field: maximum grip apertures (MGAs) were more closely matched to the object width and were smaller than for objects in the left visual field. In contrast, when people grasped with their left hand, preferences switched to the left visual field. What is more, MGA scaling with the left hand showed greater visual field differences compared to right-hand grasping. Our data suggest that, visual object analysis for unimanual grasping shows a preference for visual information from the ipsilateral visual field, and that the left hemisphere is better equipped to control grasps in both visual fields.

Parietal Area BA7 Integrates Motor Programs for Reaching, Grasping, and Bimanual Coordination

Skillful interaction with the world requires that the brain uses a multitude of sensorimotor programs and subroutines, such as for reaching, grasping, and the coordination of the two body halves. However, it is unclear how these programs operate together. Networks for reaching, grasping, and bimanual coordination might converge in common brain areas. For example, Brodmann area 7 (BA7) is known to activate in disparate tasks involving the three types of movements separately. Here, we asked whether BA7 plays a key role in integrating coordinated reach-to-grasp movements for both arms together. To test this, we applied transcranial magnetic stimulation (TMS) to disrupt BA7 activity in the left and right hemispheres, while human participants performed a bimanual size-perturbation grasping task using the index and middle fingers of both hands to grasp a rectangular object whose orientation (and thus grasp-relevant width dimension) might or might not change. We found that TMS of the right BA7 during object perturbation disrupted the bimanual grasp and transport/coordination components, and TMS over the left BA7 disrupted unimanual grasps. These results show that right BA7 is causally involved in the integration of reach-to-grasp movements of the two arms. NEW & NOTEWORTHY Our manuscript describes a role of human Brodmann area 7 (BA7) in the integration of multiple visuomotor programs for reaching, grasping, and bimanual coordination. Our results are the first to suggest that right BA7 is critically involved in the coordination of reach-to-grasp movements of the two arms. The results complement previous reports of right-hemisphere lateralization for bimanual grasps.