{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T10:27:48Z","timestamp":1787567268137,"version":"build-2736575974"},"reference-count":110,"publisher":"eLife Sciences Publications, Ltd","license":[{"start":{"date-parts":[[2026,6,8]],"date-time":"2026-06-08T00:00:00Z","timestamp":1780876800000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,6,8]],"date-time":"2026-06-08T00:00:00Z","timestamp":1780876800000},"content-version":"am","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"},{"start":{"date-parts":[[2026,6,8]],"date-time":"2026-06-08T00:00:00Z","timestamp":1780876800000},"content-version":"tdm","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["R01EY029250"],"award-info":[{"award-number":["R01EY029250"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"award":["R01EY029250"],"award-info":[{"award-number":["R01EY029250"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]},{"DOI":"10.13039\/100000002","name":"National Institutes of Health","doi-asserted-by":"crossref","award":["R01MH128393"],"award-info":[{"award-number":["R01MH128393"]}],"id":[{"id":"10.13039\/100000002","id-type":"DOI","asserted-by":"crossref"}]},{"award":["R01MH128393"],"award-info":[{"award-number":["R01MH128393"]}],"id":[{"id":"https:\/\/ror.org\/01cwqze88","id-type":"ROR","asserted-by":"publisher"}]}],"content-domain":{"domain":["elifesciences.org"],"crossmark-restriction":false},"short-container-title":[],"abstract":"<jats:p>The superior colliculus (SC) is traditionally considered a brain region that functions as an interface between processing visual inputs and generating eye movement outputs. Although its role as a primary reflex center is thought to be conserved across vertebrate species, evidence suggests that the SC has evolved to support higher-order cognitive functions, including spatial attention. When it comes to oculomotor areas, such as the SC, it is critical that high precision fixation and eye movements are maintained even in the presence of signals related to ongoing changes in cognition and brain state, both of which have the potential to interfere with eye position encoding and movement generation. In this study, we recorded spiking responses of neuronal populations in the SC while two rhesus macaque monkeys performed a memory-guided saccade task and found that the activity of some of the neurons fluctuated over tens of minutes. By leveraging the statistical power afforded by high-dimensional neuronal recordings, we were able to identify a low-dimensional pattern of activity that was correlated with pupil size and simultaneously recorded data in the prefrontal cortex (PFC), consistent with slow changes in the monkeys\u2019 arousal levels while they were performing the task. Importantly, we found that the spiking responses of deep-layer SC neurons were less correlated with this brain-wide arousal signal, and that neural activity associated with changes in pupil size and saccade tuning did not overlap in population activity space with movement initiation signals. Taken together, these findings provide a framework for understanding how signals related to cognition and arousal can be embedded in the population activity of oculomotor structures without compromising the fidelity of the motor output.<\/jats:p>","DOI":"10.7554\/elife.99278","type":"journal-article","created":{"date-parts":[[2024,8,21]],"date-time":"2024-08-21T19:27:30Z","timestamp":1724268450000},"update-policy":"https:\/\/doi.org\/10.7554\/elife.99278","source":"Crossref","is-referenced-by-count":2,"title":["Brain-wide arousal signals are segregated from movement planning in the superior colliculus of the macaque"],"prefix":"10.7554","volume":"13","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-1646-6919","authenticated-orcid":true,"given":"Richard","family":"Johnston","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05x2bcf33","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Biomedical Engineering, Carnegie Mellon University","place":["Pittsburgh, United States"]},{"id":[{"id":"https:\/\/ror.org\/05x2bcf33","id-type":"ROR","asserted-by":"publisher"}],"name":"Neuroscience Institute, Carnegie Mellon University","place":["Pittsburgh, United States"]},{"id":[{"id":"https:\/\/ror.org\/05x2bcf33","id-type":"ROR","asserted-by":"publisher"}],"name":"Center for the Neural Basis of Cognition, Carnegie Mellon University","place":["Pittsburgh, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"data-curation","vocabulary":"credit"},{"role":"formal-analysis","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"methodology","vocabulary":"credit"},{"role":"resources","vocabulary":"credit"},{"role":"software","vocabulary":"credit"},{"role":"validation","vocabulary":"credit"},{"role":"visualization","vocabulary":"credit"},{"role":"writing-original-draft","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1192-9942","authenticated-orcid":true,"given":"Matthew A","family":"Smith","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/05x2bcf33","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Biomedical Engineering, Carnegie Mellon University","place":["Pittsburgh, United States"]},{"id":[{"id":"https:\/\/ror.org\/05x2bcf33","id-type":"ROR","asserted-by":"publisher"}],"name":"Neuroscience Institute, Carnegie Mellon University","place":["Pittsburgh, United States"]},{"id":[{"id":"https:\/\/ror.org\/05x2bcf33","id-type":"ROR","asserted-by":"publisher"}],"name":"Center for the Neural Basis of Cognition, Carnegie Mellon University","place":["Pittsburgh, United States"]}],"role":[{"role":"author","vocabulary":"crossref"},{"role":"corresponding-author","vocabulary":"crossref"},{"role":"conceptualization","vocabulary":"credit"},{"role":"investigation","vocabulary":"credit"},{"role":"project-administration","vocabulary":"credit"},{"role":"supervision","vocabulary":"credit"},{"role":"writing-review-editing","vocabulary":"credit"}]}],"member":"4374","published-online":{"date-parts":[[2026,6,8]]},"reference":[{"key":"bib1","doi-asserted-by":"publisher","first-page":"119","DOI":"10.1016\/j.conb.2021.10.005","type":"journal-article","article-title":"Orienting our view of the superior colliculus: specializations and general functions","volume":"71","author":"Allen","year":"2021","journal-title":"Current Opinion in Neurobiology"},{"key":"bib2","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.46159","type":"journal-article","article-title":"Motor cortex signals for each arm are mixed across hemispheres and neurons yet partitioned within the population response","volume":"8","author":"Ames","year":"2019","journal-title":"eLife"},{"key":"bib3","doi-asserted-by":"publisher","first-page":"1410","DOI":"10.1038\/s41593-020-0696-5","type":"journal-article","article-title":"Prefrontal cortex exhibits multidimensional dynamic encoding during decision-making","volume":"23","author":"Aoi","year":"2020","journal-title":"Nature Neuroscience"},{"key":"bib4","doi-asserted-by":"publisher","first-page":"403","DOI":"10.1146\/annurev.neuro.28.061604.135709","type":"journal-article","article-title":"An integrative theory of locus coeruleus-norepinephrine function: adaptive gain and optimal performance","volume":"28","author":"Aston-Jones","year":"2005","journal-title":"Annual Review of Neuroscience"},{"key":"bib5","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2303523120","type":"journal-article","article-title":"Distinct context- and content-dependent population codes in superior colliculus during sensation and action","volume":"120","author":"Ayar","year":"2023","journal-title":"PNAS"},{"key":"bib6","doi-asserted-by":"publisher","first-page":"197","DOI":"10.1146\/annurev-vision-102016-061234","type":"journal-article","article-title":"Circuits for action and cognition: a view from the superior colliculus","volume":"3","author":"Basso","year":"2017","journal-title":"Annual Review of Vision Science"},{"key":"bib7","doi-asserted-by":"publisher","first-page":"918","DOI":"10.1016\/j.neuron.2021.01.013","type":"journal-article","article-title":"Unraveling circuits of visual perception and cognition through the superior colliculus","volume":"109","author":"Basso","year":"2021","journal-title":"Neuron"},{"key":"bib8","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2305759120","type":"journal-article","article-title":"Sensory tuning in neuronal movement commands","volume":"120","author":"Baumann","year":"2023","journal-title":"PNAS"},{"key":"bib9","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-021-24241-2","type":"journal-article","article-title":"Organization of the inputs and outputs of the mouse superior colliculus","volume":"12","author":"Benavidez","year":"2021","journal-title":"Nature Communications"},{"key":"bib10","doi-asserted-by":"publisher","first-page":"2000","DOI":"10.1002\/cne.24187","type":"journal-article","article-title":"A central mesencephalic reticular formation projection to medial rectus motoneurons supplying singly and multiply innervated extraocular muscle fibers","volume":"525","author":"Bohlen","year":"2017","journal-title":"The Journal of Comparative Neurology"},{"key":"bib11","doi-asserted-by":"publisher","DOI":"10.1167\/jov.21.11.16","type":"journal-article","article-title":"Fixational eye movements in passive versus active sustained fixation tasks","volume":"21","author":"Bowers","year":"2021","journal-title":"Journal of Vision"},{"key":"bib12","doi-asserted-by":"crossref","first-page":"433","DOI":"10.1163\/156856897X00357","type":"journal-article","article-title":"The psychophysics toolbox","volume":"10","author":"Brainard","year":"1997","journal-title":"Spatial Vision"},{"key":"bib13","doi-asserted-by":"publisher","first-page":"218","DOI":"10.1038\/s41593-018-0305-z","type":"journal-article","article-title":"Active control of arousal by a locus coeruleus GABAergic circuit","volume":"22","author":"Breton-Provencher","year":"2019","journal-title":"Nature Neuroscience"},{"key":"bib14","doi-asserted-by":"publisher","first-page":"239","DOI":"10.1146\/annurev-vision-091517-034142","type":"journal-article","article-title":"Visual function, organization, and development of the mouse superior colliculus","volume":"4","author":"Cang","year":"2018","journal-title":"Annual Review of Vision Science"},{"key":"bib15","doi-asserted-by":"publisher","first-page":"129","DOI":"10.1007\/s00221-006-0659-4","type":"journal-article","article-title":"A startle speeds up the execution of externally guided saccades","volume":"177","author":"Castellote","year":"2007","journal-title":"Experimental Brain Research"},{"key":"bib16","doi-asserted-by":"publisher","first-page":"121","DOI":"10.2147\/EB.S53613","type":"journal-article","article-title":"Distribution of cortical neurons projecting to the superior colliculus in macaque monkeys","volume":"2014","author":"Cerkevich","year":"2014","journal-title":"Eye and Brain"},{"key":"bib17","doi-asserted-by":"publisher","DOI":"10.3389\/fncir.2018.00058","type":"journal-article","article-title":"Orientation and contrast tuning properties and temporal flicker fusion characteristics of primate superior colliculus neurons","volume":"12","author":"Chen","year":"2018","journal-title":"Frontiers in Neural Circuits"},{"key":"bib18","doi-asserted-by":"publisher","first-page":"213","DOI":"10.1038\/s41583-024-00796-z","type":"journal-article","article-title":"Preparatory activity and the expansive null-space","volume":"25","author":"Churchland","year":"2024","journal-title":"Nature Reviews. Neuroscience"},{"key":"bib19","doi-asserted-by":"publisher","first-page":"59","DOI":"10.1016\/j.pneurobio.2015.06.006","type":"journal-article","article-title":"Visual attention: Linking prefrontal sources to neuronal and behavioral correlates","volume":"132","author":"Clark","year":"2015","journal-title":"Progress in Neurobiology"},{"key":"bib20","doi-asserted-by":"publisher","first-page":"811","DOI":"10.1038\/nn.2842","type":"journal-article","article-title":"Measuring and interpreting neuronal correlations","volume":"14","author":"Cohen","year":"2011","journal-title":"Nature Neuroscience"},{"key":"bib21","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.81826","type":"journal-article","article-title":"Cholinergic and noradrenergic axonal activity contains a behavioral-state signal that is coordinated across the dorsal cortex","volume":"12","author":"Collins","year":"2023","journal-title":"eLife"},{"key":"bib22","doi-asserted-by":"publisher","first-page":"4856","DOI":"10.1523\/JNEUROSCI.0242-23.2023","type":"journal-article","article-title":"Pupil dilation and microsaccades provide complementary insights into the dynamics of arousal and instantaneous attention during effortful listening","volume":"43","author":"Contadini-Wright","year":"2023","journal-title":"The Journal of Neuroscience"},{"key":"bib23","doi-asserted-by":"publisher","first-page":"551","DOI":"10.1016\/j.neuron.2020.07.021","type":"journal-article","article-title":"Slow drift of neural activity as a signature of impulsivity in macaque visual and prefrontal cortex","volume":"108","author":"Cowley","year":"2020","journal-title":"Neuron"},{"key":"bib24","doi-asserted-by":"publisher","first-page":"181","DOI":"10.1016\/j.neuron.2017.12.006","type":"journal-article","article-title":"A role for the superior colliculus in decision criteria","volume":"97","author":"Crapse","year":"2018","journal-title":"Neuron"},{"key":"bib25","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.23232","type":"journal-article","article-title":"Dynamic modulation of decision biases by brainstem arousal systems","volume":"6","author":"de Gee","year":"2017","journal-title":"eLife"},{"key":"bib26","doi-asserted-by":"publisher","first-page":"968","DOI":"10.1016\/j.neubiorev.2013.03.011","type":"journal-article","article-title":"Saccadic velocity as an arousal index in naturalistic tasks","volume":"37","author":"Di Stasi","year":"2013","journal-title":"Neuroscience and Biobehavioral Reviews"},{"key":"bib27","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.3000951","type":"journal-article","article-title":"Rules warp feature encoding in decision-making circuits","volume":"18","author":"Ebitz","year":"2020","journal-title":"PLOS Biology"},{"key":"bib28","doi-asserted-by":"publisher","first-page":"3055","DOI":"10.1016\/j.neuron.2021.07.011","type":"journal-article","article-title":"The population doctrine in cognitive neuroscience","volume":"109","author":"Ebitz","year":"2021","journal-title":"Neuron"},{"key":"bib29","doi-asserted-by":"publisher","first-page":"309","DOI":"10.1002\/cne.901840207","type":"journal-article","article-title":"Sources of subcortical projections to the superior colliculus in the cat","volume":"184","author":"Edwards","year":"1979","journal-title":"The Journal of Comparative Neurology"},{"key":"bib30","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms13239","type":"journal-article","article-title":"Reorganization between preparatory and movement population responses in motor cortex","volume":"7","author":"Elsayed","year":"2016","journal-title":"Nature Communications"},{"key":"bib31","doi-asserted-by":"publisher","first-page":"1035","DOI":"10.1016\/s0042-6989(03)00084-1","type":"journal-article","article-title":"Microsaccades uncover the orientation of covert attention","volume":"43","author":"Engbert","year":"2003","journal-title":"Vision Research"},{"key":"bib32","doi-asserted-by":"publisher","first-page":"205","DOI":"10.1146\/annurev-neuro-061010-113728","type":"journal-article","article-title":"Motor functions of the superior colliculus","volume":"34","author":"Gandhi","year":"2011","journal-title":"Annual Review of Neuroscience"},{"key":"bib33","doi-asserted-by":"publisher","first-page":"542","DOI":"10.1152\/jn.1972.35.4.542","type":"journal-article","article-title":"Activity of superior colliculus in behaving monkey I Visual receptive fields of single neurons","volume":"35","author":"Goldberg","year":"1972","journal-title":"Journal of Neurophysiology"},{"key":"bib34","doi-asserted-by":"publisher","first-page":"1904","DOI":"10.1152\/jn.01125.2011","type":"journal-article","article-title":"Similarity of superior colliculus involvement in microsaccade and saccade generation","volume":"107","author":"Hafed","year":"2012","journal-title":"Journal of Neurophysiology"},{"key":"bib35","doi-asserted-by":"publisher","DOI":"10.3389\/fncir.2021.638429","type":"journal-article","article-title":"Dissociable cortical and subcortical mechanisms for mediating the influences of visual cues on microsaccadic eye movements","volume":"15","author":"Hafed","year":"2021","journal-title":"Frontiers in Neural Circuits"},{"key":"bib36","doi-asserted-by":"publisher","first-page":"361","DOI":"10.1146\/annurev-vision-111022-123817","type":"journal-article","article-title":"Visual functions of the primate superior colliculus","volume":"9","author":"Hafed","year":"2023","journal-title":"Annual Review of Vision Science"},{"key":"bib37","doi-asserted-by":"publisher","first-page":"727","DOI":"10.1038\/s41593-021-00822-8","type":"journal-article","article-title":"Learning is shaped by abrupt changes in neural engagement","volume":"24","author":"Hennig","year":"2021","journal-title":"Nature Neuroscience"},{"key":"bib38","doi-asserted-by":"publisher","DOI":"10.1523\/ENEURO.0347-22.2022","type":"journal-article","article-title":"Decoding the time course of spatial information from spiking and local field potential activities in the superior colliculus","volume":"9","author":"Heusser","year":"2022","journal-title":"eNeuro"},{"key":"bib39","doi-asserted-by":"publisher","first-page":"1268","DOI":"10.1152\/jn.1983.49.5.1268","type":"journal-article","article-title":"Visual and oculomotor functions of monkey substantia nigra pars reticulata. III. Memory-contingent visual and saccade responses","volume":"49","author":"Hikosaka","year":"1983","journal-title":"Journal of Neurophysiology"},{"key":"bib40","doi-asserted-by":"publisher","first-page":"2281","DOI":"10.1152\/jn.00872.2003","type":"journal-article","article-title":"Representation of an abstract perceptual decision in macaque superior colliculus","volume":"91","author":"Horwitz","year":"2004","journal-title":"Journal of Neurophysiology"},{"key":"bib41","doi-asserted-by":"publisher","first-page":"299","DOI":"10.1002\/cne.902930211","type":"journal-article","article-title":"Supplementary eye field as defined by intracortical microstimulation: connections in macaques","volume":"293","author":"Huerta","year":"1990","journal-title":"The Journal of Comparative Neurology"},{"key":"bib42","doi-asserted-by":"publisher","first-page":"56","DOI":"10.1038\/nn1169","type":"journal-article","article-title":"Neuron-specific contribution of the superior colliculus to overt and covert shifts of attention","volume":"7","author":"Ignashchenkova","year":"2004","journal-title":"Nature Neuroscience"},{"key":"bib43","doi-asserted-by":"publisher","first-page":"R741","DOI":"10.1016\/j.cub.2021.04.001","type":"journal-article","article-title":"The tectum\/superior colliculus as the vertebrate solution for spatial sensory integration and action","volume":"31","author":"Isa","year":"2021","journal-title":"Current Biology"},{"key":"bib44","doi-asserted-by":"publisher","first-page":"1472","DOI":"10.1152\/jn.00641.2019","type":"journal-article","article-title":"A neural network for online spike classification that improves decoding accuracy","volume":"123","author":"Issar","year":"2020","journal-title":"Journal of Neurophysiology"},{"key":"bib45","doi-asserted-by":"publisher","first-page":"1010","DOI":"10.1016\/j.cub.2022.01.015","type":"journal-article","article-title":"Population temporal structure supplements the rate code during sensorimotor transformations","volume":"32","author":"Jagadisan","year":"2022","journal-title":"Current Biology"},{"key":"bib46","doi-asserted-by":"publisher","first-page":"3331","DOI":"10.1093\/cercor\/bhab418","type":"journal-article","article-title":"The eyes reflect an internal cognitive state hidden in the population activity of cortical neurons","volume":"32","author":"Johnston","year":"2022","journal-title":"Cerebral Cortex"},{"key":"bib47","doi-asserted-by":"publisher","DOI":"10.1523\/ENEURO.0012-22.2022","type":"journal-article","article-title":"EEG signals index a global signature of arousal embedded in neuronal population recordings","volume":"9","author":"Johnston","year":"2022","journal-title":"eNeuro"},{"key":"bib48","doi-asserted-by":"publisher","DOI":"10.1098\/rsta.2015.0202","type":"journal-article","article-title":"Principal component analysis: a review and recent developments","volume":"374","author":"Jolliffe","year":"2016","journal-title":"Philosophical Transactions. Series A, Mathematical, Physical, and Engineering Sciences"},{"key":"bib49","doi-asserted-by":"publisher","first-page":"221","DOI":"10.1016\/j.neuron.2015.11.028","type":"journal-article","article-title":"Relationships between pupil diameter and neuronal activity in the locus coeruleus, colliculi, and cingulate cortex","volume":"89","author":"Joshi","year":"2016","journal-title":"Neuron"},{"key":"bib50","doi-asserted-by":"publisher","first-page":"466","DOI":"10.1016\/j.tics.2020.03.005","type":"journal-article","article-title":"Pupil size as a window on neural substrates of cognition","volume":"24","author":"Joshi","year":"2020","journal-title":"Trends in Cognitive Sciences"},{"key":"bib51","doi-asserted-by":"publisher","DOI":"10.7554\/eLife.63490","type":"journal-article","article-title":"Context-dependent relationships between locus coeruleus firing patterns and coordinated neural activity in the anterior cingulate cortex","volume":"11","author":"Joshi","year":"2022","journal-title":"eLife"},{"key":"bib52","doi-asserted-by":"publisher","first-page":"1121","DOI":"10.1038\/s41593-021-00878-6","type":"journal-article","article-title":"Causal role for the primate superior colliculus in the computation of evidence for perceptual decisions","volume":"24","author":"Jun","year":"2021","journal-title":"Nature Neuroscience"},{"key":"bib53","doi-asserted-by":"publisher","first-page":"E1956","DOI":"10.1073\/pnas.1504866112","type":"journal-article","article-title":"Tectal microcircuit generating visual selection commands on gaze-controlling neurons","volume":"112","author":"Kardamakis","year":"2015","journal-title":"PNAS"},{"key":"bib54","doi-asserted-by":"publisher","first-page":"1250","DOI":"10.1152\/jn.00265.2011","type":"journal-article","article-title":"Order of operations for decoding superior colliculus activity for saccade generation","volume":"106","author":"Katnani","year":"2011","journal-title":"Journal of Neurophysiology"},{"key":"bib55","doi-asserted-by":"publisher","first-page":"3074","DOI":"10.1152\/jn.00283.2010","type":"journal-article","article-title":"Topography of covert visual attention in human superior colliculus","volume":"104","author":"Katyal","year":"2010","journal-title":"Journal of Neurophysiology"},{"key":"bib56","doi-asserted-by":"publisher","first-page":"440","DOI":"10.1038\/nn.3643","type":"journal-article","article-title":"Cortical activity in the null space: permitting preparation without movement","volume":"17","author":"Kaufman","year":"2014","journal-title":"Nature Neuroscience"},{"key":"bib57","first-page":"1","type":"journal-article","article-title":"What\u2019s new in psychtoolbox-3?","volume":"36","author":"Kleiner","year":"2007","journal-title":"Perception"},{"key":"bib58","doi-asserted-by":"publisher","first-page":"1961","DOI":"10.1111\/j.1460-9568.2011.07696.x","type":"journal-article","article-title":"Control from below: the role of a midbrain network in spatial attention","volume":"33","author":"Knudsen","year":"2011","journal-title":"The European Journal of Neuroscience"},{"key":"bib59","doi-asserted-by":"publisher","first-page":"1549","DOI":"10.1038\/nn.2663","type":"journal-article","article-title":"Microsaccades precisely relocate gaze in a high visual acuity task","volume":"13","author":"Ko","year":"2010","journal-title":"Nature Neuroscience"},{"key":"bib60","doi-asserted-by":"publisher","first-page":"324","DOI":"10.1016\/0006-8993(85)91528-8","type":"journal-article","article-title":"Projections from the functional subdivisions of the frontal eye field to the superior colliculus in the monkey","volume":"327","author":"Komatsu","year":"1985","journal-title":"Brain Research"},{"key":"bib61","doi-asserted-by":"publisher","first-page":"165","DOI":"10.1146\/annurev-neuro-062012-170249","type":"journal-article","article-title":"Superior colliculus and visual spatial attention","volume":"36","author":"Krauzlis","year":"2013","journal-title":"Annual Review of Neuroscience"},{"key":"bib62","doi-asserted-by":"publisher","first-page":"859","DOI":"10.1016\/j.cub.2018.02.005","type":"journal-article","article-title":"Stress accelerates defensive responses to looming in mice and involves a locus coeruleus-superior colliculus projection","volume":"28","author":"Li","year":"2018","journal-title":"Current Biology"},{"key":"bib63","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-017-02815-3","type":"journal-article","article-title":"Subcortical evidence for a contribution of arousal to fMRI studies of brain activity","volume":"9","author":"Liu","year":"2018","journal-title":"Nature Communications"},{"key":"bib64","doi-asserted-by":"publisher","first-page":"78","DOI":"10.1038\/nature12742","type":"journal-article","article-title":"Context-dependent computation by recurrent dynamics in prefrontal cortex","volume":"503","author":"Mante","year":"2013","journal-title":"Nature"},{"key":"bib65","doi-asserted-by":"publisher","DOI":"10.1038\/s42003-019-0527-y","type":"journal-article","article-title":"Sensorimotor transformation elicits systematic patterns of activity along the dorsoventral extent of the superior colliculus in the macaque monkey","volume":"2","author":"Massot","year":"2019","journal-title":"Communications Biology"},{"key":"bib66","doi-asserted-by":"publisher","DOI":"10.5334\/joc.18","type":"journal-article","article-title":"Pupillometry: psychology, physiology, and function","volume":"1","author":"Math\u00f4t","year":"2018","journal-title":"Journal of Cognition"},{"key":"bib67","doi-asserted-by":"publisher","first-page":"321","DOI":"10.1016\/S0079-6123(05)51011-2","type":"journal-article","article-title":"The mammalian superior colliculus: laminar structure and connections","volume":"151","author":"May","year":"2006","journal-title":"Progress in Brain Research"},{"key":"bib68","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0128428","type":"journal-article","article-title":"Simultaneous recordings of human microsaccades and drifts with a contemporary video eye tracker and the search coil technique","volume":"10","author":"McCamy","year":"2015","journal-title":"PLOS ONE"},{"key":"bib69","doi-asserted-by":"publisher","first-page":"722","DOI":"10.1152\/jn.1976.39.4.722","type":"journal-article","article-title":"Organization of monkey superior colliculus: intermediate layer cells discharging before eye movements","volume":"39","author":"Mohler","year":"1976","journal-title":"Journal of Neurophysiology"},{"key":"bib70","doi-asserted-by":"publisher","first-page":"524","DOI":"10.1073\/pnas.0408311101","type":"journal-article","article-title":"Microstimulation of the superior colliculus focuses attention without moving the eyes","volume":"102","author":"M\u00fcller","year":"2005","journal-title":"PNAS"},{"key":"bib71","doi-asserted-by":"publisher","first-page":"2313","DOI":"10.1152\/jn.1995.73.6.2313","type":"journal-article","article-title":"Saccade-related activity in monkey superior colliculus. I. characteristics of burst and buildup cells","volume":"73","author":"Munoz","year":"1995","journal-title":"Journal of Neurophysiology"},{"key":"bib72","doi-asserted-by":"publisher","first-page":"4140","DOI":"10.1002\/hbm.22466","type":"journal-article","article-title":"Pupil diameter covaries with BOLD activity in human locus coeruleus","volume":"35","author":"Murphy","year":"2014","journal-title":"Human Brain Mapping"},{"key":"bib73","doi-asserted-by":"publisher","first-page":"4468","DOI":"10.1523\/JNEUROSCI.12-11-04468.1992","type":"journal-article","article-title":"Distributed but convergent ordering of corticostriatal projections: analysis of the frontal eye field and the supplementary eye field in the macaque monkey","volume":"12","author":"Parthasarathy","year":"1992","journal-title":"The Journal of Neuroscience"},{"key":"bib74","doi-asserted-by":"crossref","first-page":"437","DOI":"10.1163\/156856897X00366","type":"journal-article","article-title":"The VideoToolbox software for visual psychophysics: transforming numbers into movies","volume":"10","author":"Pelli","year":"1997","journal-title":"Spatial Vision"},{"key":"bib75","doi-asserted-by":"publisher","first-page":"1125","DOI":"10.1016\/0306-4522(84)90007-1","type":"journal-article","article-title":"Retinal ganglion cells that project to the superior colliculus and pretectum in the macaque monkey","volume":"12","author":"Perry","year":"1984","journal-title":"Neuroscience"},{"key":"bib76","doi-asserted-by":"publisher","DOI":"10.1523\/ENEURO.0010-20.2020","type":"journal-article","article-title":"An intersectional viral-genetic method for fluorescent tracing of axon collaterals reveals details of noradrenergic locus coeruleus structure","volume":"7","author":"Plummer","year":"2020","journal-title":"eNeuro"},{"key":"bib77","doi-asserted-by":"publisher","first-page":"300","DOI":"10.1016\/j.exer.2005.07.002","type":"journal-article","article-title":"Development of the human superior colliculus and the retinocollicular projection","volume":"82","author":"Qu","year":"2006","journal-title":"Experimental Eye Research"},{"key":"bib78","doi-asserted-by":"publisher","first-page":"1392","DOI":"10.1152\/jn.01049.2002","type":"journal-article","article-title":"A comparison of macaque behavior and superior colliculus neuronal activity to predictions from models of two-choice decisions","volume":"90","author":"Ratcliff","year":"2003","journal-title":"Journal of Neurophysiology"},{"key":"bib79","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms13289","type":"journal-article","article-title":"Pupil fluctuations track rapid changes in adrenergic and cholinergic activity in cortex","volume":"7","author":"Reimer","year":"2016","journal-title":"Nature Communications"},{"key":"bib80","doi-asserted-by":"publisher","first-page":"1795","DOI":"10.1016\/0042-6989(72)90070-3","type":"journal-article","article-title":"Eye movements evoked by collicular stimulation in the alert monkey","volume":"12","author":"Robinson","year":"1972","journal-title":"Vision Research"},{"key":"bib81","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.2422331122","type":"journal-article","article-title":"Multiplexing of cognitive encoding by oculomotor networks leads to incidental gaze shifts","volume":"122","author":"Rosen","year":"2025","journal-title":"PNAS"},{"key":"bib82","doi-asserted-by":"publisher","first-page":"1669","DOI":"10.1038\/s41593-019-0477-1","type":"journal-article","article-title":"Simultaneous multi-area recordings suggest that attention improves performance by reshaping stimulus representations","volume":"22","author":"Ruff","year":"2019","journal-title":"Nature Neuroscience"},{"key":"bib83","doi-asserted-by":"publisher","first-page":"376","DOI":"10.1038\/s41583-022-00582-9","type":"journal-article","article-title":"Priority coding in the visual system","volume":"23","author":"Rust","year":"2022","journal-title":"Nature Reviews. Neuroscience"},{"key":"bib84","doi-asserted-by":"publisher","first-page":"130","DOI":"10.1016\/j.neuron.2012.09.011","type":"journal-article","article-title":"Orienting and reorienting: the locus coeruleus mediates cognition through arousal","volume":"76","author":"Sara","year":"2012","journal-title":"Neuron"},{"key":"bib85","doi-asserted-by":"publisher","first-page":"249","DOI":"10.1016\/j.neuron.2019.01.026","type":"journal-article","article-title":"Cortical areas interact through a communication subspace","volume":"102","author":"Semedo","year":"2019","journal-title":"Neuron"},{"key":"bib86","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-28552-w","type":"journal-article","article-title":"Feedforward and feedback interactions between visual cortical areas use different population activity patterns","volume":"13","author":"Semedo","year":"2022","journal-title":"Nature Communications"},{"key":"bib87","doi-asserted-by":"publisher","DOI":"10.1038\/s41467-022-33010-8","type":"journal-article","article-title":"A temporal sequence of thalamic activity unfolds at transitions in behavioral arousal state","volume":"13","author":"Setzer","year":"2022","journal-title":"Nature Communications"},{"key":"bib88","doi-asserted-by":"publisher","first-page":"287","DOI":"10.1111\/ejn.12395","type":"journal-article","article-title":"Task difficulty in mental arithmetic affects microsaccadic rates and magnitudes","volume":"39","author":"Siegenthaler","year":"2014","journal-title":"The European Journal of Neuroscience"},{"key":"bib89","doi-asserted-by":"publisher","first-page":"5422","DOI":"10.1523\/JNEUROSCI.4782-12.2013","type":"journal-article","article-title":"Spatial and temporal scales of neuronal correlation in visual area V4","volume":"33","author":"Smith","year":"2013","journal-title":"The Journal of Neuroscience"},{"key":"bib90","type":"software","article-title":"Nasnet","volume-title":"Github","author":"SmithLabNeuro","year":"2025","unstructured":"SmithLabNeuro. 2025. Nasnet. Github. 16729ad. https:\/\/github.com\/SmithLabNeuro\/nasnet."},{"key":"bib91","doi-asserted-by":"publisher","first-page":"118","DOI":"10.1152\/physrev.1986.66.1.118","type":"journal-article","article-title":"Translation of sensory signals into commands for control of saccadic eye movements: role of primate superior colliculus","volume":"66","author":"Sparks","year":"1986","journal-title":"Physiological Reviews"},{"key":"bib92","first-page":"213","type":"journal-article","article-title":"The deep layers of the superior colliculus","volume":"3","author":"Sparks","year":"1989","journal-title":"Reviews of Oculomotor Research"},{"key":"bib93","doi-asserted-by":"publisher","first-page":"309","DOI":"10.1146\/annurev.ne.13.030190.001521","type":"journal-article","article-title":"Signal transformations required for the generation of saccadic eye movements","volume":"13","author":"Sparks","year":"1990","journal-title":"Annual Review of Neuroscience"},{"key":"bib94","doi-asserted-by":"publisher","first-page":"5299","DOI":"10.1016\/j.cub.2021.09.076","type":"journal-article","article-title":"Attention improves information flow between neuronal populations without changing the communication subspace","volume":"31","author":"Srinath","year":"2021","journal-title":"Current Biology"},{"key":"bib95","doi-asserted-by":"publisher","first-page":"493","DOI":"10.1002\/cne.902710403","type":"journal-article","article-title":"Frontal eye field efferents in the macaque monkey: II. Topography of terminal fields in midbrain and pons","volume":"271","author":"Stanton","year":"1988","journal-title":"The Journal of Comparative Neurology"},{"key":"bib96","doi-asserted-by":"publisher","first-page":"15272","DOI":"10.1073\/pnas.1907962116","type":"journal-article","article-title":"The role of the optic tectum for visually evoked orienting and evasive movements","volume":"116","author":"Suzuki","year":"2019","journal-title":"PNAS"},{"key":"bib97","doi-asserted-by":"publisher","first-page":"940","DOI":"10.1016\/j.neuron.2018.01.032","type":"journal-article","article-title":"The basal forebrain regulates global resting-state fMRI fluctuations","volume":"97","author":"Turchi","year":"2018","journal-title":"Neuron"},{"key":"bib98","doi-asserted-by":"publisher","DOI":"10.3389\/fnhum.2019.00340","type":"journal-article","article-title":"Brainstem modulation of large-scale intrinsic cortical activity correlations","volume":"13","author":"van den Brink","year":"2019","journal-title":"Frontiers in Human Neuroscience"},{"key":"bib99","doi-asserted-by":"publisher","first-page":"3629","DOI":"10.1523\/JNEUROSCI.5512-11.2012","type":"journal-article","article-title":"Microstimulation of the monkey superior colliculus induces pupil dilation without evoking saccades","volume":"32","author":"Wang","year":"2012","journal-title":"The Journal of Neuroscience"},{"key":"bib100","doi-asserted-by":"publisher","first-page":"408","DOI":"10.1523\/JNEUROSCI.3550-13.2014","type":"journal-article","article-title":"Transient pupil response is modulated by contrast-based saliency","volume":"34","author":"Wang","year":"2014","journal-title":"The Journal of Neuroscience"},{"key":"bib101","doi-asserted-by":"publisher","first-page":"10446","DOI":"10.1073\/pnas.1809668115","type":"journal-article","article-title":"Neural basis of location-specific pupil luminance modulation","volume":"115","author":"Wang","year":"2018","journal-title":"PNAS"},{"key":"bib102","doi-asserted-by":"publisher","first-page":"919","DOI":"10.1162\/jocn_a_01688","type":"journal-article","article-title":"Coordination of pupil and saccade responses by the superior colliculus","volume":"33","author":"Wang","year":"2021","journal-title":"Journal of Cognitive Neuroscience"},{"key":"bib103","doi-asserted-by":"publisher","DOI":"10.3389\/fncir.2022.792959","type":"journal-article","article-title":"Functional organisation of the mouse superior colliculus","volume":"16","author":"Wheatcroft","year":"2022","journal-title":"Frontiers in Neural Circuits"},{"key":"bib104","doi-asserted-by":"publisher","DOI":"10.1038\/ncomms14263","type":"journal-article","article-title":"Superior colliculus neurons encode a visual saliency map during free viewing of natural dynamic video","volume":"8","author":"White","year":"2017","journal-title":"Nature Communications"},{"key":"bib105","doi-asserted-by":"publisher","first-page":"1387","DOI":"10.1016\/0042-6989(76)90156-5","type":"journal-article","article-title":"Microsaccades during finely guided visuomotor tasks","volume":"16","author":"Winterson","year":"1976","journal-title":"Vision Research"},{"key":"bib106","doi-asserted-by":"publisher","first-page":"189","DOI":"10.1146\/annurev.ne.03.030180.001201","type":"journal-article","article-title":"Visual-motor function of the primate superior colliculus","volume":"3","author":"Wurtz","year":"1980","journal-title":"Annual Review of Neuroscience"},{"key":"bib107","doi-asserted-by":"publisher","first-page":"328","DOI":"10.1037\/a0016826","type":"journal-article","article-title":"The optic tectum of birds: mapping our way to understanding visual processing","volume":"63","author":"Wylie","year":"2009","journal-title":"Canadian Journal of Experimental Psychology"},{"key":"bib108","doi-asserted-by":"publisher","first-page":"434","DOI":"10.1038\/nature11497","type":"journal-article","article-title":"Attention deficits without cortical neuronal deficits","volume":"489","author":"Z\u00e9non","year":"2012","journal-title":"Nature"},{"key":"bib109","doi-asserted-by":"publisher","first-page":"1459","DOI":"10.1126\/science.150.3702.1459","type":"journal-article","article-title":"Microsaccades and the velocity-amplitude relationship for saccadic eye movements","volume":"150","author":"Zuber","year":"1965","journal-title":"Science"},{"key":"dataset1","doi-asserted-by":"publisher","DOI":"10.1184\/R1\/32300970","type":"dataset","article-title":"Data from \"Brain-wide arousal signals are segregated from movement planning in the superior colliculus\"","volume-title":"Carnegie Mellon University","author":"Johnston","year":"2026","unstructured":"Johnston R, Smith MA. 2026. Data from \"Brain-wide arousal signals are segregated from movement planning in the superior colliculus\". Carnegie Mellon University."}],"container-title":["eLife"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/cdn.elifesciences.org\/articles\/99278\/elife-99278-v1.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/cdn.elifesciences.org\/articles\/99278\/elife-99278-v1.xml","content-type":"application\/xml","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/elifesciences.org\/articles\/99278","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T08:58:55Z","timestamp":1787561935000},"score":1,"resource":{"primary":{"URL":"https:\/\/elifesciences.org\/articles\/99278"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,6,8]]},"references-count":110,"alternative-id":["10.7554\/eLife.99278"],"URL":"https:\/\/doi.org\/10.7554\/elife.99278","archive":["CLOCKSS"],"relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2024.04.26.591284","asserted-by":"subject"}],"is-same-as":[{"id-type":"doi","id":"10.7554\/eLife.99278.4","asserted-by":"subject"}],"is-version-of":[{"id-type":"doi","id":"10.7554\/eLife.99278.1","asserted-by":"subject"},{"id-type":"doi","id":"10.7554\/eLife.99278.2","asserted-by":"subject"},{"id-type":"doi","id":"10.7554\/eLife.99278.3","asserted-by":"subject"}],"is-supplemented-by":[{"id-type":"doi","id":"10.1184\/R1\/32300970","asserted-by":"subject"}]},"ISSN":["2050-084X"],"issn-type":[{"value":"2050-084X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,6,8]]},"assertion":[{"value":"single anonymised","name":"peer_review_transparency","label":"Peer review transparency","group":{"name":"peer_review_taxonomy"}},{"value":"other reviewer(s), editor","name":"peer_review_interaction","label":"Peer review interaction","group":{"name":"peer_review_taxonomy"}},{"value":"review summaries, review reports, author\/editor communication, reviewer identities reviewer opt in, editor identities","name":"peer_review_published","label":"Peer review published","group":{"name":"peer_review_taxonomy"}},{"value":"open (sign in with ORCID iD required)","name":"post_publication_commenting","label":"Post publication commenting","group":{"name":"post_publication_commenting"}}],"article-number":"RP99278"}}