David H. Brainard


417 Goddard Labs
3710 Hamilton Walk


David H. Brainard is the RRL Professor of Psychology at the University of Pennsylvania. He received an AB in Physics (Magna Cum Laude) from Harvard University (1982) and an MS (Electrical Engineering) and PhD (Psychology) from Stanford University in 1989. His research focuses on color vision, intrinsically photosensitive retinal ganglion cells, retinal imaging, as well as computational models thereof. He is a fellow of the Optical SocietyARVO and the Association for Psychological Science, as well as an elected member of the Society of Experimental Psychologists and the National Academy of Sciences USA. He is currently a member of the editorial board of the Journal of Vision and co-editor of the Annual Review of Vision Science.

Research Interests

The Brainard Lab studies human vision, both experimentally and through computational modeling of visual processing. Our primary concern is with how the visual system estimates object properties from the information available in the light signal incident at the eye. To study this general problem, we conduct psychophysical experiments to investigate questions such as how object color appearance is related to object surface properties under a wide range of illumination conditions and how color is used to identify objects, and formulate computational models of the results. In addition, we are interested in developing machine visual systems that can mimic human performance and in understanding the neural mechanisms of vision.

Representative Recent Publications

(See Publications for more, or download Brainard’s CV in PDF Format. Or visit Brainard’s Google Scholar Page).

Twomey, C. R., Roberts, G., Brainard, D., Plotkin, J. B. (2021), What we talk about when we talk about color, PNAS,

McAdams, H., Kaiser, E. A., Igdalova, A., Haggerty, E. B.., Cucchiara, B., Brainard, D. H., Aguirre, G. K. (2020). Selective amplification of ipRGC signals accounts for interictal photophobia in migraine, PNAS, https://doi.org/10.1073/pnas.2007402117Download PDF.

Cottaris, N. P., Wandell, B. A., Reike, F., Brainard, D. H. (2020). A computational observer model of spatial contrast sensitivity: Effects of photocurrent encoding, fixational eye movements, and inference engine. Journal of Vision, 20(7):17, https://doi.org/10.1167/jov.20.7.17.

Brainard, D. H. (2019). Color, pattern, and the retinal cone mosaic. Current Opinion in Behavioral Sciences, 30:41–47, https://doi.org/10.1016/j.cobeha.2019.05.005. Download PDF.

Ruff, D. Brainard, D. H., Cohen, M. R. (2018). Neuronal population mechanisms of lightness perception. Journal of Neurophysiology, 120(5), 2296-2310, doi: 10.1152/jn.00906.2017. Download PDF. Preprint available at https://doi.org/10.1101/294280.

Tuten, W. S., Cooper, R. F., Tiruveedhula, P., Dubra, A., Roorda, A., Cottaris, N. P., Brainard, D. H., Morgan, J. I. W. (2018). Spatial summation in the human fovea: do normal optical aberrations and fixational eye movements have an effect? Journal of Vision, 18(8):6, doi: 10.1167/18.8.6, https://jov.arvojournals.org/article.aspx?articleid=2697364.

Brainard, D. H., Cottaris, N. P., Radonjić, A. (2018). The perception of color and material in natural tasks. Royal Society Interface Focus, 8(4), doi: 10.1098/rsfs.2018.0012. Download PDF.

Cooper, R. F., Tuten, W. S., Dubra, A., Brainard, D. H., Morgan, J. I. W. (2017). Non-invasive assessment of human cone photoreceptor function. Biomedical Optics Express, 8(11), 5098-5112, doi: 10.1364/BOE.8.005098. https://www.osapublishing.org/boe/abstract.cfm?uri=boe-8-11-5098. Erratum: Biomed. Opt. Express, 2018, 9, 1842, https://doi.org/10.1364/BOE.9.001842.

Barnett, M. A., Aguirre, G. K., Brainard, D. H. (2021). A quadratic model captures the human V1 response to variations in chromatic direction and contrast, eLife, 2021;10:e65590, https://doi.org/10.7554/eLife.65590.

Kaiser, E. A., McAdams, H., Igdalova, A., Haggerty, E. B., Cucchiara, B., Brainard, D. H., Aguirre, G. K. (2021). Reflexive eye closure in response to cone and melanopsin stimulation: A study of implicit measures of light sensitivity in migraine, Neurology, https://doi.org/10.1212/WNL.0000000000012734. Download PDF.

Zhang, L., Cottaris, N. P., Brainard, D. H. (2022). An image reconstruction framework for characterizing early vision. eLife, 2022;11:e71132, https://doi.org/10.7554/eLife.71132.

Singh, V., Burge, J., Brainard, D. H. (2022). Equivalent noise characterization of human lightness constancy. Journal of Vision, 22(5):2, doi: https://doi.org/10.1167/jov.22.5.2.

Cooper, R. F., Brainard, D. H., Morgan, J. I. W. (2020). Optoretinography of individual human cone photoreceptors. Optics Express, 28, 39326-39339, https://www.osapublishing.org/oe/fulltext.cfm?uri=oe-28-26-39326&id=444748.

Vincent, J., Haggerty, E. B., Brainard, D. H., Aguirre, G. K. (2021). Melanopic stimulation does not affect psychophysical threshold sensitivity for luminance flicker. Scientific Reports, 11, 20167, https://www.nature.com/articles/s41598-021-99684-0.

Wandell, B. A., Brainard, D. H. (2023). Principles and consequences of the initial visual encoding. In The New Handbook of Mathematical Psychology, Vol. 3 (Ashby, F. G., Colonius, H., Dzhafarov, E. N., eds.).  Cambridge University Press.  Download PDF.

Godat, T., Cottaris, N. P., Patterson, S., Kohout, K., Parkins, K., Yang, Q., Strazzeri, J. M., McGregor, J. E., Brainard, D. H., Merigan, W. H., Williams, D. R. (2022). In vivo physiology of foveal retinal ganglion cells in Macaca fascicularis. PLoS One, 17(11), e0278261, https://doi.org/10.1371/journal.pone.0278261.

Chen, M., Jiang, Y. Y., Gee J. C., Brainard, D. H., Morgan, J. I. W. (2022), Automated assessment of photoreceptor visibility in adaptive optics split-detection images using edge detection. Trans. Vis. Sci. Tech., 11(5):25. doi: https://doi.org/10.1167/tvst.11.5.25.

Murray R. F., Brainard D. H., Patel J. Y., Weiss E., Patel K. Y. (2022). An intrinsic image network with properties of human lightness perception. Proceedings of the Color and Imaging Conference 30. Download PDF.

Murray R. F., Brainard D. H,. Flachot A., Patel J. Y. (2023). An intrinsic image network evaluated as a model of human lightness perception. Proceedings of Human Vision and Electronic Imaging 2023. Download PDF.

Twomey, C. R., Brainard, D. H., Plotkin, J. B. (2024). Historical constraints on the evolution of color naming. PNAS, 121 (10) e2313603121, https://doi.org/10.1073/pnas.2313603121.  Preprint available here: arXiv:2305.04345.  Story about this work in Penn Today.

Wandell, B. A., Goossens, T., Brainard, D. H. (2024). Deriving the cone fundamentals: a subspace intersection method. https://www.biorxiv.org/content/10.1101/2024.02.04.577470v1.

Srinath, R., Ni, A. M., Marucci, C., Cohen, M. R., Brainard, D. H. (2024). Orthogonal neural representations support perceptual judgements of natural stimuli. https://doi.org/10.1101/2024.02.14.580134.

Barnett, M. A., Chin, B. M., Aguirre, G. K., Burge, J., Brainard, D. H. (2024). Temporal dynamics of human color processing measured using a continuous tracking task, https://www.biorxiv.org/content/10.1101/2024.03.01.582975v1.

Nankivil, D., Cottaris, N. P., & Brainard, D. H. (2024). Theoretical impact of chromatic aberration correction on visual acuity.  Biomedical Optics Express, 15, 3265-3284, https://doi.org/10.1364/BOE.516049. Code and data associated with this paper: https://github.com/DavidBrainard/ISETBioJandJ.git.

Zhang, L., Kadkhodaie, Z., Simoncelli, E. P., Brainard, D. H. (2024). Optimized Linear Measurements for Inverse Problems using Diffusion-Based Image Generation, arXiv.2405.17456, https://arxiv.org/abs/2405.17456.

Preregistered Experiments

Here is a link to our list of preregistered experiments.