Evidence implicates ventral parieto-premotor cortices in representing the goal of grasping independent of the movements or effectors involved [Umilta, M. A., Escola, L., Intskirveli, I., Grammont, F., Rochat, M., Caruana, F., et al. When pliers become fingers in the monkey motor system. Proceedings of the National Academy of Sciences, U.S.A., 105, 2209–2213, 2008; Tunik, E., Frey, S. H., & Grafton, S. T. Virtual lesions of the anterior intraparietal area disrupt goal-dependent on-line adjustments of grasp. Nature Neuroscience, 8, 505–511, 2005]. Modern technologies that enable arbitrary causal relationships between hand movements and tool actions provide a strong test of this hypothesis. We capitalized on this unique opportunity by recording activity with fMRI during tasks in which healthy adults performed goal-directed reach and grasp actions manually or by depressing buttons to initiate these same behaviors in a remotely located robotic arm (arbitrary causal relationship). As shown previously [Binkofski, F., Dohle, C., Posse, S., Stephan, K. M., Hefter, H., Seitz, R. J., et al. Human anterior intraparietal area subserves prehension: A combined lesion and functional MRI activation study. Neurology, 50, 1253–1259, 1998], we detected greater activity in the vicinity of the anterior intraparietal sulcus (aIPS) during manual grasp versus reach. In contrast to prior studies involving tools controlled by nonarbitrarily related hand movements [Gallivan, J. P., McLean, D. A., Valyear, K. F., & Culham, J. C. Decoding the neural mechanisms of human tool use. Elife, 2, e00425, 2013; Jacobs, S., Danielmeier, C., & Frey, S. H. Human anterior intraparietal and ventral premotor cortices support representations of grasping with the hand or a novel tool. Journal of Cognitive Neuroscience, 22, 2594–2608, 2010], however, responses within the aIPS and premotor cortex exhibited no evidence of selectivity for grasp when participants employed the robot. Instead, these regions showed comparable increases in activity during both the reach and grasp conditions. Despite equivalent sensorimotor demands, the right cerebellar hemisphere displayed greater activity when participants initiated the robot's actions versus when they pressed a button known to be nonfunctional and watched the very same actions undertaken autonomously. This supports the hypothesis that the cerebellum predicts the forthcoming sensory consequences of volitional actions [Blakemore, S. J., Frith, C. D., & Wolpert, D. M. The cerebellum is involved in predicting the sensory consequences of action. NeuroReport, 12, 1879–1884, 2001]. We conclude that grasp-selective responses in the human aIPS and premotor cortex depend on the existence of nonarbitrary causal relationships between hand movements and end-effector actions.

Humans display a remarkable capacity to use tools instead of their biological effectors. Yet, little is known about the mechanisms that support these behaviors. Here, participants learned to grasp objects, appearing in a variety of orientations, with a novel, handheld mechanical tool. Following training, psychophysical functions relating grip preferences (i.e., pronated vs. supinated) to stimulus orientations indicate a reliance on distinct, effector-specific internal representations when planning grasping actions on the basis of the tool versus the hands. Accompanying fMRI data show that grip planning in both hand and tool conditions was associated with similar increases in activity within the same regions of the anterior intraparietal and caudal ventral premotor cortices, a putative homologue of the macaque anterior intraparietal–ventral premotor (area F5) “grasp circuit.” These findings suggest that tool use is supported by effector-specific representations of grasping with the tool that are functionally independent of previously existing representations of the hand and yet occur within the same parieto-frontal regions involved in manual prehension. These levels of representation are critical for accurate planning and execution of actions in a manner that is sensitive to the respective properties of these effectors. These effector-specific representations likely coexist with effector-independent representations. The latter were recently reported in macaque F5 [Umiltà, M. A., Escola, L., Intskirveli, I., Grammont, F., Rochat, M., Caruana, F., et al. When pliers become fingers in the monkey motor system. Proceedings of the National Academy of Sciences, U.S.A., 105, 2209–2213, 2008] and appear to be established by tool use training through modification of existing representations of grasping with the hand. These more abstract levels of representation may facilitate the transfer of skills between hand and tool.

Longitudinal changes in brain activity during second language (L2) acquisition of a miniature finite-state grammar, named Wernickese, were identified with functional magnetic resonance imaging (fMRI). Participants learned either a visual sign language form or an auditory-verbal form to equivalent proficiency levels. Brain activity during sentence comprehension while hearing/viewing stimuli was assessed at low, medium, and high levels of proficiency in three separate fMRI sessions. Activation in the left inferior frontal gyrus (Broca's area) correlated positively with improving L2 proficiency, whereas activity in the right-hemisphere (RH) homologue was negatively correlated for both auditory and visual forms of the language. Activity in sequence learning areas including the premotor cortex and putamen also correlated with L2 proficiency. Modality-specific differences in the blood oxygenation level-dependent signal accompanying L2 acquisition were localized to the planum temporale (PT). Participants learning the auditory form exhibited decreasing reliance on bilateral PT sites across sessions. In the visual form, bilateral PT sites increased in activity between Session 1 and Session 2, then decreased in left PT activity from Session 2 to Session 3. Comparison of L2 laterality (as compared to L1 laterality) in auditory and visual groups failed to demonstrate greater RH lateralization for the visual versus auditory L2. These data establish a common role for Broca's area in language acquisition irrespective of the perceptual form of the language and suggest that L2s are processed similar to first languages even when learned after the “critical period.” The right frontal cortex was not preferentially recruited by visual language after accounting for phonetic/structural complexity and performance.

The overwhelming majority of evidence indicates that the left cerebral hemisphere of right-handed humans is dominant both for manual control and the representation of acquired skills, including tool use. It is, however, unclear whether these functions involve common or dissociable mechanisms. Here we demonstrate that the disconnected left hemispheres of both right- and left-handed split-brain patients are specialized for representing acquired tool-use skills. When required to pantomime actions associated with familiar tools (Experiment 2), both patients show a right-hand (left hemisphere) advantage in response to tool names, pictures, and actual objects. Accuracy decreases as stimuli become increasingly symbolic when using the left hand (right hemisphere). Tested in isolation with lateralized pictures (Experiment 3), each patient's left hemisphere demonstrates a significant advantage over the right hemisphere for pantomiming tool-use actions with the contralateral hand. The fact that this asymmetry occurs even in a left-handed patient suggests that the left hemisphere specialization for representing praxis skills can be dissociated from mechanisms involved in hand dominance located in the right hemisphere. This effect is not attributable to differences at the conceptual level, as the left and right hemispheres are equally and highly competent at associating tools with observed pantomimes (Experiment 4).