Gorana Pobric

Anterior temporal lobes mediate semantic representation: Mimicking semantic dementia by using rTMS in normal participants.

Studies of semantic dementia and PET neuroimaging investigations suggest that the anterior temporal lobes (ATL) are a critical substrate for semantic representation. In stark contrast, classical neurological models of comprehension do not include ATL, and likewise functional MRI studies often fail to show activations in the ATL, reinforcing the classical view. Using a novel application of low-frequency, repetitive transcranial magnetic stimulation (rTMS) over the ATL, we demonstrate that the behavioral pattern of semantic dementia can be mirrored in neurologically intact participants: Specifically, we show that temporary disruption to neural processing in the ATL produces a selective semantic impairment leading to significant slowing in both picture naming and word comprehension but not to other equally demanding, nonsemantic cognitive tasks.

Conceptual Knowledge Is Underpinned by the Temporal Pole Bilaterally: Convergent Evidence from rTMS

Conceptual knowledge provides the basis on which we bring meaning to our world. Studies of semantic dementia patients and some functional neuroimaging studies indicate that the anterior temporal lobes, bilaterally, are the core neural substrate for the formation of semantic representations. This hypothesis remains controversial, however, as traditional neurological models of comprehension do not posit a role for these regions. To adjudicate on this debate, we conducted 2 novel experiments that used off-line, low-frequency, repetitive transcranial magnetic stimulation to disrupt neural processing temporarily in the left or right temporal poles (TPs). The time required to make semantic decisions was slowed considerably, yet specifically, by this procedure. The results confirm that both TPs form a critical substrate within the neural network that supports conceptual knowledge.

Action Understanding Requires the Left Inferior Frontal Cortex

Numerous studies have established that inferior frontal cortex is active when hand actions are planned, imagined, remembered, imitated, and even observed. Furthermore, it has been proposed that these activations reflect a process of simulating the observed action to allow it to be understood and thus fully perceived. However, direct evidence for a perceptual role for left inferior frontal cortex is rare, and linguistic or motor contributions to the reported activations have not been ruled out. We used repetitive transcranial magnetic stimulation (rTMS) over inferior frontal gyrus during a perceptual weight-judgement task to test the hypothesis that this region contributes to action understanding. rTMS at this site impaired judgments of the weight of a box lifted by a person, but not judgements of the weight of a bouncing ball or of stimulus duration, and rTMS at control sites had no impact. This demonstrates that the integrity of left inferior frontal gyrus is necessary to make accurate perceptual judgments about other peoples actions.

Category-specific versus category-general semantic impairment induced by transcranial magnetic stimulation.

Summary Semantic cognition permits us to bring meaning to our verbal and nonverbal experiences and to generate context- and time-appropriate behavior [1–2]. It is core to language and nonverbal skilled behaviors and, when impaired after brain damage, it generates significant disability [3]. A fundamental neuroscience question is, therefore, how does the brain code and generate semantic cognition? Historical and some contemporary theories emphasize that conceptualization stems from the joint action of modality-specific association cortices (the “distributed” theory) [4, 5] reflecting our accumulated verbal, motor, and sensory experiences. Parallel studies of semantic dementia, rTMS in normal participants, and neuroimaging indicate that the anterior temporal lobe (ATL) plays a crucial and necessary role in conceptualization by merging experience into an amodal semantic representation [1, 2, 6–8]. Some contemporary computational models suggest that concepts reflect a hub-and-spoke combination of information—modality-specific association areas support sensory, verbal, and motor sources (the spokes) while anterior temporal lobes act as an amodal hub. We demonstrate novel and striking evidence in favor of this hypothesis by applying rTMS to normal participants: ATL stimulation generates a category-general impairment whereas IPL stimulation induces a category-specific deficit for man-made objects, reflecting the coding of praxis in this neural region.

The role of the anterior temporal lobes in the comprehension of concrete and abstract words: rTMS evidence

Conceptual knowledge allows us to bring meaning to our world. Studies of semantic dementia (SD) patients and some functional neuroimaging studies indicate that the anterior temporal lobes, bilaterally, are a core neural substrate for the formation of conceptual representations. The majority of SD patients (who have circumscribed atrophy of the anterior temporal lobes) have better comprehension of concrete than abstract words. However, this finding remains controversial, as some individual SD patients have exhibited reverse imageability effects, i.e., relative preservation of abstract knowledge. This would imply that the anterior temporal lobes are particularly crucial for processing sensory aspects of semantic knowledge, which are an important part of concrete but not abstract concepts. To adjudicate on this debate, we used offline, low-frequency, repetitive transcranial magnetic stimulation to disrupt neural processing temporarily in the left or right temporal poles (TPs). We examined this effect using a synonym judgement task, comprising high, medium and low imageability items, which we have previously employed with a case-series of SD patients. The time required to make semantic decisions was slowed considerably, particularly for low imageability items, consistent with the pattern we observed in SD. These results confirm that both TPs make a critical contribution to semantic processing, even for abstract concepts that do not have strong sensory representations.

The Nature and Neural Correlates of Semantic Association versus Conceptual Similarity

The ability to represent concepts and the relationships between them is critical to human cognition. How does the brain code relationships between items that share basic conceptual properties (e.g., dog and wolf) while simultaneously representing associative links between dissimilar items that co-occur in particular contexts (e.g., dog and bone)? To clarify the neural bases of these semantic components in neurologically intact participants, both types of semantic relationship were investigated in an fMRI study optimized for anterior temporal lobe (ATL) coverage. The clear principal finding was that the same core semantic network (ATL, superior temporal sulcus, ventral prefrontal cortex) was equivalently engaged when participants made semantic judgments on the basis of association or conceptual similarity. Direct comparisons revealed small, weaker differences for conceptual similarity > associative decisions (e.g., inferior prefrontal cortex) and associative > conceptual similarity (e.g., ventral parietal cortex) which appear to reflect graded differences in task difficulty. Indeed, once reaction time was entered as a covariate into the analysis, no associative versus category differences remained. The paper concludes with a discussion of how categorical/feature-based and associative relationships might be represented within a single, unified semantic system.

Posterior middle temporal gyrus is involved in verbal and non-verbal semantic cognition: Evidence from rTMS

Background: Left posterior middle temporal gyrus (pMTG) is reliably activated in functional neuroimaging studies of semantic processing and is frequently damaged in patients with comprehension impairments following stroke (e.g., Wernickes aphasia). Its precise function remains elusive, however. Some researchers take the view that pMTG is a multimodal semantic area, involved in verbal and non-verbal semantic cognition. Others ascribe a lexical-semantic function to the region, positing that it is involved in mapping between phonology and conceptual knowledge. Aims: We investigated whether pMTG was involved in non-verbal as well as verbal semantic cognition by using rTMS to induce temporary, focal “virtual lesions” to this region in healthy participants. Methods & Procedures: Participants completed picture and word versions of a semantic association test before and after receiving 10 minutes of 1-Hz offline rTMS to left pMTG. They also completed a difficulty-matched visual decision task on scrambled pictures. An occipital lobe control site was stimulated in a separate session. Outcomes & Results: TMS slowed responses to word and picture versions of the test to an equal degree. There was no slowing on a non-semantic visual-matching task, or following TMS to the control site. Conclusions: These results indicate that pMTG is involved in both verbal and non-verbal semantic cognition. This region could be key to understanding the multimodal semantic deficits often observed following stroke.

Functional representation of living and nonliving domains across the cerebral hemispheres: A combined event-related potential/transcranial magnetic stimulation study

Transcranial magnetic stimulation (TMS) over the left hemisphere has been shown to disrupt semantic processing but, to date, there has been no direct demonstration of the electrophysiological correlates of this interference. To gain insight into the neural basis of semantic systems, and in particular, study the temporal and functional organization of object categorization processing, we combined repetitive TMS (rTMS) and ERPs. Healthy volunteers performed a picture–word matching task in which Snodgrass drawings of natural (e.g., animal) and artifactual (e.g., tool) categories were associated with a word. When short trains of high-frequency rTMS were applied over Wernickes area (in the region of the CP5 electrode) immediately before the stimulus onset, we observed delayed response times to artifactual items, and thus, an increased dissociation between natural and artifactual domains. This behavioral effect had a direct ERP correlate. In the response period, the stimuli from the natural domain elicited a significant larger late positivity complex than those from the artifactual domain. These differences were significant over the centro-parietal region of the right hemisphere. These findings demonstrate that rTMS interferes with postperceptual categorization processing of natural and artifactual stimuli that involve separate subsystems in distinct cortical areas.

The timing of anterior temporal lobe involvement in semantic processing

Despite indications that regions within the anterior temporal lobe (ATL) might make a crucial contribution to pan-modal semantic representation, to date there have been no investigations of when during semantic processing the ATL plays a critical role. To test the timing of the ATL involvement in semantic processing, we studied the effect of double-pulse TMS on behavioral responses in semantic and difficulty-matched control tasks. Chronometric TMS was delivered over the left ATL (10 mm from the tip of the temporal pole along the middle temporal gyrus). During each trial, two pulses of TMS (40 msec apart) were delivered either at baseline (before stimulus presentation) or at one of the experimental time points 100, 250, 400, and 800 msec poststimulus onset. A significant disruption to performance was identified from 400 msec on the semantic task but not on the control assessment. Our results not only reinforce the key role of the left ATL in semantic representation but also indicate that its contribution is especially important around 400 msec poststimulus onset. Together, these facts suggest that the ATL may be one of the neural sources of the N400 ERP component.

The neural network for tool-related cognition: An activation likelihood estimation meta-analysis of 70 neuroimaging contrasts

ABSTRACT The ability to recognize and use a variety of tools is an intriguing human cognitive function. Multiple neuroimaging studies have investigated neural activations with various types of tool-related tasks. In the present paper, we reviewed tool-related neural activations reported in 70 contrasts from 56 neuroimaging studies and performed a series of activation likelihood estimation (ALE) meta-analyses to identify tool-related cortical circuits dedicated either to general tool knowledge or to task-specific processes. The results indicate the following: (a) Common, task-general processing regions for tools are located in the left inferior parietal lobule (IPL) and ventral premotor cortex; and (b) task-specific regions are located in superior parietal lobule (SPL) and dorsal premotor area for imagining/executing actions with tools and in bilateral occipito-temporal cortex for recognizing/naming tools. The roles of these regions in task-general and task-specific activities are discussed with reference to evidence from neuropsychology, experimental psychology and other neuroimaging studies.