{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,5,7]],"date-time":"2026-05-07T01:56:26Z","timestamp":1778118986491,"version":"3.51.4"},"reference-count":49,"publisher":"MIT Press - Journals","issue":"4","content-domain":{"domain":["direct.mit.edu"],"crossmark-restriction":true},"short-container-title":[],"published-print":{"date-parts":[[2020,4,1]]},"abstract":"<jats:title>Abstract<\/jats:title>\n               <jats:p>Perceptual decision-making has been shown to be influenced by reward expected from alternative options or actions, but the underlying neural mechanisms are currently unknown. More specifically, it is debated whether reward effects are mediated through changes in sensory processing, later stages of decision-making, or both. To address this question, we conducted two experiments in which human participants made saccades to what they perceived to be either the first or second of two visually identical but asynchronously presented targets while we manipulated expected reward from correct and incorrect responses on each trial. By comparing reward-induced bias in target selection (i.e., reward bias) during the two experiments, we determined whether reward caused changes in sensory or decision-making processes. We found similar reward biases in the two experiments indicating that reward information mainly influenced later stages of decision-making. Moreover, the observed reward biases were independent of the individual's sensitivity to sensory signals. This suggests that reward effects were determined heuristically via modulation of decision-making processes instead of sensory processing. To further explain our findings and uncover plausible neural mechanisms, we simulated our experiments with a cortical network model and tested alternative mechanisms for how reward could exert its influence. We found that our experimental observations are more compatible with reward-dependent input to the output layer of the decision circuit. Together, our results suggest that, during a temporal judgment task, reward exerts its influence via changing later stages of decision-making (i.e., response bias) rather than early sensory processing (i.e., perceptual bias).<\/jats:p>","DOI":"10.1162\/jocn_a_01516","type":"journal-article","created":{"date-parts":[[2019,12,18]],"date-time":"2019-12-18T18:09:09Z","timestamp":1576692549000},"page":"674-690","update-policy":"https:\/\/doi.org\/10.1162\/mitpressjournals.corrections.policy","source":"Crossref","is-referenced-by-count":11,"title":["Influence of Expected Reward on Temporal Order Judgment"],"prefix":"10.1162","volume":"32","author":[{"given":"Mohsen","family":"Rakhshan","sequence":"first","affiliation":[{"name":"Dartmouth College"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Vivian","family":"Lee","sequence":"additional","affiliation":[{"name":"Dartmouth College"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Emily","family":"Chu","sequence":"additional","affiliation":[{"name":"Dartmouth College"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Lauren","family":"Harris","sequence":"additional","affiliation":[{"name":"Florida State University"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Lillian","family":"Laiks","sequence":"additional","affiliation":[{"name":"Texas A&M University"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Peyman","family":"Khorsand","sequence":"additional","affiliation":[{"name":"Dartmouth College"}],"role":[{"role":"author","vocabulary":"crossref"}]},{"given":"Alireza","family":"Soltani","sequence":"additional","affiliation":[{"name":"Dartmouth College"}],"role":[{"role":"author","vocabulary":"crossref"}]}],"member":"281","published-online":{"date-parts":[[2020,4,1]]},"reference":[{"key":"2022042815031102700_bib1","doi-asserted-by":"crossref","unstructured":"Bang,  J. W., & Rahnev,  D. (2017). Stimulus expectation alters decision criterion but not sensory signal in perceptual decision making. Scientific Reports, 7, 17072.","DOI":"10.1038\/s41598-017-16885-2"},{"key":"2022042815031102700_bib2","doi-asserted-by":"crossref","unstructured":"Brainard,  D. H.\n           (1997). The Psychophysics Toolbox. Spatial Vision, 10, 433\u2013436.","DOI":"10.1163\/156856897X00357"},{"key":"2022042815031102700_bib3","doi-asserted-by":"crossref","unstructured":"Carrasco,  M., & Barbot,  A. (2019). Spatial attention alters visual appearance. Current Opinion in Psychology, 29, 56\u201364.","DOI":"10.1016\/j.copsyc.2018.10.010"},{"key":"2022042815031102700_bib4","doi-asserted-by":"crossref","unstructured":"Carrasco,  M., Ling,  S., & Read,  S. (2004). Attention alters appearance. Nature Neuroscience, 7, 308\u2013313.","DOI":"10.1038\/nn1194"},{"key":"2022042815031102700_bib5","doi-asserted-by":"crossref","unstructured":"Christopoulos,  V., Bonaiuto,  J., & Andersen,  R. A. (2015). A biologically plausible computational theory for value integration and action selection in decisions with competing alternatives. PLoS Computational Biology, 11, e1004104.","DOI":"10.1371\/journal.pcbi.1004104"},{"key":"2022042815031102700_bib6","doi-asserted-by":"crossref","unstructured":"Christopoulos,  V., & Schrater,  P. R. (2015). Dynamic integration of value information into a common probability currency as a theory for flexible decision making. PLoS Computational Biology, 11, e1004402.","DOI":"10.1371\/journal.pcbi.1004402"},{"key":"2022042815031102700_bib7","doi-asserted-by":"crossref","unstructured":"Cicmil,  N., Cumming,  B. G., Parker,  A. J., & Krug,  K. (2015). Reward modulates the effect of visual cortical microstimulation on perceptual decisions. eLife, 4, e07832.","DOI":"10.7554\/eLife.07832"},{"key":"2022042815031102700_bib8","doi-asserted-by":"crossref","unstructured":"Diederich,  A.\n           (2008). A further test of sequential-sampling models that account for payoff effects on response bias in perceptual decision tasks. Perception & Psychophysics, 70, 229\u2013256.","DOI":"10.3758\/PP.70.2.229"},{"key":"2022042815031102700_bib9","doi-asserted-by":"crossref","unstructured":"Diederich,  A., & Busemeyer,  J. R. (2006). Modeling the effects of payoff on response bias in a perceptual discrimination task: Bound-change, drift-rate-change, or two-stage-processing hypothesis. Perception & Psychophysics, 68, 194\u2013207.","DOI":"10.3758\/BF03193669"},{"key":"2022042815031102700_bib10","doi-asserted-by":"crossref","unstructured":"Farashahi,  S., Azab,  H., Hayden,  B., & Soltani,  A. (2018). On the flexibility of basic risk attitudes in monkeys. Journal of Neuroscience, 38, 4383\u20134398.","DOI":"10.1523\/JNEUROSCI.2260-17.2018"},{"key":"2022042815031102700_bib11","doi-asserted-by":"crossref","unstructured":"Farashahi,  S., Donahue,  C. H., Khorsand,  P., Seo,  H., Lee,  D., & Soltani,  A. (2017). Metaplasticity as a neural substrate for adaptive learning and choice under uncertainty. Neuron, 94, 401\u2013414.","DOI":"10.1016\/j.neuron.2017.03.044"},{"key":"2022042815031102700_bib12","doi-asserted-by":"crossref","unstructured":"Farashahi,  S., Ting,  C.-C., Kao,  C.-H., Wu,  S.-W., & Soltani,  A. (2018). Dynamic combination of sensory and reward information under time pressure. PLoS Computational Biology, 14, e1006070.","DOI":"10.1371\/journal.pcbi.1006070"},{"key":"2022042815031102700_bib13","doi-asserted-by":"crossref","unstructured":"Feng,  S., Holmes,  P., Rorie,  A., & Newsome,  W. T. (2009). Can monkeys choose optimally when faced with noisy stimuli and unequal rewards?PLoS Computational Biology, 5, e1000284.","DOI":"10.1371\/journal.pcbi.1000284"},{"key":"2022042815031102700_bib14","doi-asserted-by":"crossref","unstructured":"Gao,  J., Tortell,  R., & McClelland,  J. L. (2011). Dynamic integration of reward and stimulus information in perceptual decision-making. PLoS One, 6, e16749.","DOI":"10.1371\/journal.pone.0016749"},{"key":"2022042815031102700_bib15","doi-asserted-by":"crossref","unstructured":"Gold,  J. I., & Shadlen,  M. N. (2001). Neural computations that underlie decisions about sensory stimuli. Trends in Cognitive Sciences, 5, 10\u201316.","DOI":"10.1016\/S1364-6613(00)01567-9"},{"key":"2022042815031102700_bib16","doi-asserted-by":"crossref","unstructured":"Hickey,  C., Chelazzi,  L., & Theeuwes,  J. (2010). Reward changes salience in human vision via the anterior cingulate. Journal of Neuroscience, 30, 11096\u201311103.","DOI":"10.1523\/JNEUROSCI.1026-10.2010"},{"key":"2022042815031102700_bib17","doi-asserted-by":"crossref","unstructured":"Hikosaka,  O., Miyauchi,  S., & Shimojo,  S. (1993). Focal visual attention produces illusory temporal order and motion sensation. Vision Research, 33, 1219\u20131240.","DOI":"10.1016\/0042-6989(93)90210-N"},{"key":"2022042815031102700_bib18","doi-asserted-by":"crossref","unstructured":"Khorsand,  P., & Soltani,  A. (2017). Optimal structure of metaplasticity for adaptive learning. PLoS Computational Biology, 13, e1005630.","DOI":"10.1371\/journal.pcbi.1005630"},{"key":"2022042815031102700_bib19","unstructured":"Kleiner,  M., Brainard,  D. H., Pelli,  D. G., Ingling,  A., Murray,  R., & Broussard,  C. (2007). What's new in Psychtoolbox-3?Perception, 36, 1\u201316."},{"key":"2022042815031102700_bib20","doi-asserted-by":"crossref","unstructured":"Liston,  D. B., & Stone,  L. S. (2008). Effects of prior information and reward on oculomotor and perceptual choices. Journal of Neuroscience, 28, 13866\u201313875.","DOI":"10.1523\/JNEUROSCI.3120-08.2008"},{"key":"2022042815031102700_bib21","doi-asserted-by":"crossref","unstructured":"Maddox,  W. T.\n           (2002). Toward a unified theory of decision criterion learning in perceptual categorization. Journal of the Experimental Analysis of Behavior, 78, 567\u2013595.","DOI":"10.1901\/jeab.2002.78-567"},{"key":"2022042815031102700_bib22","doi-asserted-by":"crossref","unstructured":"Maunsell,  J. H. R.\n           (2004). Neuronal representations of cognitive state: Reward or attention?Trends in Cognitive Sciences, 8, 261\u2013265.","DOI":"10.1016\/j.tics.2004.04.003"},{"key":"2022042815031102700_bib23","doi-asserted-by":"crossref","unstructured":"Mulder,  M. J., Wagenmakers,  E.-J., Ratcliff,  R., Boekel,  W., & Forstmann,  B. U. (2012). Bias in the brain: A diffusion model analysis of prior probability and potential payoff. Journal of Neuroscience, 32, 2335\u20132343.","DOI":"10.1523\/JNEUROSCI.4156-11.2012"},{"key":"2022042815031102700_bib24","doi-asserted-by":"crossref","unstructured":"Navalpakkam,  V., Koch,  C., & Perona,  P. (2009). Homo economicus in visual search. Journal of Vision, 9, 31.","DOI":"10.1167\/9.1.31"},{"key":"2022042815031102700_bib25","doi-asserted-by":"crossref","unstructured":"Pelli,  D. G.\n           (1997). The VideoToolbox software for visual psychophysics: Transforming numbers into movies. Spatial Vision, 10, 437\u2013442.","DOI":"10.1163\/156856897X00366"},{"key":"2022042815031102700_bib26","doi-asserted-by":"crossref","unstructured":"Pleger,  B., Blankenburg,  F., Ruff,  C. C., Driver,  J., & Dolan,  R. J. (2008). Reward facilitates tactile judgments and modulates hemodynamic responses in human primary somatosensory cortex. Journal of Neuroscience, 28, 8161\u20138168.","DOI":"10.1523\/JNEUROSCI.1093-08.2008"},{"key":"2022042815031102700_bib27","doi-asserted-by":"crossref","unstructured":"Rajsic,  J., Perera,  H., & Pratt,  J. (2017). Learned value and object perception: Accelerated perception or biased decisions?Attention, Perception, & Psychophysics, 79, 603\u2013613.","DOI":"10.3758\/s13414-016-1242-0"},{"key":"2022042815031102700_bib28","doi-asserted-by":"crossref","unstructured":"Rorie,  A. E., Gao,  J., McClelland,  J. L., & Newsome,  W. T. (2010). Integration of sensory and reward information during perceptual decision-making in Lateral Intraparietal Cortex (LIP) of the macaque monkey. PLoS One, 5, e9308.","DOI":"10.1371\/journal.pone.0009308"},{"key":"2022042815031102700_bib29","doi-asserted-by":"crossref","unstructured":"Rungratsameetaweemana,  N., Itthipuripat,  S., Salazar,  A., & Serences,  J. T. (2018). Expectations do not alter early sensory processing during perceptual decision-making. Journal of Neuroscience, 38, 5632\u20135648.","DOI":"10.1523\/JNEUROSCI.3638-17.2018"},{"key":"2022042815031102700_bib30","doi-asserted-by":"crossref","unstructured":"Schiller,  P. H., & Chou,  I.-H. (1998). The effects of frontal eye field and dorsomedial frontal cortex lesions on visually guided eye movements. Nature Neuroscience, 1, 248\u2013253.","DOI":"10.1038\/693"},{"key":"2022042815031102700_bib31","doi-asserted-by":"crossref","unstructured":"Schiller,  P. H., & Chou,  I.-H. (2000). The effects of anterior arcuate and dorsomedial frontal cortex lesions on visually guided eye movements: 2. Paired and multiple targets. Vision Research, 40, 1627\u20131638.","DOI":"10.1016\/S0042-6989(00)00058-4"},{"key":"2022042815031102700_bib32","doi-asserted-by":"crossref","unstructured":"Schiller,  P. H., & Tehovnik,  E. J. (2003). Cortical inhibitory circuits in eye-movement generation. European Journal of Neuroscience, 18, 3127\u20133133.","DOI":"10.1111\/j.1460-9568.2003.03036.x"},{"key":"2022042815031102700_bib33","doi-asserted-by":"crossref","unstructured":"Schneider,  K. A., & Bavelier,  D. (2003). Components of visual prior entry. Cognitive Psychology, 47, 333\u2013366.","DOI":"10.1016\/S0010-0285(03)00035-5"},{"key":"2022042815031102700_bib34","doi-asserted-by":"crossref","unstructured":"Serences,  J. T.\n           (2008). Value-based modulations in human visual cortex. Neuron, 60, 1169\u20131181.","DOI":"10.1016\/j.neuron.2008.10.051"},{"key":"2022042815031102700_bib35","doi-asserted-by":"crossref","unstructured":"Shore,  D. I., Spence,  C., & Klein,  R. M. (2001). Visual prior entry. Psychological Science, 12, 205\u2013212.","DOI":"10.1111\/1467-9280.00337"},{"key":"2022042815031102700_bib36","doi-asserted-by":"crossref","unstructured":"Soltani,  A., Khorsand,  P., Guo,  C., Farashahi,  S., & Liu,  J. (2016). Neural substrates of cognitive biases during probabilistic inference. Nature Communications, 7, 11393.","DOI":"10.1038\/ncomms11393"},{"key":"2022042815031102700_bib37","doi-asserted-by":"crossref","unstructured":"Soltani,  A., Noudoost,  B., & Moore,  T. (2013). Dissociable dopaminergic control of saccadic target selection and its implications for reward modulation. Proceedings of the National Academy of Sciences, U.S.A., 110, 3579\u20133584.","DOI":"10.1073\/pnas.1221236110"},{"key":"2022042815031102700_bib38","doi-asserted-by":"crossref","unstructured":"Spitmaan,  M., Chu,  E., & Soltani,  A. (2019). Salience-driven value construction for adaptive choice under risk. Journal of Neuroscience, 39, 5195\u20135209.","DOI":"10.1523\/JNEUROSCI.2522-18.2019"},{"key":"2022042815031102700_bib39","doi-asserted-by":"crossref","unstructured":"Stanford,  T. R., Shankar,  S., Massoglia,  D. P., Costello,  M. G., & Salinas,  E. (2010). Perceptual decision making in less than 30 milliseconds. Nature Neuroscience, 13, 379\u2013385.","DOI":"10.1038\/nn.2485"},{"key":"2022042815031102700_bib40","doi-asserted-by":"crossref","unstructured":"St\u0103ni\u015for,  L., van der Togt,  C., Pennartz,  C. M. A., & Roelfsema,  P. R. (2013). A unified selection signal for attention and reward in primary visual cortex. Proceedings of the National Academy of Sciences, U.S.A., 110, 9136\u20139141.","DOI":"10.1073\/pnas.1300117110"},{"key":"2022042815031102700_bib41","doi-asserted-by":"crossref","unstructured":"Stelmach,  L. B., & Herdman,  C. M. (1991). Directed attention and perception of temporal order. Journal of Experimental Psychology: Human Perception and Performance, 17, 539\u2013550.","DOI":"10.1037\/0096-1523.17.2.539"},{"key":"2022042815031102700_bib42","doi-asserted-by":"crossref","unstructured":"Sugrue,  L. P., Corrado,  G. S., & Newsome,  W. T. (2005). Choosing the greater of two goods: Neural currencies for valuation and decision making. Nature Reviews Neuroscience, 6, 363\u2013375.","DOI":"10.1038\/nrn1666"},{"key":"2022042815031102700_bib43","doi-asserted-by":"crossref","unstructured":"Summerfield,  C., & Koechlin,  E. (2010). Economic value biases uncertain perceptual choices in the parietal and prefrontal cortices. Frontiers in Human Neuroscience, 4, 208.","DOI":"10.3389\/fnhum.2010.00208"},{"key":"2022042815031102700_bib44","doi-asserted-by":"crossref","unstructured":"Teichert,  T., & Ferrera,  V. P. (2010). Suboptimal integration of reward magnitude and prior reward likelihood in categorical decisions by monkeys. Frontiers in Neuroscience, 4, 186.","DOI":"10.3389\/fnins.2010.00186"},{"key":"2022042815031102700_bib45","doi-asserted-by":"crossref","unstructured":"Tosoni,  A., Committeri,  G., Calluso,  C., & Galati,  G. (2017). The effect of reward expectation on the time course of perceptual decisions. European Journal of Neuroscience, 45, 1152\u20131164.","DOI":"10.1111\/ejn.13555"},{"key":"2022042815031102700_bib46","doi-asserted-by":"crossref","unstructured":"Tversky,  A., & Kahneman,  D. (1992). Advances in prospect theory: Cumulative representation of uncertainty. Journal of Risk and Uncertainty, 5, 297\u2013323.","DOI":"10.1007\/BF00122574"},{"key":"2022042815031102700_bib47","doi-asserted-by":"crossref","unstructured":"Vassena,  E., Deraeve,  J., & Alexander,  W. H. (2019). Task-specific prioritization of reward and effort information: Novel insights from behavior and computational modeling. Cognitive, Affective, & Behavioral Neuroscience, 19, 619\u2013636.","DOI":"10.3758\/s13415-018-00685-w"},{"key":"2022042815031102700_bib48","doi-asserted-by":"crossref","unstructured":"Voss,  A., Rothermund,  K., & Brandtst\u00e4dter,  J. (2008). Interpreting ambiguous stimuli: Separating perceptual and judgmental biases. Journal of Experimental Social Psychology, 44, 1048\u20131056.","DOI":"10.1016\/j.jesp.2007.10.009"},{"key":"2022042815031102700_bib49","doi-asserted-by":"crossref","unstructured":"Wong,  K.-F., & Wang,  X.-J. (2006). A recurrent network mechanism of time integration in perceptual decisions. Journal of Neuroscience, 26, 1314\u20131328.","DOI":"10.1523\/JNEUROSCI.3733-05.2006"}],"container-title":["Journal of Cognitive Neuroscience"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/direct.mit.edu\/jocn\/article-pdf\/32\/4\/674\/2013176\/jocn_a_01516.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"https:\/\/direct.mit.edu\/jocn\/article-pdf\/32\/4\/674\/2013176\/jocn_a_01516.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2022,4,29]],"date-time":"2022-04-29T00:16:47Z","timestamp":1651191407000},"score":1,"resource":{"primary":{"URL":"https:\/\/direct.mit.edu\/jocn\/article\/32\/4\/674\/95399\/Influence-of-Expected-Reward-on-Temporal-Order"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2020,4,1]]},"references-count":49,"journal-issue":{"issue":"4","published-online":{"date-parts":[[2020,4,1]]},"published-print":{"date-parts":[[2020,4,1]]}},"URL":"https:\/\/doi.org\/10.1162\/jocn_a_01516","relation":{},"ISSN":["0898-929X","1530-8898"],"issn-type":[{"value":"0898-929X","type":"print"},{"value":"1530-8898","type":"electronic"}],"subject":[],"published-other":{"date-parts":[[2020,4]]},"published":{"date-parts":[[2020,4,1]]}}}