Professional Education

  • Master of Arts, University of Saint Andrews (2012)
  • Doctor of Philosophy, University of Pennsylvania (2016)

Stanford Advisors

Lab Affiliations

All Publications

  • Variation of outdoor illumination as a function of solar elevation and light pollution SCIENTIFIC REPORTS Spitschan, M., Aguirre, G. K., Brainard, D. H., Sweeney, A. M. 2016; 6


    The illumination of the environment undergoes both intensity and spectral changes during the 24 h cycle of a day. Daylight spectral power distributions are well described by low-dimensional models such as the CIE (Commission Internationale de l'Éclairage) daylight model, but the performance of this model in non-daylight regimes is not characterised. We measured downwelling spectral irradiance across multiple days in two locations in North America: One rural location (Cherry Springs State Park, PA) with minimal anthropogenic light sources, and one city location (Philadelphia, PA). We characterise the spectral, intensity and colour changes and extend the existing CIE model for daylight to capture twilight components and the spectrum of the night sky.

    View details for DOI 10.1038/srep26756

    View details for Web of Science ID 000377553100001

    View details for PubMedID 27272736

  • Human Visual Cortex Responses to Rapid Cone and Melanopsin-Directed Flicker JOURNAL OF NEUROSCIENCE Spitschan, M., Datta, R., Stern, A. M., Brainard, D. H., Aguirre, G. K. 2016; 36 (5): 1471-1482


    Signals from cones are recombined in postreceptoral channels [luminance, L + M; red-green, L - M; blue-yellow, S - (L + M)]. The melanopsin-containing retinal ganglion cells are also active at daytime light levels and recent psychophysical results suggest that melanopsin contributes to conscious vision in humans. Here, we measured BOLD fMRI responses to spectral modulations that separately targeted the postreceptoral cone channels and melanopsin. Responses to spatially uniform (27.5° field size, central 5° obscured) flicker at 0.5, 1, 2, 4, 8, 16, 32, and 64 Hz were recorded from areas V1, V2/V3, motion-sensitive area MT, and the lateral occipital complex. In V1 and V2/V3, higher temporal sensitivity was observed to L + M + S (16 Hz) compared with L - M flicker (8 Hz), consistent with psychophysical findings. Area MT was most sensitive to rapid (32 Hz) flicker of either L + M + S or L - M. We found S cone responses only in areas V1 and V2/V3 (peak frequency: 4-8 Hz). In addition, we studied an L + M modulation and found responses that were effectively identical at all temporal frequencies to those recorded for the L + M + S modulation. Finally, we measured the cortical response to melanopsin-directed flicker and compared this response with control modulations that addressed stimulus imprecision and the possibility of stimulation of cones in the shadow of retinal blood vessels (penumbral cones). For our stimulus conditions, melanopsin flicker did not elicit a cortical response exceeding that of the control modulations. We note that failure to control for penumbral cone stimulation could be mistaken for a melanopsin response.The retina contains cone photoreceptors and ganglion cells that contain the photopigment melanopsin. Cones provide brightness and color signals to visual cortex. Melanopsin influences circadian rhythm and the pupil, but its contribution to cortex and perception is less clear. We measured the response of human visual cortex with fMRI using spectral modulations tailored to stimulate the cones and melanopsin separately. We found that cortical responses to cone signals vary systematically across visual areas. Differences in temporal sensitivity for achromatic, red-green, and blue-yellow stimuli generally reflect the known perceptual properties of vision. We found that melanopsin signals do not produce a measurable response in visual cortex at temporal frequencies between 0.5 and 64 Hz at daytime light levels.

    View details for DOI 10.1523/JNEUROSCI.193-15.2016

    View details for Web of Science ID 000369182200005

    View details for PubMedID 26843631

  • Eye fixation during multiple object attention is based on a representation of discrete spatial foci. Scientific reports Fluharty, M., Jentzsch, I., Spitschan, M., Vishwanath, D. 2016; 6: 31832-?


    We often look at and attend to several objects at once. How the brain determines where to point our eyes when we do this is poorly understood. Here we devised a novel paradigm to discriminate between different models of spatial selection guiding fixation. In contrast to standard static attentional tasks where the eye remains fixed at a predefined location, observers selected their own preferred fixation position while they tracked static targets that were arranged in specific geometric configurations and which changed identity over time. Fixations were best predicted by a representation of discrete spatial foci, not a polygonal grouping, simple 2-foci division of attention or a circular spotlight. Moreover, attentional performance was incompatible with serial selection. Together with previous studies, our findings are compatible with a view that attentional selection and fixation rely on shared spatial representations and suggest a more nuanced definition of overt vs. covert attention.

    View details for DOI 10.1038/srep31832

    View details for PubMedID 27561413

  • Selective Stimulation of Penumbral Cones Reveals Perception in the Shadow of Retinal Blood Vessels PLOS ONE Spitschan, M., Aguirre, G. K., Brainard, D. H. 2015; 10 (4)


    In 1819, Johann Purkinje described how a moving light source that displaces the shadow of the retinal blood vessels to adjacent cones can produce the entopic percept of a branching tree. Here, we describe a novel method for producing a similar percept. We used a device that mixes 56 narrowband primaries under computer control, in conjunction with the method of silent substitution, to present observers with a spectral modulation that selectively targeted penumbral cones in the shadow of the retinal blood vessels. Such a modulation elicits a clear Purkinje-tree percept. We show that the percept is specific to penumbral L and M cone stimulation and is not produced by selective penumbral S cone stimulation. The Purkinje-tree percept was strongest at 16 Hz and fell off at lower (8 Hz) and higher (32 Hz) temporal frequencies. Selective stimulation of open-field cones that are not in shadow, with penumbral cones silenced, also produced the percept, but it was not seen when penumbral and open-field cones were modulated together. This indicates the need for spatial contrast between penumbral and open-field cones to create the Purkinje-tree percept. Our observation provides a new means for studying the response of retinally stabilized images and demonstrates that penumbral cones can support spatial vision. Further, the result illustrates a way in which silent substitution techniques can fail to be silent. We show that inadvertent penumbral cone stimulation can accompany melanopsin-directed modulations that are designed only to silence open-field cones. This in turn can result in visual responses that might be mistaken as melanopsin-driven.

    View details for DOI 10.1371/journal.pone.0124328

    View details for Web of Science ID 000353212600063

    View details for PubMedID 25897842

  • Opponent melanopsin and S-cone signals in the human pupillary light response PROCEEDINGS OF THE NATIONAL ACADEMY OF SCIENCES OF THE UNITED STATES OF AMERICA Spitschan, M., Jain, S., Brainard, D. H., Aguirre, G. K. 2014; 111 (43): 15568-15572


    In the human, cone photoreceptors (L, M, and S) and the melanopsin-containing, intrinsically photosensitive retinal ganglion cells (ipRGCs) are active at daytime light intensities. Signals from cones are combined both additively and in opposition to create the perception of overall light and color. Similar mechanisms seem to be at work in the control of the pupil's response to light. Uncharacterized however, is the relative contribution of melanopsin and S cones, with their overlapping, short-wavelength spectral sensitivities. We measured the response of the human pupil to the separate stimulation of the cones and melanopsin at a range of temporal frequencies under photopic conditions. The S-cone and melanopsin photoreceptor channels were found to be low-pass, in contrast to a band-pass response of the pupil to L- and M-cone signals. An examination of the phase relationships of the evoked responses revealed that melanopsin signals add with signals from L and M cones but are opposed by signals from S cones in control of the pupil. The opposition of the S cones is revealed in a seemingly paradoxical dilation of the pupil to greater S-cone photon capture. This surprising result is explained by the neurophysiological properties of ipRGCs found in animal studies.

    View details for DOI 10.1073/pnas.1400942111

    View details for Web of Science ID 000343729500073

    View details for PubMedID 25313040

  • Perceptual integration across natural monocular regions JOURNAL OF VISION Zeiner, K. M., Spitschan, M., Harris, J. M. 2014; 14 (3)


    Natural scenes contain hidden regions, or occlusions, that differ in the two eyes, resulting in monocular regions that can only be seen by one eye. Such monocular regions appear to not be suppressed but seem to be integrated into the scene percept. Here we explore how the two eyes' views are combined to represent a scene that contains monocular regions, partially hidden behind a foreground occluding "fence." We measured performance in a density/numerosity discrimination task for scenes containing differing amounts of binocular and monocular information. We find that information from a number of separate monocular regions can be integrated into our overall percept of dot density/numerosity, although different observers use different strategies. If, however, both monocular and binocular information is present, observers appear to ignore the purely monocular regions, relying solely on the binocular information when making density/numerosity judgments. Our work suggests that binocular regions are favored over monocular regions, such that information from monocular regions is effectively ignored when binocular regions are present in a scene.

    View details for DOI 10.1167/14.3.5

    View details for Web of Science ID 000334344500005

    View details for PubMedID 24599943