Jennifer M. Rodd

Processing Objects at Different Levels of Specificity

How objects are represented and processed in the brain is a central topic in cognitive neuroscience. Previous studies have shown that knowledge of objects is represented in a featurebased distributed neural system primarily involving occipital and temporal cortical regions. Research with nonhuman primates suggest that these features are structured in a hierarchical system with posterior neurons in the inferior temporal cortex representing simple features and anterior neurons in the perirhinal cortex representing complex conjunctions of features (Bussey & Saksida, 2002; Murray & Bussey, 1999). On this account, the perirhinal cortex plays a crucial role in object identification by integrating information from different sensory systems into more complex polymodal feature conjunctions. We tested the implications of these claims for human object processing in an event-related fMRI study in which we presented colored pictures of common objects for 19 subjects to name at two levels of specificity-basic and domain. We reasoned that domain-level naming requires access to a coarsergrained representation of objects, thus involving only posterior regions of the inferior temporal cortex. In contrast, basic-level naming requires finer-grained discrimination to differentiate between similar objects, and thus should involve anterior temporal regions, including the perirhinal cortex. We found that object processing always activated the fusiform gyrus bilaterally, irrespective of the task, whereas the perirhinal cortex was only activated when the task required finer-grained discriminations. These results suggest that the same kind of hierarchical structure, which has been proposed for object processing in the monkey temporal cortex, functions in the human.

Dissociating speech perception and comprehension at reduced levels of awareness.

We used functional MRI and the anesthetic agent propofol to assess the relationship among neural responses to speech, successful comprehension, and conscious awareness. Volunteers were scanned while listening to sentences containing ambiguous words, matched sentences without ambiguous words, and signal-correlated noise (SCN). During three scanning sessions, participants were nonsedated (awake), lightly sedated (a slowed response to conversation), and deeply sedated (no conversational response, rousable by loud command). Bilateral temporal-lobe responses for sentences compared with signal-correlated noise were observed at all three levels of sedation, although prefrontal and premotor responses to speech were absent at the deepest level of sedation. Additional inferior frontal and posterior temporal responses to ambiguous sentences provide a neural correlate of semantic processes critical for comprehending sentences containing ambiguous words. However, this additional response was absent during light sedation, suggesting a marked impairment of sentence comprehension. A significant decline in postscan recognition memory for sentences also suggests that sedation impaired encoding of sentences into memory, with left inferior frontal and temporal lobe responses during light sedation predicting subsequent recognition memory. These findings suggest a graded degradation of cognitive function in response to sedation such that “higher-level” semantic and mnemonic processes can be impaired at relatively low levels of sedation, whereas perceptual processing of speech remains resilient even during deep sedation. These results have important implications for understanding the relationship between speech comprehension and awareness in the healthy brain in patients receiving sedation and in patients with disorders of consciousness.

Residual auditory function in persistent vegetative state: a combined pet and fmri study

In recent years, a number of studies have demonstrated an important role for functional neuroimaging in the identification of residual cognitive function in persistent vegetative state. Such studies, when successful, may be particularly useful where there is concern about the accuracy of the diagnosis and the possibility that residual cognitive function has remained undetected. Unfortunately, functional neuroimaging in persistent vegetative state is extremely complex and subject to numerous methodological, clinical and theoretical difficulties. Here, we describe the strategy used to study residual auditory and speech processing in a single patient with a clinical diagnosis of persistent vegetative state. Identical positron emission tomography studies, conducted nine months apart, revealed preserved and consistent responses in predicted regions of auditory cortex in response to intelligible speech stimuli. Moreover, a preliminary functional magnetic resonance imaging examination at the time of the second session revealed partially intact responses to semantically ambiguous stimuli, which are known to tap higher aspects of speech comprehension. In spite of the multiple logistic and procedural problems involved, these results have major clinical and theoretical implications and provide a strong basis for the systematic study of possible residual cognitive function in patients diagnosed as being in a persistent vegetative state.

Distinctiveness and correlation in conceptual structure: behavioral and computational studies.

Patients with category-specific deficits have motivated a range of hypotheses about the structure of the conceptual system. One class of models claims that apparent category dissociations emerge from the internal structure of concepts rather than fractionation of the system into separate substores. This account claims that distinctive properties of concepts in the living domain are vulnerable because of their weak correlation with other features. Given the assumption that mutual activation among correlated properties produces faster activation in the normal system, the authors predicted a disadvantage for the distinctive features of living things for unimpaired adults. Results of a speeded feature verification study supported this prediction, as did a computational simulation in which networks mapped from orthography to semantics.

The Functional Organisation of the Fronto-Temporal Language System: Evidence from Syntactic and Semantic Ambiguity.

Spoken language comprehension is known to involve a large left-dominant network of fronto-temporal brain regions, but there is still little consensus about how the syntactic and semantic aspects of language are processed within this network. In an fMRI study, volunteers heard spoken sentences that contained either syntactic or semantic ambiguities as well as carefully matched low-ambiguity sentences. Results showed ambiguity-related responses in the posterior left inferior frontal gyrus (pLIFG) and posterior left middle temporal regions. The pLIFG activations were present for both syntactic and semantic ambiguities suggesting that this region is not specialised for processing either semantic or syntactic information, but instead performs cognitive operations that are required to resolve different types of ambiguity irrespective of their linguistic nature, for example by selecting between possible interpretations or reinterpreting misparsed sentences. Syntactic ambiguities also produced activation in the posterior middle temporal gyrus. These data confirm the functional relationship between these two brain regions and their importance in constructing grammatical representations of spoken language.

Why Clowns Taste Funny: The Relationship between Humor and Semantic Ambiguity

What makes us laugh? One crucial component of many jokes is the disambiguation of words with multiple meanings. In this functional MRI study of normal participants, the neural mechanisms that underlie our experience of getting a joke that depends on the resolution of semantically ambiguous words were explored. Jokes that contained ambiguous words were compared with sentences that contained ambiguous words but were not funny, as well as to matched verbal jokes that did not depend on semantic ambiguity. The results confirm that both the left inferior temporal gyrus and left inferior frontal gyrus are involved in processing the semantic aspects of language comprehension, while a more widespread network that includes both of these regions and the temporoparietal junction bilaterally is involved in processing humorous verbal jokes when compared with matched nonhumorous material. In addition, hearing jokes was associated with increased activity in a network of subcortical regions, including the amygdala, the ventral striatum, and the midbrain, that have been implicated in experiencing positive reward. Moreover, activity in these regions correlated with the subjective ratings of funniness of the presented material. These results allow a more precise account of how the neural and cognitive processes that are involved in ambiguity resolution contribute to the appreciation of jokes that depend on semantic ambiguity.

Dissociating Frontotemporal Contributions to Semantic Ambiguity Resolution in Spoken Sentences

Comprehension of sentences containing semantically ambiguous words requires listeners to select appropriate interpretations, maintain linguistic material in working memory, and to reinterpret sentences that have been misinterpreted. All these functions appear to involve frontal cortical regions. Here, we attempt to differentiate these functions by varying the relative timing of an ambiguous word and disambiguating information in spoken sentences. We compare the location, magnitude, and timing of evoked activity using a fast-acquisition semisparse functional magnetic resonance imaging sequence. The left inferior frontal gyrus (LIFG) shows a strong response to sentences that are initially ambiguous (disambiguated by information that occurs either soon after the ambiguity or that is delayed until the end of the sentence). Response profiles indicate that activity, in both anterior and posterior LIFG regions, is triggered both by the ambiguous word and by the subsequent disambiguating information. The LIFG also responds to ambiguities that are preceded by disambiguating context. These results suggest that the LIFG subserves multiple cognitive processes including selecting an appropriate meaning and reinterpreting sentences that have been misparsed. In contrast, the left inferior temporal gyrus responds to the disambiguating information but not to the ambiguous word itself and may be involved in reprocessing sentences that were initially misinterpreted.

When do leotards get their spots? Semantic activation of lexical neighbors in visual word recognition.

Shadowing and priming studies have provided strong evidence that during spoken word recognition, the meanings of different words that share their onset (e.g.,captain andcaptive) are activated in parallel. In contrast, for visual word recognition, there is little evidence that the meanings of visually similar words are activated in parallel. This is consistent with the idea that for reading (in contrast to listening), since all the sensory information necessary to identify a word is available at once, any competition between visually similar words is resolved before their meanings are retrieved. However, Forster and Hector (2002) have recently shown that for nonwords (e.g.,turple), some aspect of the meanings of lexical neighbors (e.g.,turtle) can be activated. However, this finding is limited to nonwords. The activation ofturtle’s meaning in response toturple may occur becauseturple has no meaning. In normal reading, we do not encounter nonwords, and there is strong pressure on the reading system to produce meaningful representations for every word (even misspelled words). The two semantic categorization experiments reported here extend this finding to real words. Participants are slower to decide thatleotard is not an animal because of its animal neighborleopard. This shows that information about a word’s meaning can be available before it has been uniquely recognized.

The role of domain-general frontal systems in language comprehension: Evidence from dual-task interference and semantic ambiguity

Neuroimaging studies have shown that the left inferior frontal gyrus (LIFG) plays a critical role in semantic and syntactic aspects of speech comprehension. It appears to be recruited when listeners are required to select the appropriate meaning or syntactic role for words within a sentence. However, this region is also recruited during tasks not involving sentence materials, suggesting that the systems involved in processing ambiguous words within sentences are also recruited for more domain-general tasks that involve the selection of task-relevant information. We use a novel dual-task methodology to assess whether the cognitive system(s) that are engaged in selecting word meanings are also involved in non-sentential tasks. In Experiment 1, listeners were slower to decide whether a visually presented letter is in upper or lower case when the sentence that they are simultaneously listening to contains words with multiple meanings (homophones), compared to closely matched sentences without homophones. Experiment 2 indicates that this interference effect is not tied to the occurrence of the homophone itself, but rather occurs when listeners must reinterpret a sentence that was initially misparsed. These results suggest some overlap between the cognitive system involved in semantic disambiguation and the domain-general process of response selection required for the case-judgement task. This cognitive overlap may reflect neural overlap in the networks supporting these processes, and is consistent with the proposal that domain-general selection processes in inferior frontal regions are critical for language comprehension.

Localising semantic and syntactic processing in spoken and written language comprehension: An Activation Likelihood Estimation meta-analysis

We conducted an Activation Likelihood Estimation (ALE) meta-analysis to identify brain regions that are recruited by linguistic stimuli requiring relatively demanding semantic or syntactic processing. We included 54 functional MRI studies that explicitly varied the semantic or syntactic processing load, while holding constant demands on earlier stages of processing. We included studies that introduced a syntactic/semantic ambiguity or anomaly, used a priming manipulation that specifically reduced the load on semantic/syntactic processing, or varied the level of syntactic complexity. The results confirmed the critical role of the posterior left Inferior Frontal Gyrus (LIFG) in semantic and syntactic processing. These results challenge models of sentence comprehension highlighting the role of anterior LIFG for semantic processing. In addition, the results emphasise the posterior (but not anterior) temporal lobe for both semantic and syntactic processing.