Abstract
Neurons in the superior colliculus (SC), like those in cortex, are modulated by shifts in attention but contribute differently from visual cortex neurons. It remains debated whether SC attention-related activity represents enhanced perceptual sensitivity, decision criterion shifts, or motor bias linked to saccade planning at neurons’ response fields. We dissociated these components by independently controlling behavioral sensitivity, perceptual decision criterion, and motor response criterion in a visual spatial attention task in two male rhesus monkeys. SC activity correlated strongly with behavioral sensitivity and motor criterion for selecting a saccade target within the response field, independent of saccade execution. In contrast, SC responses remained unchanged despite large shifts in decision criterion when motor bias was excluded. Notably, SC activity did not predict trial-by-trial choice accuracy. These results demonstrate that the SC specifically supports components of attention related to heightened perceptual sensitivity and response selection, while being largely independent of internal perceptual decision criterion and trial outcome.
Similar content being viewed by others
Data availability
All data are available in the main text, source data or the supplementary information. Source data are provided with this paper.
Code availability
All behavioral and neuronal data analysis was done using Matlab (MathWorks Inc.). Behavioral task was controlled using custom-written software (https://github.com/MaunsellLab/Lablib-Public-05-February-2026).
References
Luo, T. Z. & Maunsell, J. H. Attention can be subdivided into neurobiological components corresponding to distinct behavioral effects. Proc. Natl. Acad. Sci. 116, 26187–26194 (2019).
Knudsen, E. I. Fundamental components of attention. Annu. Rev. Neurosci. 30, 57–78 (2007).
MacDonald, A. W., Cohen, J. D., Stenger, V. A. & Carter, C. S. Dissociating the role of the dorsolateral prefrontal and anterior cingulate cortex in cognitive control. Science 288, 1835–1838 (2000).
Moran, J. & Desimone, R. Selective attention gates visual processing in the extrastriate cortex. Science 229, 782–784 (1985).
Krauzlis, R. J., Lovejoy, L. P. & Zénon, A. Superior colliculus and visual spatial attention. Annu. Rev. Neurosci. 36, 165–182 (2013).
Goldberg, M. E. & Wurtz, R. H. Activity of superior colliculus in behaving monkey. II. Effect of attention on neuronal responses. J. Neurophysiol. 35, 560–574 (1972).
Macmillan, N. A., and Creelman, C. D. (2004). Detection theory: A user’s guide (Psychology press).
Luo, T. Z. & Maunsell, J. H. R. Neuronal modulations in visual cortex are associated with only one of multiple components of attention. Neuron 86, 1182–1188 (2015).
Ghosh, S. & Maunsell, J. H. R. Neuronal correlates of selective attention and effort in visual area V4 are invariant of motivational context. Sci. Adv. 8, eabc8812 (2022).
Lovejoy, L. P. & Krauzlis, R. J. Inactivation of primate superior colliculus impairs covert selection of signals for perceptual judgments. Nat. Neurosci. 13, 261–266 (2010).
Jun, E. J. et al. Causal role for the primate superior colliculus in the computation of evidence for perceptual decisions. Nat. Neurosci. 24, 1121–1131 (2021).
Sridharan, D., Steinmetz, N. A., Moore, T. & Knudsen, E. I. Does the superior colliculus control perceptual sensitivity or choice bias during attention? Evidence from a multialternative decision framework. J. Neurosci. 37, 480–511 (2017).
Zénon, A. & Krauzlis, R. J. Attention deficits without cortical neuronal deficits. Nature 489, 434–437 (2012).
Basso, M. A. & May, P. J. Circuits for action and cognition: a view from the superior colliculus. Annu. Rev. Vis. Sci. 3, 197–226 (2017).
Ignashchenkova, A., Dicke, P. W., Haarmeier, T. & Thier, P. Neuron-specific contribution of the superior colliculus to overt and covert shifts of attention. Nat. Neurosci. 7, 56–64 (2004).
Crapse, T. B., Lau, H. & Basso, M. A. A role for the superior colliculus in decision criteria. Neuron 97, 181–194.e186 (2018).
McPeek, R. M. & Keller, E. L. Saccade target selection in the superior colliculus during a visual search task. J. Neurophysiol. 88, 2019–2034 (2002).
Ghosh, S. & Maunsell, J. H. R. Locus coeruleus norepinephrine contributes to visual-spatial attention by selectively enhancing perceptual sensitivity. Neuron 112, 2231–2240.e2235 (2024).
Peysakhovich, B. et al. Primate superior colliculus is causally engaged in abstract higher-order cognition. Nat. Neurosci. 27, 1999–2008 (2024).
Luo, T. Z. & Maunsell, J. H. R. Attentional changes in either criterion or sensitivity are associated with robust modulations in lateral prefrontal cortex. Neuron 97, 1382–1393.e1387 (2018).
Baruchin, L. J., Alleman, M. & Schröder, S. Reward modulates visual responses in the superficial superior colliculus of mice. J. Neurosci. 43, 8663–8680 (2023).
McPeek, R. M. & Keller, E. L. Deficits in saccade target selection after inactivation of superior colliculus. Nat. Neurosci. 7, 757–763 (2004).
Stine, G. M., Trautmann, E. M., Jeurissen, D. & Shadlen, M. N. A neural mechanism for terminating decisions. Neuron 111, 2601–2613.e2605 (2023).
Katz, L. N. et al. Optogenetic Manipulation of Covert Attention in the Nonhuman Primate. J. Cogn. Neurosci. 37, 266–285 (2025).
Carrasco, M. Visual attention: The past 25 years. Vis. Res. 51, 1484–1525 (2011).
Freedman, D. J. & Assad, J. A. Experience-dependent representation of visual categories in parietal cortex. Nature 443, 85–88 (2006).
Bisley, J. W. & Mirpour, K. The neural instantiation of a priority map. Curr. Opin. Psychol. 29, 108–112 (2019).
Bollimunta, A., Bogadhi, A. R. & Krauzlis, R. J. Comparing frontal eye field and superior colliculus contributions to covert spatial attention. Nat. Commun. 9, 3553 (2018).
Katz, L. N., Yu, G., Herman, J. P. & Krauzlis, R. J. Correlated variability in primate superior colliculus depends on functional class. Commun. Biol. 6, 540 (2023).
Sommer, M. A. & Wurtz, R. H. What the Brain Stem Tells the Frontal Cortex. I. Oculomotor Signals Sent From Superior Colliculus to Frontal Eye Field Via Mediodorsal Thalamus. J. Neurophysiol. 91, 1381–1402 (2004).
Ayar, E. C., Heusser, M. R., Bourrelly, C. & Gandhi, N. J. Distinct context- and content-dependent population codes in superior colliculus during sensation and action. Proc. Natl. Acad. Sci. 120, e2303523120 (2023).
Ghosh, S. & Maunsell, J. H. R. Single trial neuronal activity dynamics of attentional intensity in monkey visual area V4. Nat. Commun. 12, 1–15 (2021).
Maunsell, J. H. Neuronal representations of cognitive state: reward or attention? Trends Cogn. Sci. 8, 261–265 (2004).
Nilsson, N., Håkansson, B. & Ortiz-Catalan, M. Classification complexity in myoelectric pattern recognition. J. Neuroeng.Rehabilitation 14, 1–18 (2017).
Kobak, D. et al. Demixed principal component analysis of neural population data. Elife 5, e10989 (2016).
Acknowledgements
We thank Chery J. Cherian and Lai Wei for critical feedback on the manuscript; Rachel Parker for technical help with monkey procedures. This work was supported by National Institutes of Health grant R01EY005911 (J.H.R.M.) and Brain and Behavior Research Foundation grant NARSAD 28812 (S.G.). The funder had no role in study design, data collection and interpretation, or the decision to submit the work for publication.
Author information
Authors and Affiliations
Contributions
S.G. and J.H.R.M. designed the experiments, performed the surgeries, and wrote the paper. S.G. performed the experiments and analyzed the data.
Corresponding authors
Ethics declarations
Competing interests
The authors declare no competing interests.
Peer review
Peer review information
Nature Communications thanks Xiaomo Chen and James Herman for their contribution to the peer review of this work. A peer review file is available.
Additional information
Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.
Source data
Rights and permissions
Open Access This article is licensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License, which permits any non-commercial use, sharing, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if you modified the licensed material. You do not have permission under this licence to share adapted material derived from this article or parts of it. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by-nc-nd/4.0/.
About this article
Cite this article
Ghosh, S., Maunsell, J.H.R. Attention-related modulation in the superior colliculus encodes perceptual sensitivity, but not perceptual choice. Nat Commun (2026). https://doi.org/10.1038/s41467-026-69954-4
Received:
Accepted:
Published:
DOI: https://doi.org/10.1038/s41467-026-69954-4


