Saturation in phosphene size with increasing current levels delivered to human visual cortex
Open accessAbstract
Electrically stimulating early visual cortex results in a visual percept known as a phosphene. Although phosphenes can be evoked by a wide range of electrode sizes and current amplitudes, they are invariably described as small. To better understand this observation, we electrically stimulated 93 electrodes implanted in the visual cortex of 13 human subjects who reported phosphene size while stimulation current was varied. Phosphene size increased as the stimulation current was initially raised above threshold, but then rapidly reached saturation. Phosphene size also depended on the location of the stimulated site, with size increasing with distance from the foveal representation. We developed a model relating phosphene size to the amount of activated cortex and its location within the retinotopic map. First, a sigmoidal curve was used to predict the amount of activated cortex at a given current. Second, the amount of active cortex was converted to degrees of visual angle by multiplying by the inverse cortical magnification factor for that retinotopic location. This simple model accurately predicted phosphene size for a broad range of stimulation currents and cortical locations. The unexpected saturation in phosphene sizes suggests that the functional architecture of cerebral cortex may impose fundamental restrictions on the spread of artificially evoked activity and this may be an important consideration in the design of cortical prosthetic devices. SIGNIFICANCE STATEMENT Understanding the neural basis for phosphenes, the visual percepts created by electrical stimulation of visual cortex, is fundamental to the development of a visual cortical prosthetic. Our experiments in human subjects implanted with electrodes over visual cortex show that it is the activity of a large population of cells spread out across several millimeters of tissue that supports the perception of a phosphene. In addition, we describe an important feature of the production of phosphenes by electrical stimulation: phosphene size saturates at a relatively low current level. This finding implies that, with current methods, visual prosthetics will have a limited dynamic range available to control the production of spatial forms and that more advanced stimulation methods may be required.
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The present results support previous findings and show that the border between V1 and V2, located near the occipital pole, is a suitable place for a cortical visual prosthesis.
“Our present results support previous findings (5, 10, 38, 39, 41, 42) and show that the border between V1 and V2, which is located near the occipital pole, is a suitable place for a cortical visual prosthesis (37).”
From Visual percepts evoked with an intracortical 96-channel microelectrode array inserted in human occipital cortex (2021) · cites this paper as [39]
The Electrodes section (page unavailable) places V1 and V2 near the occipital pole, and the prosthetic-devices discussion describes stimulation of early visual areas as an approach to cortical visual prostheses. The paper does not identify the V1–V2 border specifically or establish that this border is a suitable prosthesis site. (LLM, conf=0.84)
Experimental — accuracy not yet measuredThere often exists a numeric or symbolic forward model, constrained by empirical data, that can predict a neuronal or ideally perceptual response to the applied stimulus.
“In addition, there often exists a numeric or symbolic forward model, constrained by empirical data, that can predict a neuronal or ideally perceptual response to the applied stimulus (Bosking et al., 2017; Beyeler et al., 2019).”
From Towards a <i>Smart Bionic Eye</i> : AI-powered artificial vision for the treatment of incurable blindness (2022) · cites this paper as Bosking et al., 2017
The effects elicited by iES vary with the frequency and amplitude of stimulation, with higher parameter values tending to elicit more potent responses across many domains.
“This is critical because the effects elicited by iES are known to vary with the frequency and amplitude of stimulation (Mohan et al., 2020), with higher parameter values tending to elicit more potent responses across many domains, including visual (Winawer and Parvizi, 2016;Bosking et al., 2017), physiological (Inman et al., 2018), and emotional effects (Yih et al., 2019).”
From Does the Prefrontal Cortex Play an Essential Role in Consciousness? Insights from Intracranial Electrical Stimulation of the Human Brain (2021) · cites this paper as Bosking et al., 2017
Previous studies in monkeys and humans demonstrated that the thresholds for phosphene perception in areas V1 and V2 are similar and that the quality of the percepts elicited is comparable.
“Previous studies in monkeys and humans demonstrated that the thresholds for phosphene perception in areas V1 and V2 are similar and that the quality of the percepts that are elicited are comparable (14, 3840).”
From Visual percepts evoked with an intracortical 96-channel microelectrode array inserted in human occipital cortex (2021) · cites this paper as [39]
The Summary describes human phosphene-size measurements from V1 stimulation, and the Electrodes section mentions V2 only as part of the surrounding early visual cortex (pages unavailable). Searches for V1–V2 perception thresholds and percept quality found no comparison supporting either part of the claim. (LLM, conf=0.75)
Experimental — accuracy not yet measuredThe model predictions are similar to those made by others using simpler models that assume phosphene size is inversely proportional to cortical magnification.
“our model predictions are similar to those made by others24,83 using simpler models that assume that phosphene size is inversely proportional to cortical magnification.”
From A virtual patient simulation modeling the neural and perceptual effects of human visual cortical stimulation, from pulse trains to percepts (2024) · cites this paper as [24]
The previous observation of Bosking et al. that the size of the phosphenes drawn by their patients could be predicted by cortical magnification is explained by the relationship between receptive field areas and cortical magnification.
“Across much of cortex receptive field areas approximate the areal cortical magnification to the − 2/3 power82; explaining the previous observation of Bosking et al.24 that the size of the phosphenes drawn by their patients could be predicted by cortical magnification.”
From A virtual patient simulation modeling the neural and perceptual effects of human visual cortical stimulation, from pulse trains to percepts (2024) · cites this paper as [24]
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“Our central finding, that current magnitude correlates with subjective intensity, parallels our previous observations of a clear relationship in the primary visual cortex between the magnitude of electrical stimulation, the size of cortical area being activated and the size of perceived visual phosphenes (Winawer and…”
ResultOswalt, D. N., Bosking, W. H., Sun, P., Sheth, S. A., Niketeghad, S., Salas, M. A., Patel, U. K., Greenberg, R. J., Dorn, J. D., Pouratian, N., et al. (2021) · Brain stimulation · doi:10.1016/j.brs.2021.08.024 · Cited by 40 · see citing works
“A standard mapping equation was used to estimate th e xpected relationship between CMF and 334 eccentricity for each subject (see solid lines in F igure 8) [15], [17], [31].”
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“…studies in non-human primates have explored how different stimulation parameters and electrode configurations can be used to alter phosphene characteristics such as shape, brightness, longevity and color (Bak et al 1990, Tehovnik et al 2005, Torab et al 2011, Davis et al 2012, Bosking et al 2017).”
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“In parallel, also the ICMS-evoked phosphene size depend on the location of the stimulated site in the visual cortex, with size increasing with distance from the foveal representation (Bosking et al., 2017).”
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“Our model successfully predicts the finding that phosphene size increases as a function of eccentricity in the visual field.”
BackgroundMethodInfluentialIntracortical current steering shifts the location of evoked neural activity
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“Based on the correlations between single-electrode neural responses and the evoked phosphene location and size [15, 16, 34], predicting the current steering response will enable clinicians to determine the parameters necessary for evoking a phosphene in a desired location of the visual field.”
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“Additionally, stimulating with an increased current will only increase the size of a phosphene until saturation in phosphene size is reached (Bosking et al. 2017).”
BackgroundMethodInfluentialRules governing perception of multiple phosphenes by human observers
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Postmortem investigation of a human cortical visual prosthesis that was implanted for 36 years
Towle, V. L., Pytel, P., Lane, F., Plass, J., Frim, D. M., & Troyk, P. R. (2020) · Journal of Neural Engineering · doi:10.1088/1741-2552/ab9d11 · Cited by 11 · see citing works
Continuity within the somatosensory cortical map facilitates learning
Lassagne, H., Goueytes, D., Shulz, D. E., Estebanez, L., & Ego‐Stengel, V. (2022) · Cell Reports · doi:10.1016/j.celrep.2022.110617 · Cited by 12 · see citing works
“Stimulation of distinct cortical zones in a topographic sensory map elicited localized percepts matching the expected peripheral locations on the sensory organ (Bosking et al., 2017; Ceballo et al., 2019a; Chen et al., 2020; Flesher et al., 2016; Winawer and Parvizi, 2016).”
BackgroundFour-dimensional map of the human early visual system
Nakai, Y., Nagashima, A., Hayakawa, A., Osuki, T., Jeong, J., Sugiura, A., Brown, E. C., & Asano, E. (2017) · Clinical Neurophysiology · doi:10.1016/j.clinph.2017.10.019 · Cited by 12 · see citing works
Comparison of electrical microstimulation artifact removal methods for high-channel-count prostheses
Wang, F., Chen, X., & Roelfsema, P. R. (2024) · Journal of Neuroscience Methods · doi:10.1016/j.jneumeth.2024.110169 · Cited by 11 · see citing works
“…the eye and induce visual percepts that resemble dots of light via electrical stimulation, known as ‘phosphenes ’ (Bak, 1990; Bartlett et al., 2005; Bosking et al., 2017; Brindley and Lewin, 1968; Button and Put-nam, 1962; Dagnino et al., 2015; Davis et al., 2012; DeYoe et al., 2005; Dobelle et…”
Grani, F., Soto‐Sánchez, C., Doblado, A. R., López-Peco, R., González-López, P., & Fernández, E. (2025) · Science Advances · doi:10.1126/sciadv.adv8846 · Cited by 10 · see citing works
Raising the stakes for cortical visual prostheses
Beauchamp, M. S., Bosking, W. H., Oswalt, D. N., & Yoshor, D. (2021) · Journal of Clinical Investigation · doi:10.1172/jci154983 · Cited by 9 · see citing works
Self-citationMuramatsu, S., Toda, M., Nishikawa, J., & Tateno, T. (2019) · Brain Research · doi:10.1016/j.brainres.2019.146312 · Cited by 9 · see citing works
Biased Orientation and Color Tuning of the Human Visual Gamma Rhythm
Li, Y., Bosking, W. H., Beauchamp, M. S., Sheth, S. A., Yoshor, D., Bartoli, E., & Foster, B. L. (2021) · Journal of Neuroscience · doi:10.1523/jneurosci.1085-21.2021 · Cited by 8 · see citing works
Self-citationBuilding egocentric models of local space from retinal input
Martins, D. M., Manda, J. M., Goard, M. J., & Parker, P. R. L. (2024) · Current Biology · doi:10.1016/j.cub.2024.10.057 · Cited by 7 · see citing works
Improving Understanding of Visual Snow by Quantifying its Appearance and Effect on Vision
Brooks, C. J., Chan, Y. M., Fielding, J., White, O. B., Badcock, D. R., & McKendrick, A. M. (2024) · Investigative Ophthalmology & Visual Science · doi:10.1167/iovs.65.5.38 · Cited by 6 · see citing works
“41 The retinal image of the single-pixel simulation dot is expected to be larger in size than a foveal cone but approaching the midget ganglion cell receptive field.”
BackgroundBayesian optimization of cortical neuroprosthetic vision using perceptual feedback
Küçükoğlu, B., Soo, L., Leeftink, D., Grani, F., Soto‐Sánchez, C., Güçlü, U., van Gerven, M. A. J., & Fernández, E. (2025) · Journal of Neural Engineering · doi:10.1088/1741-2552/adeae9 · Cited by 5 · see citing works
“For example, as stimulation currents are raised above the threshold, the size of phosphenes is found to reach a rapid saturation after an initial increase (Bosking et al 2017).”
Towards precise synthetic neural codes: high-dimensional stimulation with flexible electrodes
Kim, R., Liu, Y., Zhang, J., Xie, C., & Luan, L. (2025) · npj Flexible Electronics · doi:10.1038/s41528-025-00447-y · Cited by 4 · see citing works
Esquenazi, R., Meier, K., Beyeler, M., Boynton, G. M., & Fine, I. (2021) · Journal of Vision · doi:10.1167/jov.21.13.10 · Cited by 4 · see citing works
“Other groups are actively implanting (Beauchamp et al., 2020; Bosking, Beauchamp, & Yoshor, 2017; Bosking, Sun, et al., 2017; Morillas et al., 2007; Murphey, Maunsell, Beauchamp, & Yoshor, 2009) or developing implants for cortical stimulation (Chen, Wang, Fernandez, & Roelfsema, 2020; Troyk, 2017).”
Pavloski, R. P. (2022) · Prosthesis · doi:10.3390/prosthesis4040049 · Cited by 4 · see citing works
“The techniques reported in [30,32] appear to be very promising candidates for making the required estimations.”
BackgroundOptimization of sputtered iridium oxide microelectrodes for long-term intracortical stimulation
Schelles, M., Goyvaerts, L., Merken, L., Janssen, P., Ceyssens, F., & Kraft, M. D. (2025) · Biomedical Physics & Engineering Express · doi:10.1088/2057-1976/adfdec · Cited by 3 · see citing works
Lozano, A., Chen, X., Grouw, M. L., Li, B., Wang, F., van der Grinten, M., Soto‐Sánchez, C., Morales-Gregorio, A., Fernández, E., & Roelfsema, P. R. (2025) · Brain stimulation · doi:10.1016/j.brs.2025.103019 · Cited by 6 · see citing works
A novel simulation paradigm utilising MRI-derived phosphene maps for cortical prosthetic vision
Wang, H. Z., & Wong, Y. T. (2023) · Journal of Neural Engineering · doi:10.1088/1741-2552/aceca2 · Cited by 3 · see citing works
“All parameters were taken from the final model in [52], which provides the best prediction result for all cases.”
BackgroundMethodInfluentialLi, Y., Tan, Z., Wang, J., Wang, M., & Wang, L. (2022) · Frontiers in Neuroscience · doi:10.3389/fnins.2022.918767 · Cited by 3 · see citing works
Sadrzadeh-Afsharazar, F., & Douplik, A. (2024) · Sensors · doi:10.3390/s24082512 · Cited by 3 · see citing works
Sense Element Engagement Theory Explains How Neural Networks Produce Cortical Prosthetic VisionPreprint
Pavloski, R. P. (2021) · doi:10.36227/techrxiv.17161187 · Cited by 3 · see citing works
Large-scale RF mapping without visual input for neuroprostheses in macaque and human visual cortexPreprint
Lozano, A., Chen, X., Grouw, M. L., Li, B., Wang, F., van der Grinten, M., Soto‐Sánchez, C., Morales-Gregorio, A., Fernández, E., & Roelfsema, P. R. (2024) · medRxiv · doi:10.1101/2024.12.22.24319047 · Cited by 3 · see citing works
Perceptual learning of prosthetic vision using video game training
Esquenazi, R., Meier, K., Beyeler, M., Wright, D., Boynton, G. M., & Fine, I. (2025) · Journal of Vision · doi:10.1167/jov.25.12.12 · Cited by 2 · see citing works
“…Le Mer, Mohand-Said, & Sahel, 2022; Rizzo et al., 2014; Saunders et al., 2014; Stingl et al., 2015), cortical implants (Beauchamp et al., 2020; Bosking et al., 2017; Fernandez et al., 2021; Troyk, 2017), and optogenetic technologies (AbbVie, 2022; Antolik, Sabatier, Galle, Fregnac, &…”
Electrical stimulation thresholds differ between V1 and V2
Meikle, S. J., Allison-Walker, T., Hagan, M. A., Price, N. S. C., & Wong, Y. T. (2023) · Annual International Conference of the IEEE Engineering in Medicine and Biology Society (EMBC) · doi:10.1109/embc40787.2023.10340103 · Cited by 2 · see citing works
“However, the threshold level for V2 is still much lower than subdural stimulation, which has been used in humans with no reported side effects [18].”
BackgroundKaas, J. H., & Stepniewska, I. (2021) · Contemporary clinical neuroscience · doi:10.1007/978-3-030-54564-2_1 · Cited by 2 · see citing works
Spatial resolution of local field potential signals in macaque V4
Foroushani, A. N., Neupane, S., Pastor, P. D. H., Pack, C. C., & Sawan, M. (2020) · Journal of Neural Engineering · doi:10.1088/1741-2552/ab7321 · Cited by 2 · see citing works
Bipolar intracortical microstimulation expands the repertoire of neuronal activation patterns
Schelles, M., Verhaege, A., Rompaey, N. V., Goyvaerts, L., Wierda, K. D. B., Ceyssens, F., Kraft, M. D., & Bonin, V. (2026) · Journal of Neural Engineering · doi:10.1101/2025.03.21.644593 · Cited by 2 · see citing works
“Higher current amplitudes recruit larger neural populations, resulting in larger and less precise phosphenes [8,35], highlighting the need for more selective stimulation.”
Electrophysiological Signatures of Visual Sensations Elicited by Direct Electrical Stimulation
Li, Y., Zhang, B., Wang, J., Saalmann, Y. B., Afrasiabi, M., Lv, P.-C., Wang, H.-X., Xiang, H.-H., Wang, M., Luan, G., et al. (2025) · Neuroscience Bulletin · doi:10.1007/s12264-025-01464-7 · Cited by 1 · see citing works
Kozai, T. D. Y., Hooks, B. M., Vazquez, A. L., Gharbawie, O. A., Huang, C., Graczyk, E. L., Smith, M. A., Taylor, D. M., Hernandez-Reynoso, A. G., London, A. J., et al. (2026) · Journal of Neural Engineering · doi:10.1088/1741-2552/ae54cf · Cited by 2 · see citing works
Electrical stimulation mapping of the visual cortex for epilepsy surgery
Jonas, J., Aupy, J., PETRESCU, A. M., Marchive, M., Perguilhem, S., Guinet, V., Laguitton, V., Sabadell, V., Chipaux, M., & Dangles, M.-T. (2026) · Neurophysiologie Clinique · doi:10.1016/j.neucli.2026.103145 · Cited by 1 · see citing works
Restoring vision and touch with cortical microstimulation
Valle, G., Oswalt, D. N., Gaunt, R. A., Roelfsema, P. R., Greenspon, C. M., & Fernandez, E. (2026) · Nature Reviews Bioengineering · doi:10.1038/s44222-026-00449-z · Cited by 1 · see citing works
“As visual and somatosensory cortices exhibit retinotopic and somatotopic maps, respectively 16,60 , localized activation of neurons results in punctate percepts appropriate to the stimulated region 61–63 .”
InfluentialOswalt, D. N., Datta, P., Talbot, N., Mirzadeh, Z., & Greger, B. E. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.12.15.422891 · Cited by 1 · see citing works
“[36] W.”
BackgroundSense Element Engagement Theory Explains How Neural Networks Produce Cortical Prosthetic VisionPreprint
Pavloski, R. P. (2021) · doi:10.36227/techrxiv.17161187.v1 · Cited by 1 · see citing works
Fine, I., & Boynton, G. M. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.03.18.532424 · Cited by 1 · see citing works
“Across much of cortex receptive field areas approximate the areal cortical magnification to the −2/3 power(55); explaining the previous observation of Bosking et al.(20) that the size of the phosphenes drawn by their patients could be predicted by cortical magnification.”
BackgroundMethodResultInfluentialContinuity within somatosensory cortical map shapes the integration of optogenetic inputPreprint
Lassagne, H., Goueytes, D., Shulz, D. E., Estebanez, L., & Ego‐Stengel, V. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.03.26.437211
Restoring Vision Naturally and Noninvasively
Wang, C., Deng, H., & Kuang, S. (2021) · Neuroscience Bulletin · doi:10.1007/s12264-021-00760-2
Yih, J., Beam, D. E., Fox, K. C. R., & Parvizi, J. (2019) · doi:10.31231/osf.io/8jnxm
“…observations of a clear relationship in the primary visual cortex between the magnitude of electrical stimulation, the size of cortical area being activated, and the size of perceived visual phosphenes (Winawer & Parvizi, 2016) – a finding that was recently replicated (Bosking et al., 2017).”
BackgroundSadrzadeh-Afsharazar, F., & Douplik, A. (2023) · Research Square · doi:10.21203/rs.3.rs-3213860/v1
Optimal Frequency for Biomedical Wireless Power TransferPreprint
van Nunen, T., Mestrom, R., & Visser, H. J. (2023) · Research Square · doi:10.21203/rs.3.rs-3527426/v1
Wang, H. Z., & Wong, Y. T. (2023) · Annual International Conference of the IEEE Engineering in Medicine and Biology Society (EMBC) · doi:10.1109/embc40787.2023.10341189
“Studies using subdural electrodes have developed a mathematical model that can predict phosphene size based on the stimulation current level and eccentricity of the electrode [15].”
BackgroundMethodInfluentialOptimizing cortical feedback strategies for a closed-loop brain machine interface
Lassagne, H. (2023) · theses.fr (ABES) · doi:10.70675/779d5bbaz5470z4faczb771z288a715634b1
- Authors
- Bosking, W. H., Sun, P., Ozker, M., Pei, X., Foster, B. L., Beauchamp, M. S., & Yoshor, D.
- Affiliations
- Baylor College of Medicine (US)
- Venue
- The Journal of Neuroscience
- Publisher
- Society for Neuroscience
- Published
- 2017-06-26
- Volume
- 37
- Issue
- 30
- Pages
- 7188-7197
- ISSN
- 0270-6474 (ISSN-L)
- 1529-2401
- PMC
- PMC5546398
- OpenAlex
- W2674339973
- Cited by
- 75
- References
- 42
- Citations by year
- 2026: 4 · 2025: 8 · 2024: 7 · 2023: 10 · 2022: 9 · 2021: 16 · 2020: 8 · 2019: 8 · 2018: 3 · 2017: 2
- FWCI
- 2.81
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Journal of Neuroscience (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- Funders
- National Institutes of Health
- National Institute of Mental Health (R00MH103479)
- National Eye Institute (R01EY023336)
- Topics
- Life Sciences › Neuroscience › Cellular and Molecular Neuroscience › Neuroscience and Neural Engineering
- Life Sciences › Neuroscience › Cognitive Neuroscience › Tactile and Sensory Interactions
- Life Sciences › Neuroscience › Cognitive Neuroscience › EEG and Brain-Computer Interfaces
- Keywords
- phosphenes, visual cortex stimulation, retinotopic map, cortical magnification factor, visual prosthetics, foveal representation, dynamic range
- MeSH terms
- Adult, Electric Stimulation, Evoked Potentials, Visual, Female, Humans, Male, Middle Aged, Nerve Net, Phosphenes, Visual Cortex, Visual Fields
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