FMRI of two measures of phonological processing in visual word recognition: ecological validity matters

Previous functional magnetic resonance imaging (fMRI) studies have investigated the role of phonological processing by utilizing nonword rhyming decision tasks (e.g., Pugh et al., 1996). Although such tasks clearly engage phonological components of visual word recognition, it is clear that decision tasks are more cognitively involved than the simple overt naming tasks, which more closely map onto normal reading behavior. Our research aim for this study was to examine the advantages of overt naming tasks for fMRI studies of word recognition processes. Process models are presented to highlight the similarities and differences between two cognitive tasks that are used in the word recognition literature, pseudohomophone naming (e.g., pronounce BRANE) and rhyming decision (e.g., do LEAT and JEAT rhyme?). An fMRI study identified several differences in cortical activation associated with the differences observed in the process models. Specifically, the results show that the overt naming task involved the insular cortex and inferior frontal gyrus, whereas the rhyming decision task engaged the temporal-parietal regions. It is argued that future fMRI research examining the neuroanatomical components of basic visual word recognition utilize overt naming tasks.     

On the role of word frequency in the detection of component letters

The experiments reported in this study were conducted to explore the issue of race models versus holistic models of word processing. In both types of model, it is assumed that an available word-level encoding for a display will conceal letter information, and thereby inhibit component-letter detection. However, whereas in holistic models it is assumed that encoding always should occur at the word or pattern level first, in the race models it is assumed that encoding occurs at all levels (e.g., feature, letter, and word) simultaneously, with the final level of encoding being at whatever level has been completed first. If the rate of word-level encoding is facilitated by increasing word frequency, the holistic models predict a generally declining latency for letter detection, because the initial step in letter detection (i.e., word-level encoding) will be occurring more rapidly. The race models, on the other hand, predict that with increasing word frequency there will be an increasing chance that the word-level encoding will win the encoding race, resulting in an increase in the latency for letter detection (i.e., the word code will conceal the letter codes). Two experiments are reported, and the obtained pattern of latency data appears to be most consistent with the race models.     

Interactive visual and postvisual processes and their roles in form-specific memory.

Effects of depth of encoding on form-specific memory were examined. After viewing words (e.g., "bear") presented centrally during initial encoding, participants completed word stems (e.g., "BEA") presented laterally and pattern masked during subsequent test. When the encoding task was perceptual, letter-case specific memory was not observed, unlike in previous experiments without pattern masking. However, when the encoding task required both perceptual and conceptual processing, letter-case specific memory was observed in direct right-hemisphere, but not in direct left-hemisphere, test presentations, like in previous studies without pattern masking. Results were not influenced by whether stems were completed to form the first words that came to mind or words explicitly retrieved from encoding. Depth of encoding may influence form-specific memory through interactive processing of visual and postvisual information.     

Imaging a science of mind

I believe that the study of neuroimaging has supported localization of mental operations within the human brain. Most studies have shown a small number of widely distributed brain areas that must be orchestrated to carry out a cognitive task. Although, as in all sciences, there are disagreements, the convergence of results in areas of attention and language in particular seem impressive. Moreover, the anatomical data has helped us to specify the computations that are used by the brain to carry out cognitive tasks. Building upon localization of cognitive operations, imaging methods are being applied to studies of the circuitry, plasticity and individual development of neural networks. Working together with cellular and genetic methods, there is movement towards a more unified view of the role of the human brain in supporting the mind.     

Electrophysiological evidence of different loci for case-mixing and word frequency effects in visual word recognition

Do word frequency and case mixing affect different processing stages in visual word recognition? Some studies of online reading have suggested that word frequency affects an earlier, perceptual-encoding stage and that case mixing affects a later, central decision stage (e.g., Reingold, Yang, & Rayner, Journal of Experimental Psychology: Human Perception and Performance 36:1677-1683 2010); others have suggested otherwise (e.g., Allen, Smith, Lien, Grabbe, & Murphy, Journal of Experimental Psychology: Human Perception and Performance 31:713-721 2005; Besner & McCann, 1987). To determine the locus of the word frequency and case-mixing effects, we manipulated word frequency (high vs. low) and case type (consistent lower case vs. mixing case) in a lexical-decision paradigm. We measured two event-related-potential components: the N170 (an early peak occurring 140-240 ms after stimulus onset, related to structural encoding) and the P3 (a late peak occurring 400-600 ms after stimulus onset, related to stimulus categorization). The critical finding was that the N170 amplitude was sensitive to case mixing, but the P3 amplitude was sensitive to word frequency and lexicality. These results suggest that case mixing affects an earlier processing stage than does word frequency, at least with respect to lexical-decision processes.     

The sound of enemies and friends in the neighborhood

Previous studies (e.g., Pecher, Zeelenberg, & Wagenmakers, 2005) found that semantic classification performance is better for target words with orthographic neighbors that are mostly from the same semantic class (e.g., living) compared to target words with orthographic neighbors that are mostly from the opposite semantic class (e.g., nonliving). In the present study we investigated the contribution of phonology to orthographic neighborhood effects by comparing effects of phonologically congruent orthographic neighbors (book-hook) to phonologically incongruent orthographic neighbors (sand-wand). The prior presentation of a semantically congruent word produced larger effects on subsequent animacy decisions when the previously presented word was a phonologically congruent neighbor than when it was a phonologically incongruent neighbor. In a second experiment, performance differences between target words with versus without semantically congruent orthographic neighbors were larger if the orthographic neighbors were also phonologically congruent. These results support models of visual word recognition that assume an important role for phonology in cascaded access to meaning.     

Simple arithmetic processing: surface form effects in a priming task

Models of numerical processing vary on whether they assume common or separate processing pathways for problems represented in different surface forms. The present study employed a priming procedure, with target naming task, in an investigation of surface form effects in simple addition and multiplication operations. Participants were presented with Arabic digit and number word problems in one of three prime-target relationships, including congruent (e.g., '2+3' and '5'), incongruent (e.g., '9+7' and '5') and neutral (e.g., 'X+Y' and '5') conditions. The results revealed significant facilitatory effects in response to congruent digit stimuli at SOAs of 300 and 1000ms, in both operations. In contrast, inhibitory effects were observed in response to incongruent word stimuli in both the addition and multiplication operations at 300ms, and in the addition operation at 1000ms. The overall priming effects observed in the digit condition were significantly greater than in the word condition at 1000ms in the multiplication operation and at 300ms in the addition operation. The results provide support to separate pathway accounts of simple arithmetic processing for problems represented in different surface forms. An explanation for variation in processing due to differences in access to visual and phonological representations is provided.     

The unification of mind: Integration of hemispheric semantic processing

Seventy-six participants performed a visual half-field lexical decision task at two different stimulus onset asynchronies (50 or 750 ms). Word targets were primed either by a highly associated word (e.g., CLEAN-DIRTY), a weakly associated word (e.g., CLEAN-TIDY), or an unrelated word (e.g., CLEAN-FAMILY) projected to either the same or opposite visual field (VF) as the target. In the short SOA, RVF-left hemisphere primes resulted in high associate priming regardless of target location (ipsilateral or contralateral to the prime) whereas LVF-right hemisphere primes produced both high and low associate priming across both target location conditions. In the long SOA condition, contralateral priming patterns converged, demonstrating only high associate priming in both VF locations. The results of this study demonstrate the critical role of interhemispheric transfer in semantic processing and indicate a need to elaborate current models of semantic processing.     

Influence of imaging ability on word transformation

In the present study, we examined whether individual differences in imaging ability affect visual word recognition. Poor and vivid imagers performed a naming task that involved nonreversed (e.g., JUMP) and reversed (e.g., PMUJ) words (Experiment 1). Poor and vivid imagers were also tested on a naming task that was controlled for verbal ability; all the words were reversed and presentation time was varied (Experiment 2). In both experiments, imaging ability interacted with task difficulty, suggesting that individual differences in imaging ability affect visual word recognition. Specifically, the present data suggest that poor imagers may be less efficient than vivid imagers at processing words analytically. The data are interpreted within a limited-capacity, hybrid, word recognition model, in which words can be processed as either word-level or letter-level codes.     

Syntactic processing in the human brain: what we know, what we don't know, and a suggestion for how to proceed

For every claim in the neuroimaging literature about a particular brain region supporting syntactic processing, there exist other claims implicating the target region in different linguistic processes, and, in many cases, in non-linguistic cognitive processes (e.g., Blumstein, 2009). We argue that traditional group analysis methods in neuroimaging may obscure functional specificity because of inter-subject anatomical variability (Fedorenko & Kanwisher, 2009). In Fedorenko, Hsieh, Nieto-Castanon, Whitfield-Gabrieli, and Kanwisher (2010) we presented a functional localizer that allows quick and reliable identification of key language-sensitive regions in each individual brain. This approach enables pooling data from corresponding functional regions across subjects rather than from the same locations in stereotaxic space that may differ functionally due to inter-subject anatomical variability. In the current paper we demonstrate that the individual-subjects functional localization approach is superior to the traditional methods in its ability to distinguish among conditions in a brain region’s response. This ability is at the core of all neuroimaging research and is critical for answering questions of functional specialization (e.g., does a brain region specialize for processing syntactic aspects of the linguistic signal), which is in turn essential for making inferences about the precise computations conducted in each brain region. Based on our results, we argue that supplementing existing methods with an individual-subjects functional localization approach may lead to a clearer picture of the neural basis of syntactic processing, as it has in some other domains, such as high-level vision (e.g., Kanwisher, 2010) and social cognition (e.g., Saxe & Kanwisher, 2003).     

Recalling taboo and nontaboo words

People remember emotional and taboo words better than neutral words. It is well known that words that are processed at a deep (i.e., semantic) level are recalled better than words processed at a shallow (i.e., purely visual) level. To determine how depth of processing influences recall of emotional and taboo words, a levels of processing paradigm was used. Whether this effect holds for emotional and taboo words has not been previously investigated. Two experiments demonstrated that taboo and emotional words benefit less from deep processing than do neutral words. This is consistent with the proposal that memories for taboo and emotional words are a function of the arousal level they evoke, even under shallow encoding conditions. Recall was higher for taboo words, even when taboo words were cued to be recalled after neutral and emotional words. The superiority of taboo word recall is consistent with cognitive neuroscience and brain imaging research.     

Beyond arousal and valence: The importance of the biological versus social relevance of emotional stimuli

The present study addressed the hypothesis that emotional stimuli relevant to survival or reproduction (biologically emotional stimuli) automatically affect cognitive processing (e.g., attention, memory), while those relevant to social life (socially emotional stimuli) require elaborative processing to modulate attention and memory. Results of our behavioral studies showed that (1) biologically emotional images hold attention more strongly than do socially emotional images, (2) memory for biologically emotional images was enhanced even with limited cognitive resources, but (3) memory for socially emotional images was enhanced only when people had sufficient cognitive resources at encoding. Neither images’ subjective arousal nor their valence modulated these patterns. A subsequent functional magnetic resonance imaging study revealed that biologically emotional images induced stronger activity in the visual cortex and greater functional connectivity between the amygdala and visual cortex than did socially emotional images. These results suggest that the interconnection between the amygdala and visual cortex supports enhanced attention allocation to biological stimuli. In contrast, socially emotional images evoked greater activity in the medial prefrontal cortex (MPFC) and yielded stronger functional connectivity between the amygdala and MPFC than did biological images. Thus, it appears that emotional processing of social stimuli involves elaborative processing requiring frontal lobe activity.     

Neurophysiology and neuroanatomy of reflexive and volitional saccades: evidence from studies of humans

This review provides a summary of the contributions made by human functional neuroimaging studies to the understanding of neural correlates of saccadic control. The generation of simple visually guided saccades (redirections of gaze to a visual stimulus or pro-saccades) and more complex volitional saccades require similar basic neural circuitry with additional neural regions supporting requisite higher level processes. The saccadic system has been studied extensively in non-human (e.g., single-unit recordings) and human (e.g., lesions and neuroimaging) primates. Considerable knowledge of this system’s functional neuroanatomy makes it useful for investigating models of cognitive control. The network involved in pro-saccade generation (by definition largely exogenously-driven) includes subcortical (striatum, thalamus, superior colliculus, and cerebellar vermis) and cortical (primary visual, extrastriate, and parietal cortices, and frontal and supplementary eye fields) structures. Activation in these regions is also observed during endogenously-driven voluntary saccades (e.g., anti-saccades, ocular motor delayed response or memory saccades, predictive tracking tasks and anticipatory saccades, and saccade sequencing), all of which require complex cognitive processes like inhibition and working memory. These additional requirements are supported by changes in neural activity in basic saccade circuitry and by recruitment of additional neural regions (such as prefrontal and anterior cingulate cortices). Activity in visual cortex is modulated as a function of task demands and may predict the type of saccade to be generated, perhaps via top-down control mechanisms. Neuroimaging studies suggest two foci of activation within FEF - medial and lateral - which may correspond to volitional and reflexive demands, respectively. Future research on saccade control could usefully (i) delineate important anatomical subdivisions that underlie functional differences, (ii) evaluate functional connectivity of anatomical regions supporting saccade generation using methods such as ICA and structural equation modeling, (iii) investigate how context affects behavior and brain activity, and (iv) use multi-modal neuroimaging to maximize spatial and temporal resolution.     

Imageability estimates for 3,000 disyllabic words

We provide imageability estimates for 3,000 disyllabic words (as supplementary materials that may be downloaded with the article from www.springerlink.com ). Imageability is a widely studied lexical variable believed to influence semantic and memory processes (see, e.g., Paivio, 1971). In addition, imageability influences basic word recognition processes (Plaut, McClelland, Seidenberg, & Patterson, 1996). In fact, neuroimaging studies have suggested that reading high- and low-imageable words elicits distinct neural activation patterns for the two types e.g., Bedny & Thompson-Schill (Brain and Language 98:127-139, 2006; Graves, Binder, Desai, Conant, & Seidenberg NeuroImage 53:638-646, 2010). Despite the usefulness of this variable, imageability estimates have not been available for large sets of words. Furthermore, recent megastudies of word processing e.g., Balota et al. (Behavior Research Methods 39:445-459, 2007) have expanded the number of words that interested researchers can select according to other lexical characteristics (e.g., average naming latencies, lexical decision times, etc.). However, the dearth of imageability estimates (as well as those of other lexical characteristics) limits the items that researchers can include in their experiments. Thus, these imageability estimates for disyllabic words expand the number of words available for investigations of word processing, which should be useful for researchers interested in the influences of imageability both as an input and as an outcome variable.     

Working memory load and distraction: dissociable effects of visual maintenance and cognitive control

We establish a new dissociation between the roles of working memory (WM) cognitive control and visual maintenance in selective attention as measured by the efficiency of distractor rejection. The extent to which focused selective attention can prevent distraction has been shown to critically depend on the level and type of load involved in the task. High perceptual load that consumes perceptual capacity leads to reduced distractor processing, whereas high WM load that reduces WM ability to exert priority-based executive cognitive control over the task results in increased distractor processing (e.g., Lavie, Trends in Cognitive Sciences, 9(2), 75–82, 2005). WM also serves to maintain task-relevant visual representations, and such visual maintenance is known to recruit the same sensory cortices as those involved in perception (e.g., Pasternak & Greenlee, Nature Reviews Neuroscience, 6(2), 97–107, 2005). These findings led us to hypothesize that loading WM with visual maintenance would reduce visual capacity involved in perception, thus resulting in reduced distractor processing—similar to perceptual load and opposite to WM cognitive control load. Distractor processing was assessed in a response competition task, presented during the memory interval (or during encoding; Experiment 1a) of a WM task. Loading visual maintenance or encoding by increased set size for a memory sample of shapes, colors, and locations led to reduced distractor response competition effects. In contrast, loading WM cognitive control with verbal rehearsal of a random letter set led to increased distractor effects. These findings confirm load theory predictions and provide a novel functional distinction between the roles of WM maintenance and cognitive control in selective attention.     

从面孔模块到马赛克──视觉特异性加工的脑机制

“脑功能的模块性”是视觉乃至整个认知神经科学的研究热点。特异性的面孔加工模块被当作是认知模块假说的最好证据。但综述跨学科的正反两方面实验证据表明,视觉加工存在特异性,并没有普遍的模块式的亚单元,而是“马赛克”镶嵌式的组构。     

Genes, brain, and behavior: Bridging disciplines

With excitement surrounding the publication of the human genome, scientists have set out to uncover the functions of specific genes. This special issue on Genes, Brain, and Behavior attempts to present research strategies that connect major avenues of genetic research across disciplines. For example, anatomical information provided by brain imaging can serve as a convenient link between anatomical abnormalities seen in knockout/transgenic mouse models and abnormal patterns of brain activity seen in certain patient populations. Identifying genetic risk factors for disorders with carefully designed cognitive assays is another strategy that has gained increasing attention. These approaches are being combined with behavioral studies of mouse models of gene function. Alone, each of these approaches provides limited information on gene function in complex human behavior, but together, they are forming bridges between animal models and human psychiatric disorders.     

Neuroimaging evidence for agenda-dependent monitoring of different features during short-term source memory tests

A short-term source monitoring procedure with functional magnetic resonance imaging assessed neural activity when participants made judgments about the format of 1 of 4 studied items (picture, word), the encoding task performed (cost, place), or whether an item was old or new. The results support findings from long-term memory studies showing that left anterior ventrolateral prefrontal cortex (PFC) is engaged when people make source attributions about reflectively generated information (cognitive operations, conceptual features). The findings also point to a role for right lateral PFC in attention to perceptual features and/or familiarity in making source decisions. Activity in posterior regions also differed depending on what was evaluated. These results provide neuroimaging evidence for theoretical approaches emphasizing that agendas influence which features are monitored during remembering (e.g., M. K. Johnson, S. Hashtroudi, & D. S. Lindsay, 1993). They also support the hypothesis that some of the activity in left lateral PFC and posterior regions associated with remembering specific information is not unique to long-term memory but rather is associated with agenda-driven source monitoring processes common to working memory and long-term memory.     

Neurophysiology and neuroanatomy of reflexive and volitional saccades: Evidence from studies of humans

This review provides a summary of the contributions made by human functional neuroimaging studies to the understanding of neural correlates of saccadic control. The generation of simple visually guided saccades (redirections of gaze to a visual stimulus or pro-saccades) and more complex volitional saccades require similar basic neural circuitry with additional neural regions supporting requisite higher level processes. The saccadic system has been studied extensively in non-human (e.g., single-unit recordings) and human (e.g., lesions and neuroimaging) primates. Considerable knowledge of this system’s functional neuroanatomy makes it useful for investigating models of cognitive control. The network involved in pro-saccade generation (by definition largely exogenously-driven) includes subcortical (striatum, thalamus, superior colliculus, and cerebellar vermis) and cortical (primary visual, extrastriate, and parietal cortices, and frontal and supplementary eye fields) structures. Activation in these regions is also observed during endogenously-driven voluntary saccades (e.g., anti-saccades, ocular motor delayed response or memory saccades, predictive tracking tasks and anticipatory saccades, and saccade sequencing), all of which require complex cognitive processes like inhibition and working memory. These additional requirements are supported by changes in neural activity in basic saccade circuitry and by recruitment of additional neural regions (such as prefrontal and anterior cingulate cortices). Activity in visual cortex is modulated as a function of task demands and may predict the type of saccade to be generated, perhaps via top-down control mechanisms. Neuroimaging studies suggest two foci of activation within FEF - medial and lateral - which may correspond to volitional and reflexive demands, respectively. Future research on saccade control could usefully (i) delineate important anatomical subdivisions that underlie functional differences, (ii) evaluate functional connectivity of anatomical regions supporting saccade generation using methods such as ICA and structural equation modeling, (iii) investigate how context affects behavior and brain activity, and (iv) use multi-modal neuroimaging to maximize spatial and temporal resolution.     

Syntactic gender processing in the human brain: a review and a model

Despite the increasing number of neuroimaging studies of syntactic gender processing no model is currently available that includes data from visual and auditory language comprehension and language production. This paper provides a systematic review of the neural correlates of syntactic gender processing. Based on anatomical information from cytoarchitectonic probability maps it is argued that the left BA 44 plays a central role for the active use of gender information, e.g., for explicit decisions as well as for subsequent morphological encoding. The left BA 45 is involved in the strategic generation of morphological cues that facilitate gender processing. Model implications for aphasic patients with lesions including or excluding parts of Broca's speech region are discussed.