My research expertise is in the effects of light on human behaviors. During the course of my doctoral work in experimental psychology (majoring in Vision Science), I developed specialist technical knowledge of visual psychophysics, and the measurement of pupil kinetics (pupillometry) as a proxy for retinal and higher order functioning in both healthy individuals and people with disease (e.g., Parkinson's disease). My current postdoctoral training leverages retinal signalling to understand how we can modulate the light pathways that control circadian rhythms and sleep.

Honors & Awards

  • France-Stanford Visiting Junior Scholar Fellow, Lyon Neuroscience Research Center, Université Claude Bernard, Lyon (2017-2018)

Professional Education

  • Bachelor of Science, Queensland University Of Technology (2011)
  • Bachelor of Arts, University Of Queensland (2010)
  • Bachelor of Science, University Of Queensland (2010)
  • Doctor of Philosophy, Queensland University Of Technology (2016)

Stanford Advisors

Community and International Work

  • Social Media Chair, Leadership Team Member, Council Member; Stanford University Postdoctoral Association (SURPAS)

    Ongoing Project


    Opportunities for Student Involvement


All Publications

  • Joyce, D.S., Feigl, B., Kerr, G., & Zele, A.J. Intrinsically photosensitive retinal ganglion cell (ipRGC) mediated pupil function is impaired in Parkinson’s disease bioRxiv Joyce, D. S., Feigl, B., Kerr, G., Roeder, L., Zele, A. J. 2017

    View details for DOI 10.1101/169946

  • Melanopsin-mediated post-illumination pupil response in the peripheral retina. Journal of vision Joyce, D. S., Feigl, B., Zele, A. J. 2016; 16 (8): 5-?


    Intrinsically photosensitive retinal ganglion cells (ipRGCs) regulate pupil size by integrating extrinsic rod and cone signals with intrinsic melanopsin-mediated phototransduction. Light adapted pupil diameter is determined by the corneal flux density (CFD), and for central visual field stimulation the melanopsin-mediated post-illumination pupil response (PIPR) follows this same CFD relationship. Rods, cones, and ipRGCs vary in size, density, and distribution across the retina, but how these differences affect the amplitude and timing of the extrinsic and intrinsic pupil light reflex in the central and peripheral retina is unknown. We determined the relationship between stimulus area and photon flux with stimuli constant for CFD, irradiance, or area at central (0°) and peripheral (20°) eccentricities with high and low melanopsin excitation. We show that the pupil constriction amplitude was similar at both eccentricities and the time to minimum diameter increased as melanopsin excitation increased. In contrast, the peripheral PIPR follows a CFD relationship but with lower amplitude compared with that at the fovea. This indicates differences in the spatial and temporal characteristics of extrinsic and intrinsic ipRGC inputs to the pupil control pathway for the central and peripheral retina. The eccentricity-dependent change in PIPR amplitude may be analogous to the hill of vision observed in visual perimetry; such knowledge is an important precursor to the development of pupil perimetry paradigms to measure the PIPR in select regions of the visual field.

    View details for DOI 10.1167/16.8.5

    View details for PubMedID 27271992

  • The effects of short-term light adaptation on the human post-illumination pupil response Investigative Ophthalmology & Visual Science Joyce, D. S., Feigl, B., Zele, A. J. 2016; 57 (13): 5672-5680

    View details for DOI 10.1167/iovs.16-19934

  • Temporal characteristics of melanopsin inputs to the human pupil light reflex VISION RESEARCH Joyce, D. S., Feigl, B., Cao, D., Zele, A. J. 2015; 107: 58-66


    Rods, cones and melanopsin containing intrinsically photosensitive retinal ganglion cells (ipRGCs) operate in concert to regulate pupil diameter. The temporal properties of intrinsic ipRGC signalling are distinct to those of rods and cones, including longer latencies and sustained signalling after light offset. We examined whether the melanopsin mediated post-illumination pupil response (PIPR) and pupil constriction were dependent upon the inter-stimulus interval (ISI) between successive light pulses and the temporal frequency of sinusoidal light stimuli. Melanopsin excitation was altered by variation of stimulus wavelength (464 nm and 638 nm lights) and irradiance (11.4 and 15.2 log photons cm(-2) s(-1)). We found that 6s PIPR amplitude was independent of ISI and temporal frequency for all melanopsin excitation levels, indicating complete summation. In contrast to the PIPR, the maximum pupil constriction increased with increasing ISI with high and low melanopsin excitation, but time to minimum diameter was slower with high melanopsin excitation only. This melanopsin response to briefly presented pulses (16 and 100 ms) slows the temporal response of the maximum pupil constriction. We also demonstrate that high melanopsin excitation attenuates the phasic peak-trough pupil amplitude compared to conditions with low melanopsin excitation, indicating an interaction between inner and outer retinal inputs to the pupil light reflex. We infer that outer retina summation is important for rapidly controlling pupil diameter in response to short timescale fluctuations in illumination and may occur at two potential sites, one that is presynaptic to extrinsic photoreceptor input to ipRGCs, or another within the pupil control pathway if ipRGCs have differential temporal tuning to extrinsic and intrinsic signalling.

    View details for DOI 10.1016/j.visres.2014.12.001

    View details for Web of Science ID 000348632600007

    View details for PubMedID 25497360

  • Effect of rod-cone interactions on mesopic visual performance mediated by chromatic and luminance pathways JOURNAL OF THE OPTICAL SOCIETY OF AMERICA A-OPTICS IMAGE SCIENCE AND VISION Zele, A. J., Maynard, M. L., Joyce, D. S., Cao, D. 2014; 31 (4): A7-A14


    We studied the effect of rod-cone interactions on mesopic visual reaction time (RT). Rod and cone photoreceptor excitations were independently controlled using a four-primary photostimulator. It was observed that (1) lateral rod-cone interactions increase the cone-mediated RTs; (2) the rod-cone interactions are strongest when rod sensitivity is maximal in a dark surround, but weaker with increased rod activity in a light surround; and (3) the presence of a dark surround nonselectively increased the mean and variability of chromatic (+L-M, S-cone) and luminance (L+M+S) RTs independent of the level of rod activity. The results demonstrate that lateral rod-cone interactions must be considered when deriving mesopic luminous efficiency using RT.

    View details for DOI 10.1364/JOSAA.31.0000A7

    View details for Web of Science ID 000333933600002

    View details for PubMedID 24695205