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Biomedical subjects

Elizabeth M Brannon

Publications and source records attributed to Elizabeth M Brannon.

At least 19 recordsLinked to original sources

Electrophysiological evidence for notation independence in numerical processing.

BACKGROUND: A dominant view in numerical cognition is that numerical comparisons operate on a notation independent representation (Dehaene, 1992). Although previous human neurophysiological studies using scalp-recorded event-related potentials (ERPs) on the numerical distance effect have been interpreted as supporting this idea, differences in the electrophysiological correlates of the numerical distance effect in symbolic notations (e.g. Arabic numerals) and non-symbolic notations (e.g. a set of visually presented dots of a certain number) are not entirely consistent with this view. METHODS AND RESULTS: Two experiments were conducted to resolve these discrepancies. In Experiment 1, participants performed a symbolic and a non-symbolic numerical comparison task ("smaller or larger than 5?") with numerical values 1-4 and 6-9 while ERPs were recorded. Consistent with a previous report (Temple & Posner, 1998), in the symbolic condition the amplitude of the P2p ERP component (210-250 ms post-stimulus) was larger for values near to the standard than for values far from the standard whereas this pattern was reversed in the non-symbolic condition. However, closer analysis indicated that the reversal in polarity was likely due to the presence of a confounding stimulus effect on the early sensory ERP components for small versus larger numerical values in the non-symbolic condition. In Experiment 2 exclusively large numerosities (8-30) were used, thereby rendering sensory differences negligible, and with this control in place the numerical distance effect in the non-symbolic condition mirrored the symbolic condition of Experiment 1. CONCLUSION: Collectively, the results support the claim of an abstract semantic processing stage for numerical comparisons that is independent of input notation.

Journal Article↗

How much does number matter to a monkey (Macaca mulatta)?

Although many animal species can represent numerical values, little is known about how salient number is relative to other object properties for nonhuman animals. In one hypothesis, researchers propose that animals represent number only as a last resort, when no other properties differentiate stimuli. An alternative hypothesis is that animals automatically, spontaneously, and routinely represent the numerical attributes of their environments. The authors compared the influence of number versus that of shape, color, and surface area on rhesus monkeys' (Macaca mulatta) decisions by testing them on a matching task with more than one correct answer: a numerical match and a nonnumerical (color, surface area, or shape) match. The authors also tested whether previous laboratory experience with numerical discrimination influenced a monkey's propensity to represent number. Contrary to the last-resort hypothesis, all monkeys based their decisions on numerical value when the numerical ratio was favorable.

Animals↗

Adding up the effects of cultural experience on the brain.

How does the brain represent number and perform mathematical calculations? According to a recent and provocative study by Tang and colleagues, it depends on which language you learn. They found that the divergent linguistic and cultural experiences of native Chinese and native English speakers are associated with distinct patterns of brain activity during mathematical processing. Their results raise important questions about the cognitive and neural specificity of cultural influences on mathematical processes and the core nature of mathematical cognition.

Brain↗

A common representational system governed by Weber's law: nonverbal numerical similarity judgments in 6-year-olds and rhesus macaques.

This study compared nonverbal numerical processing in 6-year-olds with that in nonhuman animals using a numerical bisection task. In the study, 16 children were trained on a delayed match-to-sample paradigm to match exemplars of two anchor numerosities. Children were then required to indicate whether a sample intermediate to the anchor values was closer to the small anchor value or the large anchor value. For two sets of anchor values with the same ratio, the probability of choosing the larger anchor value increased systematically with sample number, and the psychometric functions superimposed when plotted on a logarithmic scale. The psychometric functions produced by the children also superimposed with the psychometric functions produced by rhesus monkeys in an analogous previous experiment. These examples of superimposition demonstrate that nonverbal number representations, even in children who have acquired the verbal counting system, are modulated by Weber's law.

Animals↗

Nonverbal representation of time and number in adults.

A wealth of human and animal research supports common neural processing of numerical and temporal information. Here we test whether adult humans spontaneously encode number and time in a paradigm similar to those previously used to test the mode-control model in animals. Subjects were trained to classify visual stimuli that varied in both number and duration as few/short or many/long. Subsequently subjects were tested with novel stimuli that varied time and held number constant (eight flashes in 0.8-3.2s) or varied number and held time constant (4-16 flashes in 1.6s). Adult humans classified novel stimuli as many/long as monotonic functions of both number and duration, consistent with simultaneous, nonverbal, analog encoding. Numerical sensitivity, however, was finer than temporal sensitivity, suggesting differential salience of time and number. These results support the notion that adults simultaneously represent the number and duration of stimuli but suggest a possible asymmetry in their representations.

Adult↗

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.

Adult↗

The representation of numerical magnitude.

The combined efforts of many fields are advancing our understanding of how number is represented. Researchers studying numerical reasoning in adult humans, developing humans and non-human animals are using a suite of behavioral and neurobiological methods to uncover similarities and differences in how each population enumerates and compares quantities to identify the neural substrates of numerical cognition. An important picture emerging from this research is that adult humans share with non-human animals a system for representing number as language-independent mental magnitudes and that this system emerges early in development.

Animals↗

Weber's Law influences numerical representations in rhesus macaques (Macaca mulatta).

We present the results of two experiments that probe the ability of rhesus macaques to match visual arrays based on number. Three monkeys were first trained on a delayed match-to-sample paradigm (DMTS) to match stimuli on the basis of number and ignore continuous dimensions such as element size, cumulative surface area, and density. Monkeys were then tested in a numerical bisection experiment that required them to indicate whether a sample numerosity was closer to a small or large anchor value. Results indicated that, for two sets of anchor values with the same ratio, the probability of choosing the larger anchor value systematically increased with the sample number and the psychometric functions superimposed. A second experiment employed a numerical DMTS task in which the choice values contained an exact numerical match to the sample and a distracter that varied in number. Both accuracy and reaction time were modulated by the ratio between the correct numerical match and the distracter, as predicted by Weber's Law.

Animals↗

The multisensory representation of number in infancy.

Human infants can discriminate visual and auditory stimuli solely on the basis of number, suggesting a developmental foundation for the nonverbal number representations of adult humans. Recent studies suggest that these language-independent number representations are multisensory in both adult humans and nonhuman animals. Surprisingly, however, previous studies have yielded mixed evidence concerning whether nonverbal numerical representations independent of sensory modality are present early in human development. In this article, we use a paradigm that avoids stimulus confounds present in previous studies of cross-modal numerical mapping in infants. We show that 7-month-old infants preferentially attend to visual displays of adult humans that numerically match the number of adult humans they hear speaking. These data provide evidence that by 7 months of age, infants connect numerical representations across different sensory modalities when presented with human faces and voices. Results support the possibility of a shared system between preverbal infants and nonverbal animals for representing number.

Acoustic Stimulation↗

The role of reference points in ordinal numerical comparisons by rhesus macaques (Macaca mulatta).

Two experiments examined ordinal numerical knowledge in rhesus macaques (Macaca mulatta). Experiment 1 replicated the finding (E. M. Brannon & H. S. Terrace, 2000) that monkeys trained to respond in descending numerical order (4-->3-->2-->1) did not generalize the descending rule to the novel values 5-9 in contrast to monkeys trained to respond in ascending order. Experiment 2 examined whether the failure to generalize a descending rule was due to the direction of the training sequence or to the specific values used in the training sequence. Results implicated 3 factors that characterize a monkey's numerical comparison process: Weber's law, knowledge of ordinal direction, and a comparison of each value in a test pair with the reference point established by the first value of the training sequence.

Animals↗

The development of area discrimination and its implications for number representation in infancy.

This paper investigates the ability of infants to attend to continuous stimulus variables and how this capacity relates to the representation of number. We examined the change in area needed by 6-month-old infants to detect a difference in the size of a single element (Elmo face). Infants successfully discriminated a 1:4, 1:3 and 1:2 change in the area of the Elmo face but failed to discriminate a 2:3 change. In addition, the novelty preference was linearly related to the ratio difference between the novel and familiar area. Results suggest that Weber's Law holds for area discriminations in infancy and also reveal that at 6 months of age infants are equally sensitive to number, time and area.

Attention↗

Shared system for ordering small and large numbers in monkeys and humans.

There is increasing evidence that animals share with adult humans and perhaps human infants a system for representing objective number as psychological magnitudes that are an analogue of the quantities they represent. Here we show that rhesus monkeys can extend a numerical rule learned with the values 1 through 9 to the values 10, 15, 20, and 30, which suggests that there is no upper limit on a monkey's numerical capacity. Instead, throughout the numerical range tested, both accuracy and latency in ordering two numerical values were systematically controlled by the ratio of the values compared. In a second experiment, we directly compared humans' and monkeys' performance in the same ordinal comparison task. The qualitative and quantitative similarity in their performance provides the strongest evidence to date of a single nonverbal, evolutionarily primitive mechanism for representing and comparing numerical values.

Animals↗

Re-visiting the competence/performance debate in the acquisition of the counting principles.

Advocates of the "continuity hypothesis" have argued that innate non-verbal counting principles guide the acquisition of the verbal count list (Gelman & Galistel, 1978). Some studies have supported this hypothesis, but others have suggested that the counting principles must be constructed anew by each child. Defenders of the continuity hypothesis have argued that the studies that failed to support it obscured children's understanding of counting by making excessive demands on their fragile counting skills. We evaluated this claim by testing two-, three-, and four-year-olds both on "easy" tasks that have supported continuity and "hard" tasks that have argued against it. A few noteworthy exceptions notwithstanding, children who failed to show that they understood counting on the hard tasks also failed on the easy tasks. Therefore, our results are consistent with a growing body of evidence that shows that the count list as a representation of the positive integers transcends pre-verbal representations of number.

Association Learning↗

Semantic congruity affects numerical judgments similarly in monkeys and humans.

Monkeys (Macaca mulatta) were trained to order visual arrays based on their number of elements and to conditionally choose the array with the larger or smaller number of elements dependent on a color cue. When the screen background was red, monkeys were reinforced for choosing the smaller numerical value first. When the screen background was blue, monkeys were reinforced for choosing the larger numerical value first. Monkeys showed a semantic congruity effect analogous to that reported for human comparison judgments. Specifically, decision time was systematically influenced by the semantic congruity between the cue ("choose smaller" or "choose larger") and the magnitude of the choice stimuli (small or large numbers of dots). This finding demonstrates a semantic congruity effect in a nonlinguistic animal and provides strong evidence for an evolutionarily primitive magnitude-comparison algorithm common to humans and monkeys.

Animals↗

Monkeys match the number of voices they hear to the number of faces they see.

Convergent evidence demonstrates that adult humans possess numerical representations that are independent of language [1, 2, 3, 4, 5 and 6]. Human infants and nonhuman animals can also make purely numerical discriminations, implicating both developmental and evolutionary bases for adult humans' language-independent representations of number [7 and 8]. Recent evidence suggests that the nonverbal representations of number held by human adults are not constrained by the sensory modality in which they were perceived [9]. Previous studies, however, have yielded conflicting results concerning whether the number representations held by nonhuman animals and human infants are tied to the modality in which they were established [10, 11, 12, 13, 14 and 15]. Here, we report that untrained monkeys preferentially looked at a dynamic video display depicting the number of conspecifics that matched the number of vocalizations they heard. These findings suggest that number representations held by monkeys, like those held by adult humans, are unfettered by stimulus modality.

Acoustic Stimulation↗

Analog number representations in mongoose lemurs (Eulemur mongoz): evidence from a search task.

A wealth of data demonstrating that monkeys and apes represent number have been interpreted as suggesting that sensitivity to number emerged early in primate evolution, if not before. Here we examine the numerical capacities of the mongoose lemur (Eulemur mongoz), a member of the prosimian suborder of primates that split from the common ancestor of monkeys, apes and humans approximately 47-54 million years ago. Subjects observed as an experimenter sequentially placed grapes into an opaque bucket. On half of the trials the experimenter placed a subset of the grapes into a false bottom such that they were inaccessible to the lemur. The critical question was whether lemurs would spend more time searching the bucket when food should have remained in the bucket, compared to when they had retrieved all of the food. We found that the amount of time lemurs spent searching was indicative of whether grapes should have remained in the bucket, and furthermore that lemur search time reliably differentiated numerosities that differed by a 1:2 ratio, but not those that differed by a 2:3 or 3:4 ratio. Finally, two control conditions determined that lemurs represented the number of food items, and neither the odor of the grapes, nor the amount of grape (e.g., area) in the bucket. These results suggest that mongoose lemurs have numerical representations that are modulated by Weber's Law.

Analysis of Variance↗

Timing in the baby brain.

Ten-month-old infants and adults were tested in an auditory oddball paradigm in which 50-ms tones were separated by 1500 ms (standard interval) and occasionally 500 ms (deviant interval). Both infants and adults showed marked brain responses to the tone that followed a deviant inter-stimulus interval (ISI). Specifically, the timing-deviance event-related-potential (ERP) difference waves (deviant-ISI ERP minus standard-ISI ERP) yielded a significant, fronto-centrally distributed, mismatch negativity (MMN) in the latency range of 120-240 ms post-stimulus for infants and 110-210 ms for adults. A robust, longer latency, deviance-related positivity was also obtained for infants (330-520 ms), with a much smaller and later deviance-related positivity observed for adults (585-705 ms). These results suggest that the 10-month-old infant brain has already developed some of the same mechanisms as adults for detecting deviations in the timing of stimulus events.

Acoustic Stimulation↗