Multi-representation of symbolic and nonsymbolic numerical magnitude in Chinese number processing

Numerical information can be conveyed by either symbolic or nonsymbolic representation. Some symbolic numerals can also be identified as nonsymbolic quantities defined by the number of lines (e.g., I, II, III in Roman and -, =, ≡ in Japanese Kanji and Chinese). Here we report that such multi-representation of magnitude can facilitate the processing of these numerals under certain circumstances. In a magnitude comparison task judging 1 to 9 (except 5) Chinese and Arabic numerals presented at the foveal (at the center) or parafoveal (3° left or right of the center) location, multi-representational small-value Chinese numerals showed a processing advantage over single-representational Arabic numerals and large-value Chinese numerals only in the parafoveal condition, demonstrated by lower error rates and faster reaction times. Further event-related potential (ERP) analysis showed that such a processing advantage was not reflected by traditional ERP components identified in previous studies of number processing, such as N1 or P2p. Instead, the difference was found much later in a N400 component between 300-550 msec over parietal regions, suggesting that those behavioral differences may not be due to early processing of visual identification, but later processing of subitizing or accessing mental number line when lacking attentional resources. These results suggest that there could be three stages of number processing represented separately by the N1, P2p and N400 ERP components. In addition, numerical information can be represented simultaneously by both symbolic and nonsymbolic systems, which will facilitate number processing in certain situations.     

A magnitude code common to numerosities and number symbols in human intraparietal cortex

Activation of the horizontal segment of the intraparietal sulcus (hIPS) has been observed in various number-processing tasks, whether numbers were conveyed by symbolic numerals (digits, number words) or by nonsymbolic displays (dot patterns). This suggests an abstract coding of numerical magnitude. Here, we critically tested this hypothesis using fMRI adaptation to demonstrate notation-independent coding of numerical quantity in the hIPS. Once subjects were adapted either to dot patterns or to Arabic digits, activation in the hIPS and in frontal regions recovered in a distance-dependent fashion whenever a new number was presented, irrespective of notation changes. This remained unchanged when analyzing the hIPS peaks from an independent localizer scan of mental calculation. These results suggest an abstract coding of approximate number common to dots, digits, and number words. They support the idea that symbols acquire meaning by linking neural populations coding symbol shapes to those holding nonsymbolic representations of quantities.     

Functional imaging of numerical processing in adults and 4-y-old children

Adult humans, infants, pre-school children, and non-human animals appear to share a system of approximate numerical processing for non-symbolic stimuli such as arrays of dots or sequences of tones. Behavioral studies of adult humans implicate a link between these non-symbolic numerical abilities and symbolic numerical processing (e.g., similar distance effects in accuracy and reaction-time for arrays of dots and Arabic numerals). However, neuroimaging studies have remained inconclusive on the neural basis of this link. The intraparietal sulcus (IPS) is known to respond selectively to symbolic numerical stimuli such as Arabic numerals. Recent studies, however, have arrived at conflicting conclusions regarding the role of the IPS in processing non-symbolic, numerosity arrays in adulthood, and very little is known about the brain basis of numerical processing early in development. Addressing the question of whether there is an early-developing neural basis for abstract numerical processing is essential for understanding the cognitive origins of our uniquely human capacity for math and science. Using functional magnetic resonance imaging (fMRI) at 4-Tesla and an event-related fMRI adaptation paradigm, we found that adults showed a greater IPS response to visual arrays that deviated from standard stimuli in their number of elements, than to stimuli that deviated in local element shape. These results support previous claims that there is a neurophysiological link between non-symbolic and symbolic numerical processing in adulthood. In parallel, we tested 4-y-old children with the same fMRI adaptation paradigm as adults to determine whether the neural locus of non-symbolic numerical activity in adults shows continuity in function over development. We found that the IPS responded to numerical deviants similarly in 4-y-old children and adults. To our knowledge, this is the first evidence that the neural locus of adult numerical cognition takes form early in development, prior to sophisticated symbolic numerical experience. More broadly, this is also, to our knowledge, the first cognitive fMRI study to test healthy children as young as 4 y, providing new insights into the neurophysiology of human cognitive development.     

A Two-Minute Paper-and-Pencil Test of Symbolic and Nonsymbolic Numerical Magnitude Processing Explains Variability in Primary School Children's Arithmetic Competence

Recently, there has been a growing emphasis on basic number processing competencies (such as the ability to judge which of two numbers is larger) and their role in predicting individual differences in school-relevant math achievement. Children’s ability to compare both symbolic (e.g. Arabic numerals) and nonsymbolic (e.g. dot arrays) magnitudes has been found to correlate with their math achievement. The available evidence, however, has focused on computerized paradigms, which may not always be suitable for universal, quick application in the classroom. Furthermore, it is currently unclear whether both symbolic and nonsymbolic magnitude comparison are related to children’s performance on tests of arithmetic competence and whether either of these factors relate to arithmetic achievement over and above other factors such as working memory and reading ability. In order to address these outstanding issues, we designed a quick (2 minute) paper-and-pencil tool to assess children’s ability to compare symbolic and nonsymbolic numerical magnitudes and assessed the degree to which performance on this measure explains individual differences in achievement. Children were required to cross out the larger of two, single-digit numerical magnitudes under time constraints. Results from a group of 160 children from grades 1–3 revealed that both symbolic and nonsymbolic number comparison accuracy were related to individual differences in arithmetic achievement. However, only symbolic number comparison performance accounted for unique variance in arithmetic achievement. The theoretical and practical implications of these findings are discussed which include the use of this measure as a possible tool for identifying students at risk for future difficulties in mathematics.     

Maps of space in human frontoparietal cortex

Prefrontal cortex (PFC) and posterior parietal cortex (PPC) are neural substrates for spatial cognition. We here review studies in which we tested the hypothesis that human frontoparietal cortex may function as a priority map. According to priority map theory, objects or locations in the visual world are represented by neural activity that is proportional to their attentional priority. Using functional magnetic resonance imaging (fMRI), we first identified topographic maps in PFC and PPC as candidate priority maps of space. We then measured fMRI activity in candidate priority maps during the delay periods of a covert attention task, a spatial working memory task, and a motor planning task to test whether the activity depended on the particular spatial cognition. Our hypothesis was that some, but not all, candidate priority maps in PFC and PPC would be agnostic with regard to what was being prioritized, in that their activity would reflect the location in space across tasks rather than a particular kind of spatial cognition (e.g., covert attention). To test whether patterns of delay period activity were interchangeable during the spatial cognitive tasks, we used multivariate classifiers. We found that decoders trained to predict the locations on one task (e.g., working memory) cross-predicted the locations on the other tasks (e.g., covert attention and motor planning) in superior precentral sulcus (sPCS) and in a region of intraparietal sulcus (IPS2), suggesting that these patterns of maintenance activity may be interchangeable across the tasks. Such properties make sPCS in frontal cortex and IPS2 in parietal cortex viable priority map candidates, and suggest that these areas may be the human homologs of the monkey frontal eye field (FEF) and lateral intraparietal area (LIP).     

The number domain- can we count on parietal cortex?

Does the primate brain contain a dedicated and localized neural circuitry for processing generic numerical information? The human parietal cortex, particularly the intraparietal sulcus (IPS), has long been implicated in processing symbolic (verbal) number information. If the IPS is indeed the site of generic numerical processing, however, its neurons should also encode nonsymbolic numerosity information. Two recent studies by Shuman and Kanwisher and by Piazza et al. published in this issue of Neuron tested this assumption...with quite different results.     

The Processing of Symbolic and Nonsymbolic Ratios in School-Age Children

This study tested the processing of ratios of natural numbers in school-age children. Nine- and eleven-year-olds were presented collections made up of orange and grey dots (i.e., nonsymbolic format) and fractions (i.e., symbolic format). They were asked to estimate ratios between the number of orange dots and the total number of dots and fractions by producing an equivalent ratio of surface areas (filling up a virtual glass). First, we tested whether symbolic notation of ratios affects their processing by directly comparing performance on fractions with that on dot sets. Second, we investigated whether children’s estimates of nonsymbolic ratios of natural numbers relied at least in part on ratios of surface areas by contrasting a condition in which the ratio of surface areas occupied by dots covaried with the ratio of natural numbers and a condition in which this ratio of surface areas was kept constant across ratios of natural numbers. The results showed that symbolic notation did not really have a negative impact on performance among 9-year-olds, while it led to more accurate estimates in 11-year-olds. Furthermore, in dot conditions, children’s estimates increased consistently with ratios between the number of orange dots and the total number of dots even when the ratio of surface areas was kept constant but were less accurate in that condition than when the ratio of surface areas covaried with the ratio of natural numbers. In summary, these results indicate that mental magnitude representation is more accurate when it is activated from symbolic ratios in children as young as 11 years old and that school-age children rely at least in part on ratios of surface areas to process nonsymbolic ratios of natural numbers when given the opportunity to do so.     

Domain-Specific Control of Selective Attention

Previous research has shown that loading information on working memory affects selective attention. However, whether the load effect on selective attention is domain-general or domain-specific remains unresolved. The domain-general effect refers to the findings that load in one content (e.g. phonological) domain in working memory influences processing in another content (e.g., visuospatial) domain. Attentional control supervises selection regardless of information domain. The domain-specific effect refers to the constraint of influence only when maintenance and processing operate in the same domain. Selective attention operates in a specific content domain. This study is designed to resolve this controversy. Across three experiments, we manipulated the type of representation maintained in working memory and the type of representation upon which the participants must exert control to resolve conflict and select a target into the focus of attention. In Experiments 1a and 1b, participants maintained digits and nonverbalized objects, respectively, in working memory while selecting a target in a letter array. In Experiment 2, we presented auditory digits with a letter flanker task to exclude the involvement of resource competition within the same input modality. In Experiments 3a and 3b, we replaced the letter flanker task with an object flanker task while manipulating the memory load on object and digit representation, respectively. The results consistently showed that memory load modulated distractibility only when the stimuli of the two tasks were represented in the same domain. The magnitude of distractor interference was larger under high load than under low load, reflecting a lower efficacy of information prioritization. When the stimuli of the two tasks were represented in different domains, memory load did not modulate distractibility. Control of processing priority in selective attention demands domain-specific resources.     

How distributed is visual category information in human occipito-temporal cortex? An fMRI study

We used fMRI to study the distribution of object category information in the ventral visual pathway. Extending the findings of, we find that categories of stimuli can be distinguished by the pattern of activation they elicit across this entire pathway, even when the stimuli within a category differ in viewpoint, exemplar, or image format. However, regions within the ventral visual pathway are neither interchangeable nor equipotential. Although the FFA and PPA permit excellent discrimination between preferred versus nonpreferred stimuli (e.g., faces-bottles and houses-bottles, respectively), we find that neither region alone permits accurate discrimination between pairs of nonpreferred stimuli (e.g., bottles-shoes). These findings indicate that the ventral visual pathway is not homogeneous, but contains some regions (including FFA and PPA) that are primarily involved in the analysis of a single class of stimulus.     

The Mediating Relation between Symbolic and Nonsymbolic Foundations of Math Competence

This study investigated the relation between symbolic and nonsymbolic magnitude processing abilities with 2 standardized measures of math competence (WRAT Arithmetic and KeyMath Numeration) in 150 3^rd- grade children (mean age 9.01 years). Participants compared sets of dots and pairs of Arabic digits with numerosities 1–9 for relative numerical magnitude. In line with previous studies, performance on both symbolic and nonsymbolic magnitude processing was related to math ability. Performance metrics combining reaction and accuracy, as well as weber fractions, were entered into mediation models with standardized math test scores. Results showed that symbolic magnitude processing ability fully mediates the relation between nonsymbolic magnitude processing and math ability, regardless of the performance metric or standardized test.     

Perceptual blending in odor mixtures depends on the nature of odorants and human olfactory expertise

Our olfactory system is confronted with complex mixtures of odorants, often recognized as single entities due to odor blending (e.g., coffee). In contrast, we are also able to discriminate odors from complex mixtures (e.g., off-odors). Therefore, the olfactory system is able to engage either configural or elemental processes when confronted with mixtures. However, the rules that govern the involvement of these processes during odor perception remain poorly understood. In our first experiment, we examined whether simple odorant mixtures (binary/ternary) could elicit configural perception. Twenty untrained subjects were asked to evaluate the odor typicality of mixtures and their constituents. The results revealed a significant increase in odor typicality in some but not all mixtures as compared with the single components, which suggest that perceptual odor blending can occur only in specific mixtures (configural processing). In our second experiment, we tested the hypothesis that general olfactory expertise can improve elemental perception of mixtures. Thirty-two trained subjects evaluated the odor typicality of the stimuli presented during the first experiment, and their responses were compared with those obtained from the untrained panelists. The results support the idea that general training with odors increases the elemental perception of binary and ternary blending mixtures.     

汉字字形认知研究的刺激源设计及在fMRI研究中的应用

目的:评估汉字字形刺激源在汉字认知fMRI研究中的有效性,并对参与汉字处理的脑皮层区域进行定位及初步的量化分析。方法:选择母语为汉语、经利手测试后为右利手且裸眼视力正常(大于等于1．0)的在校大学生10例(男6例,女4例)作为被试。试验任务采用组块设计,将汉字(非字、假字、真字)投射到屏幕上,受试者接受汉字字形图片的视觉刺激,按非字-假字-真字-非字-假字-真字顺序呈现,共6个block。数据处理及统计分析采用国际通用的AFNI软件。结果:左额叶上、中、下回(包括Broca's area)、左中央前回(BA6)、左顶上小叶及顶下小叶(包括缘上回及角回)及双侧枕叶、楔前叶显著激活;左颞叶梭状回(BA37)、右额下回及双侧颞中、上回及扣带回显著激活,左大脑半球的激活体积明显大于右侧大脑半球。结论:本研究设计的汉字字形刺激源结合功能磁共振成像技术可以对汉字处理的相关大脑皮层区域进行定位,为研究人脑加工处理汉字的神经机制提供了一种有效的无创性影像学方法,并应用fMRI技术进一步证实其优势半球为左半球,且需要多种脑区共同参与完成。本试验模式可望成为一种对语言障碍病人进行脑功能检查的有效手段,从而为指导临床治疗和评价预后提供更丰富的信息。     

The Perception of Dynamic and Static Facial Expressions of Happiness and Disgust Investigated by ERPs and fMRI Constrained Source Analysis

A recent functional magnetic resonance imaging (fMRI) study by our group demonstrated that dynamic emotional faces are more accurately recognized and evoked more widespread patterns of hemodynamic brain responses than static emotional faces. Based on this experimental design, the present study aimed at investigating the spatio-temporal processing of static and dynamic emotional facial expressions in 19 healthy women by means of multi-channel electroencephalography (EEG), event-related potentials (ERP) and fMRI-constrained regional source analyses. ERP analysis showed an increased amplitude of the LPP (late posterior positivity) over centro-parietal regions for static facial expressions of disgust compared to neutral faces. In addition, the LPP was more widespread and temporally prolonged for dynamic compared to static faces of disgust and happiness. fMRI constrained source analysis on static emotional face stimuli indicated the spatio-temporal modulation of predominantly posterior regional brain activation related to the visual processing stream for both emotional valences when compared to the neutral condition in the fusiform gyrus. The spatio-temporal processing of dynamic stimuli yielded enhanced source activity for emotional compared to neutral conditions in temporal (e.g., fusiform gyrus), and frontal regions (e.g., ventromedial prefrontal cortex, medial and inferior frontal cortex) in early and again in later time windows. The present data support the view that dynamic facial displays trigger more information reflected in complex neural networks, in particular because of their changing features potentially triggering sustained activation related to a continuing evaluation of those faces. A combined fMRI and EEG approach thus provides an advanced insight to the spatio-temporal characteristics of emotional face processing, by also revealing additional neural generators, not identifiable by the only use of an fMRI approach.     

A deficit in covert attention after parietal cortex inactivation in the monkey

Although the parietal cortex has been repeatedly implicated in controlling attention, the nature and importance of this contribution remain unclear. Here we show that inactivating the lateral intraparietal area in monkeys delays the detection of a visual target located in the contralateral visual field. This effect was observed using different visual scene configurations, e.g., with distractors that differ in number or that differ from the target by a conjunction of shape and color or by a single feature. Since eye movements were not allowed during the searching tasks, these results argue for an unambiguous role of the parietal cortex in the top-down control of attentional deployment in space.     

Social Distance Evaluation in Human Parietal Cortex

Across cultures, social relationships are often thought of, described, and acted out in terms of physical space (e.g. “close friends” “high lord”). Does this cognitive mapping of social concepts arise from shared brain resources for processing social and physical relationships? Using fMRI, we found that the tasks of evaluating social compatibility and of evaluating physical distances engage a common brain substrate in the parietal cortex. The present study shows the possibility of an analytic brain mechanism to process and represent complex networks of social relationships. Given parietal cortex''s known role in constructing egocentric maps of physical space, our present findings may help to explain the linguistic, psychological and behavioural links between social and physical space.     

Synaesthetic Colour in the Brain: Beyond Colour Areas. A Functional Magnetic Resonance Imaging Study of Synaesthetes and Matched Controls

Background

In synaesthesia, sensations in a particular modality cause additional experiences in a second, unstimulated modality (e.g., letters elicit colour). Understanding how synaesthesia is mediated in the brain can help to understand normal processes of perceptual awareness and multisensory integration. In several neuroimaging studies, enhanced brain activity for grapheme-colour synaesthesia has been found in ventral-occipital areas that are also involved in real colour processing. Our question was whether the neural correlates of synaesthetically induced colour and real colour experience are truly shared.

Methodology/Principal Findings

First, in a free viewing functional magnetic resonance imaging (fMRI) experiment, we located main effects of synaesthesia in left superior parietal lobule and in colour related areas. In the left superior parietal lobe, individual differences between synaesthetes (projector-associator distinction) also influenced brain activity, confirming the importance of the left superior parietal lobe for synaesthesia. Next, we applied a repetition suppression paradigm in fMRI, in which a decrease in the BOLD (blood-oxygenated-level-dependent) response is generally observed for repeated stimuli. We hypothesized that synaesthetically induced colours would lead to a reduction in BOLD response for subsequently presented real colours, if the neural correlates were overlapping. We did find BOLD suppression effects induced by synaesthesia, but not within the colour areas.

Conclusions/Significance

Because synaesthetically induced colours were not able to suppress BOLD effects for real colour, we conclude that the neural correlates of synaesthetic colour experience and real colour experience are not fully shared. We propose that synaesthetic colour experiences are mediated by higher-order visual pathways that lie beyond the scope of classical, ventral-occipital visual areas. Feedback from these areas, in which the left parietal cortex is likely to play an important role, may induce V4 activation and the percept of synaesthetic colour.     

Differential neural activation for updating rule versus stimulus information in working memory

Establishing what information is actively maintained in working memory (WM) and how it is represented and controlled is essential to understanding how such information guides future behavior. WM has traditionally been investigated in terms of the maintenance of stimulus-specific information, such as locations or words. More recently, investigators have emphasized the importance of rules that establish relationships between those stimuli and the pending response. The current study used a mental arithmetic task with fMRI to test whether updating of numbers (i.e., stimuli) and updating of mathematical operations (i.e., rules) in WM relies on the same neural system. Results indicate that, while a common network is activated by both types of updating, rule updating preferentially activates prefrontal cortex while number updating preferentially activates parietal cortex. The results suggest that both numbers and rules are maintained in WM but that they are different types of information that are controlled independently.     

Influence of Repressive Coping Style on Cortical Activation during Encoding of Angry Faces

Background

Coping plays an important role for emotion regulation in threatening situations. The model of coping modes designates repression and sensitization as two independent coping styles. Repression consists of strategies that shield the individual from arousal. Sensitization indicates increased analysis of the environment in order to reduce uncertainty. According to the discontinuity hypothesis, repressors are sensitive to threat in the early stages of information processing. While repressors do not exhibit memory disturbances early on, they manifest weak memory for these stimuli later. This study investigates the discontinuity hypothesis using functional magnetic resonance imaging (fMRI).

Methods

Healthy volunteers (20 repressors and 20 sensitizers) were selected from a sample of 150 students on the basis of the Mainz Coping Inventory. During the fMRI experiment, subjects evaluated and memorized emotional and neutral faces. Subjects performed two sessions of face recognition: immediately after the fMRI session and three days later.

Results

Repressors exhibited greater activation of frontal, parietal and temporal areas during encoding of angry faces compared to sensitizers. There were no differences in recognition of facial emotions between groups neither immediately after exposure nor after three days.

Conclusions

The fMRI findings suggest that repressors manifest an enhanced neural processing of directly threatening facial expression which confirms the assumption of hyper-responsivity to threatening information in repression in an early processing stage. A discrepancy was observed between high neural activation in encoding-relevant brain areas in response to angry faces in repressors and no advantage in subsequent memory for these faces compared to sensitizers.     

Stereopsis activates V3A and caudal intraparietal areas in macaques and humans

Stereopsis, the perception of depth from small differences between the images in the two eyes, provides a rich model for investigating the cortical construction of surfaces and space. Although disparity-tuned cells have been found in a large number of areas in macaque visual cortex, stereoscopic processing in these areas has never been systematically compared using the same stimuli and analysis methods. In order to examine the global architecture of stereoscopic processing in primate visual cortex, we studied fMRI activity in alert, fixating human and macaque subjects. In macaques, we found strongest activation to near/far compared to zero disparity in areas V3, V3A, and CIPS. In humans, we found strongest activation to the same stimuli in areas V3A, V7, the V4d topolog (V4d-topo), and a caudal parietal disparity region (CPDR). Thus, in both primate species a small cluster of areas at the parieto-occipital junction appears to be specialized for stereopsis.     

Distributed and overlapping cerebral representations of number, size, and luminance during comparative judgments

How are comparative judgments performed in the human brain? We scanned subjects with fMRI while they compared stimuli for size, luminance, or number. Regions involved in comparative judgments were identified using three criteria: task-related activation, presence of a distance effect, and interference of one dimension onto the other. We observed considerable overlap in the neural substrates of the three comparison tasks. Interestingly, the amount of overlap predicted the amount of cross-dimensional interference: in both behavior and fMRI, number interfered with size, and size with luminance, but number did not interfere with luminance. The results suggest that during comparative judgments, the relevant continuous quantities are represented in distributed and overlapping neural populations, with number and size engaging a common parietal spatial code, while size and luminance engage shared occipito-temporal perceptual representations.