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How does our brain understand the number five when it is written as an Arabic numeral, and when presented as five fingers held up? Four facets have been implicated in adult numerical processing: semantic, visual, manual, and phonological/verbal. Here, we ask how the brain represents each, using a combination of tasks and stimuli. We collected fMRI data from adult participants while they completed our novel "four number code" paradigm. In this paradigm, participants viewed one of two stimulus types to tap into the visual and manual number codes, respectively. Concurrently, they completed one of two tasks to tap into the semantic and phonological/verbal number codes, respectively. Classification analyses revealed that neural codes representing distinctions between the number comparison and phonological tasks were generalizable across format (e.g., Arabic numerals to hands) within intraparietal sulcus (IPS), angular gyrus, and precentral gyrus. Neural codes representing distinctions between formats were generalizable across tasks within visual areas such as fusiform gyrus and calcarine sulcus, as well as within IPS. Our results identify the neural facets of numerical processing within a single paradigm and suggest that IPS is sensitive to distinctions between semantic and phonological/verbal, as well as visual and manual, facets of number representations., (© 2022 Massachusetts Institute of Technology.)

Writing systems vary in many ways, making it difficult to account for cross-linguistic neural differences. For example, orthographic processing of Chinese characters activates the mid-fusiform gyri (mFG) bilaterally, whereas the processing of English words predominantly activates the left mFG. Because Chinese and English vary in visual processing (holistic vs. analytical) and linguistic mapping principle (morphosyllabic vs. alphabetic), either factor could account for mFG laterality differences. We used artificial orthographies representing English to investigate the effect of mapping principle on mFG lateralization. The fMRI data were compared for two groups that acquired foundational proficiency: one for an alphabetic and one for an alphasyllabic artificial orthography. Greater bilateral mFG activation was observed in the alphasyllabic versus alphabetic group. The degree of bilaterality correlated with reading fluency for the learned orthography in the alphasyllabic but not alphabetic group. The results suggest that writing systems with a syllable-based mapping principle recruit bilateral mFG to support orthographic processing. Implications for individuals with left mFG dysfunction will be discussed.

Numerous functional neuroimaging studies have shown that most orthographic stimuli, such as printed English words, produce a left-lateralized response within the fusiform gyrus (FG) at a characteristic location termed the visual word form area (VWFA). We developed an experimental alphabet (FaceFont) comprising 35 face-phoneme pairs to disentangle phonological and perceptual influences on the lateralization of orthographic processing within the FG. Using functional imaging, we found that a region in the vicinity of the VWFA responded to FaceFont words more strongly in trained versus untrained participants, whereas no differences were observed in the right FG. The trained response magnitudes in the left FG region correlated with behavioral reading performance, providing strong evidence that the neural tissue recruited by training supported the newly acquired reading skill. These results indicate that the left lateralization of the orthographic processing is not restricted to stimuli with particular visual-perceptual features. Instead, lateralization may occur because the anatomical projections in the vicinity of the VWFA provide a unique interconnection between the visual system and left-lateralized language areas involved in the representation of speech.

Adults have difficulty discriminating nonnative phonetic contrasts, but under certain circumstances training can lead to improvement in this ability. Despite the ubiquitous use of performance feedback in training paradigms in this and many other domains, the mechanisms by which feedback affects learning are not well understood. In this event-related functional magnetic resonance imaging study, we examined how performance feedback is processed during perceptual learning. Thirteen Japanese speakers for whom the English phonemes [r] and [l] were nondistinct performed an identification task of the words "road" and "load" that has been shown to be effective in inducing learning only when performance feedback is present. Each subject performed alternating runs of training with and without feedback, followed by performance of a card-guessing task with monetary reward and punishment outcomes. We found that the caudate nucleus was more robustly activated bilaterally when performing the perceptual identification task with feedback than without feedback, and the right caudate nucleus also showed a differential response to positive and negative feedback. Moreover, using a within-subjects design, we found that the caudate nucleus also showed a similar activation pattern to monetary reward and punishment outcomes in the card-guessing task. These results demonstrate that the caudate responds to positive and negative feedback during learning in a manner analogous to its processing of extrinsic affective reinforcers and indicate that this region may be a critical moderator of the influence of feedback on learning. These findings impact our broader understanding of the mechanisms underlying nondeclarative learning and language acquisition.

To distinguish areas involved in the processing of word meaning (semantics) from other regions involved in lexical processing more generally, subjects were scanned with positron emission tomography (PET) while performing lexical tasks, three of which required varying degrees of semantic analysis and one that required phonological analysis. Three closely apposed regions in the left inferior frontal cortex and one in the right cerebellum were significantly active above baseline in the semantic tasks, but not in the nonsemantic task. The activity in two of the frontal regions was modulated by the difficulty of the semantic judgment. Other regions, including some in the left temporal cortex and the cerebellum, were active across all four language tasks. Thus, in addition to a number of regions known to be active during language processing, regions in the left inferior frontal cortex were specifically recruited during semantic processing in a task-dependent manner. A region in the right cerebellum may be functionally related to those in the left inferior frontal cortex. Discussion focuses on the implications of these results for current views regarding neural substrates of semantic processing.

Nine previous positron emission tomography (PET) studies of human visual information processing were reanalyzed to determine the consistency across experiments of blood flow decreases during active tasks relative to passive viewing of the same stimulus array. Areas showing consistent decreases during active tasks included posterior cingulate/precuneous (Brodmann area, BA 31/7), left (BAS 40 and 39/19) and right (BA 40) inferior parietal cortex, left dorsolateral frontal cortex (BA S), left lateral inferior frontal cortex (BA 10/47), left inferior temporal gyrus @A 20), a strip of medial frontal regions running along a dorsal-ventral axis (BAs 8, 9, 10, and 32), and the right amygdala. Experiments involving language-related processes tended to show larger decreases than nonlanguage experiments. This trend mainly reflected blood flow increases at certain areas in the passive conditions of the language experiments (relative to a fixation control in which no task stimulus was present) and slight blood flow decreases in the passive conditions of the nonlanguage experiments. When the active tasks were referenced to the fixation condition, the overall size of blood flow decreases in language and nonlanguage tasks were the same, but differences were found across cortical areas. Decreases were more pronounced in the posterior cingulate/precuneous (BAS 31/7) and right inferior parietal cortex (BA 40) during language-related tasks and more pronounced in left inferior frontal cortex (BA 10/47) during nonlanguage tasks. Blood flow decreases did not generally show significant differences across the active task states within an experiment, but a verb-generation task produced larger decreases than a read task in right and left inferior parietal lobe (BA 40) and the posterior cingulate/precuneous (BA 31/7), while the read task produced larger decreases in left lateral inferior frontal cortex (BA 10/47). These effects mirrored those found between experiments in the language-nonlanguage comparison. Consistent active minus passive decreases may reflect decreased activity caused by active task processes that generalize over tasks or increased activity caused by passive task processes that are suspended during the active tasks. Increased activity during the passive condition might reflect ongoing processes, such as unconstrained verbally mediated thoughts and monitoring of the external environment, body, and emotional state.

Nine positron emission tomography (PET) studies of human visual information processing were reanalyzed to determine the consistency across experiments of blood flow increases during active tasks relative to passive viewing of the same stimulus array. No consistent blood flow increases were found in cerebral cortex outside of the visual system, but increases were seen in the thalamus and cerebellum. Although most tasks involve increases in arousal, establishing an intention or behavioral goal, setting up control structures for sequencing task operations, detecting targets, etc., these operations do not produce blood flow increases, detectable with the present methods, in localized cortical regions that are common across tasks. Common subcortical regions, however, may be involved. A left cerebellar and a medial cerebellar focus reflected motor-related processes. Blood flow increases in these regions only occurred in experiments in which the subject made an overt response and were largest when the response was made in the active but not passive condition. These motor-related processes were more complex than simple motor execution, however, since increases were still present when the response was made in both the active and passive conditions. These cerebellar increases may reflect processes related to response selection.Blood flow increases in a right cerebellar region were not motor-related. Increases were not modulated by the presence or absence of motor responses during either the active or passive conditions, and increases were sensitive to within-experiment variables that held the motor response constant. Increases occurred in both language and nonlanguage tasks and appeared to involve a general nonmotor process, but the nature of that process was difficult to specify. A right thalamic focus was sensitive to variables related to focal attention, suggesting that this region was involved in attentional engagement. Right thalamic increases were also correlated over conditions with increases in the left and medial cerebellum, perhaps reflecting additional contributions from motor-related nuclei receiving cerebellar projections. Blood flow increases in a left thalamic focus were completely uncorrelated over conditions with increases in the right thalamus, indicating that it was involved in different functions. Both the left thalamus and right cerebellum yielded larger blood flow increases when subjects performed a complex rather than simple language task, possibly reflecting a language-related pathway. Blood flow increases in the left thalamus were also observed, however, during nonlanguage tasks.

Positron emission tomography (PET) was used to investigate the functional anatomy of auditory and phonological processing. Stimulus sets were designed to determine areas of the brain significantly activated during speech and nonspeech acoustic processing for stimuli with or without rapidly changing acoustic cues. Performance of auditory target detection tasks using these stimulus sets produced increased activation in superior temporal, frontal opercular, and medial frontal (SMA) cortices, relative to a visual fixation control task. While the medial frontal and superior temporal changes are best explained by motor and sensory components of the task, respectively, the frontal opercular changes were dependent upon the task performed upon the auditory input (mere presentation of the stimuli did not result in significant activation). On the left, the frontal opercular increases were larger when subjects performed an auditory detection task upon stimuli that incorporated rapid temporal changes (words, syllables, and tone sequences) than steady-state vowels. A converging study involving performance of orthographic (ascending letter) and phonological (long vowel sound) word discrimination tasks supports anatomical and behavioral evidence suggesting the left frontal opercular region is important for certain types of auditory/temporal analysis, as well as high-level articulatory coding. In addition to the activation increases associated with performance of auditory target detection tasks, decreases in activation were observed bilaterally along the intraparietal sulcus and superior parietal cortex, in the Rolandic sulcus, and the posterior cingulate; these decreases may reflect an attentional shift away from areas involved in the fixation task during the performance of a difficult auditory task. These results demonstrate that focusing more closely on basic neural processing differences (such as temporal integration rates) may lead to a better understanding of the specific neural processes that underlie complex phonological tasks.