The Brannon Lab – Numerical Cognition



The Brannon Lab studies the development of numerical cognition across the lifespan. They conduct behavioral and neuroimaging studies with kids and adults. In this way, they hope to learn how our understanding of number changes throughout development and to discover the biological basis of this understanding.

Studies happening now

The Brannon Lab is currently conducting studies at several Philadelphia child organizations. To learn more about your child participating in a study, click here.


Fornaciai, M., Brannon, E.M., Woldorff, M.G. Park, J. Numerosity processing in early visual cortex, in press, Neuroimage

Li, R., Roberts, R. C., Huettel, S. A., & Brannon, E. M. (2017). Five-year-olds do not show ambiguity aversion in a risk and ambiguity task with physical objects. Journal of experimental child psychology, 159, 319-326.

Park, J., Berg, B., Chiang, C. , Woldorff, M. G., & Brannon, E. M. (2017). Developmental trajectory of neural specialization for letter and number visual processing. Developmental Science.

Starr, A., DeWind, N. & Brannon, E. M. The contributions of numerical acuity and non-numerical stimulus features to the development of the number sense and symbolic math achievement, Cognition

Szkudlarek, E. & Brannon, E. M. (2017) Does the Approximate Number System Serve as a Foundation for Symbolic Mathematics? Language Learning and Development. 13:2, 171-190.

Bugden, S., DeWind, N.K., & Brannon, E.M. (2016). Using cognitive training studies to unravel the mechanisms by which the approximate number system supports symbolic math ability. Current Option in Behavioral Sciences. 10:73-80.

Cantlon, J., Merritt, D., & Brannon, E.M. (2016). Monkeys display classic signatures of human symbolic arithmetic. Animal Cognition19:405-415.

DeWind, N. K. & Brannon, E. M. (2016). Significant inter-test reliability across approximate number system assessments. Frontiers in Psychology 7, 1-10.

Drucker,C.B., Rossau, M.A., & Brannon, E. M. (2016). Comparison of discrete ratios by rhesus macaques (Macaca mulatta), Animal Cognition. 19:75­89.

Park, J., Bermudez, V. Roberts, R. C., & Brannon, E. M. (2016). Non-symbolic approximate arithmetic training improves math performance in preschoolers. Journal of Experimental Child Psychology, 152, 278-293.

Dewind, N. K., Adams, G. K., Platt, M. L., & Brannon, E. M. (2015). Modeling the approximate number system to quantify the contribution of visual stimulus features. Cognition, 142, 247-265. 

Drucker, C. B., Rossa, M. A., & Brannon, E. M. (2015). Comparison of discrete ratios by rhesus macaques (Macaca mulatta). Animal Cognition Anim Cogn, 19(1), 75-89. 

Drucker, C. B., & Brannon, E. M. (2015). Commentary on: “Number-space mapping in the newborn chick resembles humans’ mental number line” Frontiers in Psychology Front. Psychol., 6.

Drucker, C. B., Baghdoyan, T., & Brannon, E. M. (2015). Implicit sequence learning in ring-tailed lemurs ( Lemur catta ). Journal of the Experimental Analysis of Behavior Jrnl Exper Analysis Behavior, 105(1), 123-132. 

Li, R., Brannon, E.M., & Huettel, S.A. (2015). Children do not exhibit ambiguity aversion despite intact familiarity bias. Frontiers in Psychology. DOI 10.3389/fpsyg.2014.01519.

Park, J., DeWind, N.K., Woldorff, M.G., & Brannon, E.M. (2015). Rapid and Direct Encoding of Numerosity in the Visual Stream. Cerebral Cortex. DOI 10.1093/cercor/bhv017.

Starr, A., & Brannon, E. M. (2015). Developmental Continuity in the Link Between Sensitivity to Numerosity and Physical Size. Journal of Numerical Cognition J. Numer. Cogn., 1(1), 7-20. 

Starr, A., & Brannon, E. M. (2015). Evidence against continuous variables driving numerical discrimination in infancy. Frontiers in Psychology Front. Psychol., 6. 

Drucker, C. & Brannon, E. M. (2014). Rhesus monkeys (Macaca mulatta) map number onto space. Cognition.

Libertus, M., Starr, A. & Brannon, E.M. (2014). Number trumps area for 7-month old infants. Developmental Psychology. DOI 10.1037/a0032986

Park, J., & Brannon, E.M. (2014). Improving arithmetic performance with number sense training: An investigation of underlying mechanism. Cognition.

Park, J., Chiang, C., Brannon, E.M & Woldorff, M. (2014). Experience-Dependent Hemispheric Specialization of Letters and Numbers is Revealed in Early Visual Processing. Journal of Cognitive Neuroscience. DOI 10.1162/jocn_a_00621

Pinhas, M., Donohue, S. H., Woldorff, M. G., & Brannon, E. M. (2014). Electrophysiological evidence for the involvement of the approximate number system in preschoolers’ processing of spoken number words. Journal of Cognitive Neuroscience.

Starr, A., & Brannon, E.M. Evolutionary and Developmental Continuities in Numerical Cognition, In Ed.s Geary, D, Berch, D.

Jones, S. M., Pearson, J., DeWind, N., Paulsen, D., Tenekedjieva, A. & Brannon, E.M. (2013). Lemurs and macaques show similar numerical sensitivity. Animal Cognition DOI 10.1007/s10071-013-0682-3

Merritt, D.J. & Brannon, E.M. (2013). Nothing to it: Precursors to a zero concept in preschool children. Behavioural Processes, 93:91-97. PMCID: PMC 3582820.

Park, J., Li, R. & Brannon, E.M. (2013). Neural connectivity patterns underlying symbolic number processing indicate mathematical achievement in children. Developmental Science. DOI 10.1111/desc.12114

Park, J. & Brannon, E.M. (2013). Training the Approximate Number System Improves Math Proficiency. Psychological Science. NIHMSID 460715

Starr, A.B., Libertus, M.E., & Brannon, E.M. (2013). Infants show ratio dependent discrimination regardless of set size. Infancy, 1-15. NIHMSID 440920.

Starr, A., Libertus, M. & Brannon, E.M. (2013). Number sense in infancy predicts mathematical abilities in childhood. Proceedings of the National Academy of Science.DOI 10.1073/pnas.1302751110

Jones, S.M., & Brannon, E.M., (2012). Prosimian primates show ratio dependence in spontaneous quantity discriminations. Frontiers in Comparative PsychologyVolume 3: Article 550, 1-8. DOI: 10.3389/fpsyg.2012.00550

Roitman, J.D., Brannon, E.M., & Platt, M.L, (2012). Representation of numerosity in posterior parietal cortex. Frontiers in Integrative Neuroscience Volume 6: Article 25, 1-9. DOI: 10.3389/fnint.2012.00025

DeWind, N.K., & Brannon, E.M. Malleability of the approximate number system: effects of feedback and training. Frontiers in Human Neuroscience. Volume 6: Article 68, 1-10. DOI: 10.3389/fnhum.2012.00068

MacLean, E.L., Mandalaywala, T.M., Brannon, E.M., (2012) Variance-sensitive choice in lemurs: constancy trumps quantity. Animal CognitionVolume 15, Number 1, 15-25, DOI: 10.1007/s10071-011-0425-2

Paulsen, D.J., Carter, M., Platt, M.L., Huettel, S.A., &. Brannon, E.M., (2012). Neurocognitive development of risk aversion from early childhood to adulthood. Frontiers in Human Neuroscience, 5:178. DOI: 10.3389/fnhum.2011.00178

Cantlon, J.F. Davis, S.W., Libertus, M.E. Kahane, J. Brannon, E.M. Pelphrey, K.A. (2011) Inter-Parietal White Matter Development Predicts Numerical Performance in Young Children, Learning and Individual Differences, 21(6), 672-680

Cordes, S. & Brannon, E. M. (2011). Attending to one of many: When infants are surprisingly poor at discriminating an item’s size. Frontiers in Developmental Psychology, 2:65. doi: 10.3389/fpsyg.2011.00065

Libertus, M.E., Brannon, E.M., Woldorff, M. (2011). Parallels in stimulus-driven oscillatory brain responses to numerosity changes in 7-month-old infants and adults.Developmental Neuropsychology, 36(6), 651-667

MacLean, E.L., Matthews, L.J., Hare, B. Nunn, C.L., Anderson, R.C., Aureli, F. Brannon, E.M., Call, J., Drea, C.M., Emery, N.J., Haun, D.B.M., Herrmann, E., Jacobs, L.F., Platt, M.L., Rosati, A.G., Sandel, A., Schroepfer, K.K., Seed, A.M., Tan, J., van Schaik, C.P., Wobber, V., (2011) How Does Cognition Evolve?: Phylogenetic Comparative Psychology, Animal Cognition, DOI 10.1007/s10071-011-0448-8

Merritt, D. MacLean, E. Crawford, J.C. Brannon, E. M. (2011). Numerical rule-learning in ring-tailed Lemurs (Lemur catta), Frontiers in Comparative Psychology, 2:23, 1-9.

Paulsen, D.J., Platt, M.L., Huettel, S.A., &. Brannon, E.M. (2011). Decision-making Under Risk in Children, Adolescents, and Young Adults, Frontiers in Developmental Psychology(2), 72, 10.3389/fpsyg.2011.00072

Brannon, E.M. (2010) Introduction to thought without language: A tribute to the contributions of H.S. Terrace. Behavioural Processes, 82 (2) 137-138.

Cantlon, J. F., Safford, K.E., & Brannon, E.M. (2010). Spontaneous analog number representations in 3-year-old children. Developmental Science, 13 (2) 289-297.

Dehaene, S. & Brannon, E.M. (2010) Space, time, and number: A Kantian research program. Trends in Cognitive Sciences.

Jones, S.M., Merritt, D., Cantlon, J., & Brannon, E.M. (2010). Context affects the numerical semantic congruity effect in rhesus monkeys.Behavioral Processes, 82(2) 191-196.

Libertus, M., Brannon, E. (2010). Stable individual differences in number discrimination in infancy. Developmental Science.1-7.

Merritt, D., Casasanto, D., Brannon, E.M. (2010). Do monkeys think in metaphors? Representations of space and time in monkeys and humans. Cognition.117, 191-202.

Paulsen, D., Woldorff, M. & Brannon, E.M. (2010). Individual Differences in Non-verbal number discrimination correlate. Neuropsychologia.48, 3687-3695.

Pearson, J., Roitman, J.D. Brannon, E.M. Platt, M.L. & Raghavachari, S. (2010). A physiologically-inspired model of numerical classification based on graded stimulus coding. Frontiers in Behavioral Neuroscience, vol. 4(1) PMCID: PMC2814553

Cantlon, J. F., Cordes, S., Libertus, M. E., & Brannon, E. M. (2009). Numerical abstraction: It ain’t broke. (commentary).Behavioral and Brain Sciences, 32:331-332.

Cantlon, J. F., Cordes, S., Libertus , M. E. , Brannon, E. M. (2009) Comment on Log or Linear? Distinct Intuitions of the Number Scale in Western and Amazonian Indigene Cultures, Science, 323(5910) 38b.

Cantlon, J. F., Libertus, M.E., Pinel, P., Dehaene, S., Brannon, E.M., Pelphrey, K.A. (2009). The neural development of an abstract concept of number. Journal of Cognitive Neuroscience 21(11), 2217-2229.

Cantlon, J. F., Platt, M. L., & Brannon, E. M. (2009). Beyond the number domain. Invited review. Trends in Cognitive Sciences, 13 (2) 83-91.

Cordes, S., & Brannon, E.M. (2009). Crossing the divide: Infants discriminate small from large numerosities. Developmental Psychology, 45(6), 1583-1594.

Cordes, S., & Brannon, E. M. (2009). The relative salience of discrete and continuous quantity in young infants.Developmental Science, 12 (3), 453-463.

Libertus, M.E., & Brannon, E.M. (2009) Behavioral and neural basis of number sense in infancy, Current Directions in Psychological Science, 18(6)346-351.

Libertus, M., Brannon, E. M. , & Pelphrey, K. (2009). Developmental changes in category-specific brain responses to numbers and letters in a working memory task . Neuroimage, 44 (4), 1404-1414.

Libertus, M., Pruitt, L., Woldorff, M., Brannon, E.M. (2009). Induced alpha-band oscillations reflect ratio-dependent number discrimination in the infant brain. Journal of Cognitive Neuroscience, 21(12), 2398-2406.

MacLean, E. L., Prior, S. R., Platt, M. L., & Brannon, E. M. (2009). Primate location preference in a double-tier cage: The effects of illumination and cage height. Journal of Applied Animal Welfare Science, 12 (1), 73-81.

Merritt, D., Rugani, R., Brannon, E.M. (2009). Empty sets and the representation of number in macaques. Journal of Experimental Psychology: General, 138(2), 258-269.

Brannon, E. M., Libertus, M., Meck, W. H., & Woldorff, M. (2008). Electrophysiological measures of time processing in infant and adult brains: Weber’s law holds. Journal of Cognitive Neuroscience, 20(2) , 193-203.

Cordes, S., & Brannon, E. M. (2008). The difficulties of representing continuous extent in infancy: Using number is just easier. Child Development, 79(2), 476-489.

Cordes, S., & Brannon, E. M. (2008). Quantitative competencies in infancy. Developmental Science, 11(6), 803-808.

Jordan, K. E., MacLean, E. L., & Brannon, E. M. (2008). Monkeys match and tally quantities across senses. Cognition, 108(3), 617-625.

Jordan, K. E., Suanda, S. H., & Brannon, E. M. (2008). Internseory redundancy accelerates preverbal numerical competence. Cognition, 108(1), 210-221.

MacLean, E., Merritt, D., & Brannon, E. M. (2008). Social complexity predicts transitive reasoning in prosimian primates. Animal Behavior, 76, 479-486.

Suanda, S. H., Tompson, W., & Brannon, E. M. (2008). Changes in the ability to detect ordinal numerical relationships between 9 and 11 months of age. Infancy, 13(4), 308-337.

Brannon, E. M., Suanda, S., & Libertus, K. (2007). Temporal discrimination increases in precision over development and parallels the development of numerosity discrimination. Developmental Science, 10(6), 770-777.

Cantlon, J. F., & Brannon, E. M. (2007). Adding up the effects of cultural experience on the brain. Trends in Cognitive Sciences, 11(1), 1-4.

Cantlon, J. F., & Brannon, E. M. (2007). Basic math in monkeys and college students. PLoS Biology, 5(12), e328.

Cantlon, J. F., & Brannon, E. M. (2007). How much does number matter to a monkey (Macaca mulatta)? Journal of Experimental Psychology: Animal Behavior Processes, 33(1), 32-41.

Cantlon, J. F., Fink, R., Safford, K., & Brannon, E. M. (2007). Heterogeneity impairs numerical matching but not numerical ordering in preschool children.. Developmental Science, 10(4), 431-440.

Libertus, M. E., Woldorff, M. G., & Brannon, E. M. (2007). Electrophysiological evidence for notation independence in numerical processing. Behavioral and Brain Functions, 3(1).

Merritt, D., MacLean, E. L., Jaffe, S., & Brannon, E. M. (2007). A comparative analysis of serial ordering in ring-tailed lemurs (Lemur catta). Journal of Comparative Psychology, 121(4), 363-371.

Roitman, J., Brannon, E. M., Andrews, J. R., & Platt, M. L. (2007). Nonverbal representation of time and number in adults. Acta Psychologica, 124, 296-318.

Roitman, J., Brannon, E. M., & Platt, M. L. (2007). Monotonic coding of numerosity in macaque lateral intraparietal area. PLoS Biolology, 5(8), e208.

Brannon, E. M. (2006). The representation of numerical magnitude. Invited review for Current Opinion in Neurobiology, 16, 222-229.

Brannon, E. M., Cantlon, J. F., & Terrace, H. S. (2006). The role of reference points in ordinal numerical comparisons by Rhesus macaques (Macaca mulatta) . Journal of Experimental Psychology: Animal Behavior Processes, 32(2), 120-134.

Brannon, E. M., Lutz, D., and Cordes, S. (2006). The development of area discrimination and its implications for number representation in infancy. Developmental Science, 9, F59-F64.

Cantlon, J., & Brannon, E. M. (2006). The effect of heterogeneity on numerical ordering in rhesus monkeys, Infancy, 9(2), 173-189.

Cantlon, J., & Brannon, E. M. (2006). Shared system for ordering small and large numbers in monkeys and humans. Psychological Science, 17(5), 401-406.

Cantlon, J., Brannon, E. M., Carter, E. J., & Pelphrey, K. (2006). Functional imaging of numerical processing in adults and 4-year-old children. PLOS Biology, 4(5), e125.

Jordan, K., & Brannon, E. M. (2006). The multisensory representation of number in infancy. Proceedings of the National Academy of Sciences, 103(9), 3486-3489.

Jordan, K., & Brannon, E.M. (2006). A common representational system governed by Weber’s Law: Nonverbal numerical similarity judgments in six-year-old children and rhesus macaques. Journal of Experimental Child Psychology, 95, 215 229.

Jordan, K., & Brannon, E. M. (2006). Weber’s Law influences numerical representations in rhesus macaques (Macaca mulatta). Animal Cognition, 9(3), 159-172.

Le Corre, M., Van de Walle, G., Brannon, E. M., & Carey, S. (2006). Re-visiting the competence/performance debate in the acquisition of the counting principles. Cognitive Psychology, 52(2), 130-169.

Brannon, E. M. (2005). The independence of language and mathematical reasoning. Proceedings of the National Academy of Sciences, 109(9), 3177-3178.

Cantlon, J., & Brannon, E. M. (2005). Semantic congruity affects numerical judgments similarly in monkeys and humans. Proceedings of the National Academy of Sciences, 102(45), 16507-16511.

Jordan , K. E., Brannon, E. M., Logothetis, N. K., Ghazanfar, A. A. (2005). Monkeys match the number of voices they hear to the number of faces they see. Current Biology, 15, 1-5.

Lewis, K., Jaffe, S., & Brannon, E. M. (2005). Analog number representations in mongoose lemurs (Eulemur mongoz): Evidence from a search task. Animal Cognition, 8(4), 247-251.

Brannon, E. M., Abbott, S., & Lutz, D. (2004). Number bias for the discrimination of large visual sets in infancy, Cognition, 93, B59-B68.

Brannon, E. M., Andrews, M., & Rosenblum, L. (2004). Effectiveness of video of conspecifics as a reward for socially housed bonnet macaques (Macaca radiata). Perceptual and Motor Skills, 98, 849-858.

Brannon, E. M., Wolfe, L., Meck, W. H., & Woldorff, M. (2004). Timing in the baby brain. Cognitive Brain Research, 21, 227-233.

Brannon, E. M. (2003). Number knows no bounds. Invited commentary in Trends in Cognitive Sciences, 7(7), 279-281.

Terrace, H. S., Son, L., & Brannon, E. M. (2003). Serial expertise of rhesus macaques. Psychological Science, 14(1), 66-73.

Brannon, E. M. (2002). The development of ordinal numerical knowledge in infancy. Cognition, 83, 223-240.

Brannon, E. M., & Van de Walle, G. (2001). Ordinal numerical knowledge in young children. Cognitive Psychology, 43, 53-81.

Brannon, E. M., Wusthoff, C. J., Gallistel, C. R., & Gibbon, J. (2001). Subtraction in the pigeon: Evidence for a linear subjective number scale. Psychological Science, 12(3), 238-243.

Gallistel, C. R., Brannon, E. M., Gibbon, J., & Wusthoff, C. J. (2001). Response to Dehaene’s Commentary.Psychological Science, 12(3), 247.

Brannon, E. M., & Terrace, H. S. (2000). Representation of the numerosities 1-9 by Rhesus Monkeys (Macaca mulatta). Journal of Experimental Psychology: Animal Behavior Processes, 26(1), 31-49.

Brannon, E. M., & Terrace, H. S. (1999). Letter to the editor. Science, 283, 1852.

Brannon, E. M., & Terrace, H. S. (1998). Ordering of the numerosities 1-9 by monkeys. Science, 282, 746-749.

Platt, M. L., Brannon , E. M., Briese, T. L. & French, J. A. (1996). Differences in feeding ecology predict differences in performance between golden lion tamarins (Leontopithecus rosalia) and Wied’s marmosets (Callithrix kuhli) on spatial and visual memory tasks. Animal Learning and Behavior, 24(4), 384-393.

Books and Book Chapters:

Brannon, E.M., & Park, J. (2015). Navigator Chapter for: Phylogeny and Ontogeny of Mathematical and Numerical understanding. In Handbook on Mathematical Cognition, Ed.s R.Cohen-Kadosh.

Merritt, D., DeWind, N., & Brannon, E.M. (in press) Comparative cognition of number representation, In Handbook of Comparative Cognition.Editors, Thomas Zentall and Ed Wasserman.

Brannon, E.M. & Merritt, D. (2011) Evolutionary foundations of the Approximate Number System. In Space, Time, and Number in the Brain: searching for the foundations of mathematical thought. Editors, Dehaene, S., & Brannon, E.M. Elsevier (2011).

Dehaene, S. Brannon, E.M. Editors (2011) Space, Time, and Number in the Brain: searching for the foundations of mathematical thought. Elsevier.

Cantlon, J. F., & Brannon, E. M. (2011). Animal Arithmetic. In Encyclopedia of Animal Behavior. Editors, Michael Breed and Janice Moore. Oxford: Elsevier Press.

Brannon, E.M., Jordan, K.E., Jones, S. (2010). Behavioral Signatures of Numerical Discrimination.Primate Neuroethology, Editors, Michael Platt and Asif Ghazanfar. Oxford Press.

Dehaene, S. & Brannon, E.M., Editors (2010). Special Issue on Space, Time, and Number, Trends in Cognitive Sciences.

Brannon, E.M, & Cantlon, J. F. (2009). A comparative perspective on the origin of numerical thinking. In Cognitive Biology: Evolutionary and Developmental Perspectives on Mind, Brain, and Behavior, Luca Tomasi, Mary A. Peterson, and Lynn Nadel (Eds.). Cambridge: MIT Press.

Jordan, K.E., & Brannon, E.M. (2009) A comparative approach to understanding human numerical cognition, In The Origins of Object Knowledge Bruce Hood and Laurie Santos (Eds.). Oxford University Press, Oxford.

Purves, D., Brannon, E., Cabeza, R., Huettel, S., LaBar, K., Platt, M., Woldorff, M. (2007). Principles of Cognitive Neuroscience. Sunderland , MA: Sinauer Associates. [Undergraduate/Graduate Textbook]

Brannon, E. M. (2005). Quantitative thinking: From monkey to human and human infant to adult. In S. Dehaene, J. Duhamel, M. D. Hauser, & G. Rizzolatti (Eds.), From Monkey Brain to Human Brain (pp. 97-116). Cambridge, MA: MIT Press.

Brannon, E. M. (2005). What Animals Know About Numbers. In J. I. D. Campbell (Ed.), Handbook of Mathematical Cognition (pp. 85-108). New York, NY: Psychology Press.

Brannon, E. M., & Roitman, J. (2003). Nonverbal representations of time and number in animals and human infants. In W. Meck (Ed.), Functional and Neural Mechanisms of Interval Timing (pp. 143-182). New York, NY: CRC Press.

Brannon, E. M., & Terrace, H. S. (2002). The Evolution and Ontogeny of Ordinal Numerical Ability. In M. Bekoff, C. Allen, & G. M. Burghardt (Eds.), The Cognitive Animal (pp. 197-204). Cambridge, MA: The MIT Press.