Patricia M. Gough

Dissociating linguistic processes in the left inferior frontal cortex with transcranial magnetic stimulation

Is the left inferior frontal cortex (LIFC) a single functional region, or can it be subdivided into distinct areas that contribute differently to word processing? Here we used transcranial magnetic stimulation (TMS) to investigate anterior and posterior LIFC when the meaning and sound of words were being processed. Relative to no stimulation, TMS of the anterior LIFC selectively increased response latencies when participants focused on the meaning of simultaneously presented words (i.e., synonym judgments) but not when they focused on the sound pattern of the words (i.e., homophone judgments). In contrast, the opposite dissociation was observed in the posterior LIFC, where stimulation selectively interfered with the phonological but not the semantic task. This double dissociation shows functionally distinct subdivisions of the LIFC that can be understood in terms of separable corticocortical connections linking the anterior LIFC to temporal pole regions associated with semantic memory and the posterior LIFC to temporoparietal regions involved in auditory speech processing.

The Inferior Frontal Gyrus and Phonological Processing: An Investigation using rTMS

Repetitive transcranial magnetic stimulation (rTMS) offers a powerful new technique for investigating the distinct contributions of the cortical language areas. We have used this method to examine the role of the left inferior frontal gyrus (IFG) in phonological processing and verbal working memory. Functional neuroimaging studies have implicated the posterior part of the left IFG in both phonological decision making and subvocal rehearsal mechanisms, but imaging is a correlational method and it is therefore necessary to determine whether this region is essential for such processes. In this paper we present the results of two experiments in which rTMS was applied over the frontal operculum while subjects performed a delayed phonological matching task. We compared the effects of disrupting this area either during the delay (memory) phase or at the response (decision) phase of the task. Delivered at a time when subjects were required to remember the sound of a visually presented word, rTMS impaired the accuracy with which they subsequently performed the task. However, when delivered later in the trial, as the subjects compared the remembered word with a given pseudoword, rTMS did not impair accuracy. Performance by the same subjects on a control task that required the processing of nonverbal visual stimuli was unaffected by the rTMS. Similarly, performance on both tasks was unaffected by rTMS delivered over a more anterior site (pars triangularis). We conclude that the opercular region of the IFG is necessary for the normal operation of phonologically based working memory mechanisms. Furthermore, this study shows that rTMS can shed further light on the precise role of cortical language areas in humans.

Supramarginal gyrus involvement in visual word recognition.

INTRODUCTION In the classic neurological model of language, the human inferior parietal lobule (IPL) plays an important role in visual word recognition. The region is both functionally and structurally heterogeneous, however, suggesting that subregions of IPL may differentially contribute to reading. The two main sub-divisions are the supramarginal (SMG) and angular gyri, which have been hypothesized to contribute preferentially to phonological and semantic aspects of word processing, respectively. METHODS Here we used single-pulse transcranial magnetic stimulation (TMS) to investigate the functional specificity and timing of SMG involvement in reading. Participants performed two reading tasks that focused attention on either the phonological or semantic relation between two simultaneously presented words. A third task focused attention on the visual relation between pairs of consonant letter strings to control for basic input and output characteristics of the paradigm using non-linguistic stimuli. TMS to SMG was delivered on every trial at 120, 180, 240 or 300 msec post-stimulus onset. RESULTS Stimulation at 180 msec produced a reliable facilitation of reaction times for both the phonological and semantic tasks, but not for the control visual task. CONCLUSION These findings demonstrate that SMG contributes to reading regardless of the specific task demands, and suggests this may be due to automatically computing the sound of a word even when the task does not explicitly require it.

Attentional selection and action selection in the ventral and orbital prefrontal cortex.

Different accounts of the ventral and orbital prefrontal cortex (PFv+o) have emphasized either its role in learning conditional rules for action selection or the attentional selection of behaviorally relevant stimuli. Although the accounts are not mutually exclusive, it is possible that the involvement of PFv+o in conditional action selection is a consequence of its role in selecting relevant stimuli or that its involvement in attentional selection is a consequence of the conditional rules present in many attentional paradigms. Five macaques learned a conditional action-selection task in which the difficulty of identifying the stimulus relevant for guiding action selection was varied in a simple manner by either altering its distance from the action or presenting additional distracting stimuli. Simply increasing the spatial separation between the instructing stimulus led to slower responses. Experiment 1 showed that bilateral PFv+o lesions impaired conditional action selection even when attentional demands were kept to a minimum, but there was evidence that the impairment was exacerbated by manipulating stimulus selection difficulty. Experiment 2 confirmed the importance of PFv+o for conditional action selection even when stimulus selection difficulty was minimal. Experiments 3 and 4 demonstrated that the action-selection impairment was significantly increased by making identification of the behaviorally relevant stimulus difficult. PFv+o is central to the use of conditional rules when selecting courses of action, but conditional rules are also represented in premotor and striatal regions. A special contribution of PFv+o may be initial selection of behaviorally relevant stimuli.

Action observation versus motor imagery in learning a complex motor task: A short review of literature and a kinematics study

Both motor imagery and action observation have been shown to play a role in learning or re-learning complex motor tasks. According to a well accepted view they share a common neurophysiological basis in the mirror neuron system. Neurons within this system discharge when individuals perform a specific action and when they look at another individual performing the same or a motorically related action. In the present paper, after a short review of literature on the role of action observation and motor imagery in motor learning, we report the results of a kinematics study where we directly compared motor imagery and action observation in learning a novel complex motor task. This involved movement of the right hand and foot in the same angular direction (in-phase movement), while at the same time moving the left hand and foot in an opposite angular direction (anti-phase movement), all at a frequency of 1Hz. Motor learning was assessed through kinematics recording of wrists and ankles. The results showed that action observation is better than motor imagery as a strategy for learning a novel complex motor task, at least in the fast early phase of motor learning. We forward that these results may have important implications in educational activities, sport training and neurorehabilitation.

Improving upper limb motor functions through action observation treatment: a pilot study in children with cerebral palsy

Aim  The aim of this randomized controlled trial was to assess whether action observation treatment may improve upper limb motor functions in children with cerebral palsy (CP).

Cortico-basal ganglia pathways are essential for the recall of well-established visuomotor associations.

Recent human neuroimaging studies, supported by lesion studies with nonhuman primates, have suggested that learning arbitrary associations between sensory cues and behavioural responses requires interactions between the infero‐temporal, prefrontal and premotor cortices. We directly tested the hypothesis suggested from our neuroimaging experiments that functional links between the basal ganglia and premotor cortex are involved in the process via which task performance becomes automatic. We made unilateral excitotoxic lesions, centred on the internal pallidum, in four macaques previously given extensive experience on the associations between nonspatial visual cues and movements of a joystick. The basal ganglia lesion was later combined with a premotor cortical lesion in the opposite hemisphere so as to interrupt the connections between them. Three of the animals were subsequently found to be impaired in relearning pre‐operatively acquired associations; they eventually succeeded but made three‐times as many errors. A fourth animal was unimpaired but its premotor cortex lesion was later found to be incomplete. Response times were only marginally increased and the learning of novel associations appeared relatively unaffected by these lesions. As a control, the effects of a unilateral premotor cortex lesion were assessed with two additional animals but this lesion did not result in a relearning impairment. We therefore suggest that when visuomotor associations have become well established through over‐training, performance depends on connections between the basal ganglia and premotor cortex.

How the motor system handles nouns: a behavioral study

It is an open question whether the motor system is involved during understanding of concrete nouns, as it is for concrete verbs. To clarify this issue, we carried out a behavioral experiment using a go-no go paradigm with an early and delayed go-signal delivery. Italian nouns referring to concrete objects (hand-related or foot-related) and abstract entities served as stimuli. Right-handed participants read the stimuli and responded when the presented word was concrete using the left or right hand. At the early go-signal, slower right-hand responses were found for hand-related nouns compared to foot-related nouns. The opposite pattern was found for the left hand. These findings demonstrate an early lateralized modulation of the motor system during noun processing, most likely crucial for noun comprehension.

Nouns referring to tools and natural objects differentially modulate the motor system

While increasing evidence points to a critical role for the motor system in language processing, the focus of previous work has been on the linguistic category of verbs. Here we tested whether nouns are effective in modulating the motor system and further whether different kinds of nouns - those referring to artifacts or natural items, and items that are graspable or ungraspable - would differentially modulate the system. A Transcranial Magnetic Stimulation (TMS) study was carried out to compare modulation of the motor system when subjects read nouns referring to objects which are Artificial or Natural and which are Graspable or Ungraspable. TMS was applied to the primary motor cortex representation of the first dorsal interosseous (FDI) muscle of the right hand at 150 ms after noun presentation. Analyses of Motor Evoked Potentials (MEPs) revealed that across the duration of the task, nouns referring to graspable artifacts (tools) were associated with significantly greater MEP areas. Analyses of the initial presentation of items revealed a main effect of graspability. The findings are in line with an embodied view of nouns, with MEP measures modulated according to whether nouns referred to natural objects or artifacts (tools), confirming tools as a special class of items in motor terms. Additionally our data support a difference for graspable versus non graspable objects, an effect which for natural objects is restricted to initial presentation of items.

Fine tuned modulation of the motor system by adjectives expressing positive and negative properties

Using transcranial magnetic stimulation (TMS), motor evoked potentials (MEPs) were recorded from two antagonistic muscles, the first dorsal interosseus (FDI) of the hand and the extensor communis digitorum (EC) of the forearm. FDI is involved in grasping actions and EC in releasing. TMS pulses were delivered while participants were reading adjectives expressing either negative or positive pragmatic properties, at 150 ms after presentation of language material. Overall findings showed an interaction of adjective type (positive, negative) and muscle (FDI, EC), the effect being driven by a significant difference for negative adjectives. Further analysis aimed at investigating the effectiveness of positive adjectives showed a similar, but opposite, pattern of effects for the positive words in the initial two blocks. The present results indicate that, as for verbs and nouns, adjectives recruit the sensorimotor system, and their processing is best explained by an embodiment rather than an amodal approach to language.