Behaviorally relevant decision coding in primary somatosensory cortex neurons
Open accessAbstract
Primary sensory cortex is thought to process incoming sensory information, while decision variables important for driving behavior are assumed to arise downstream in the processing hierarchy. Here, we used population two-photon calcium imaging and targeted two-photon optogenetic stimulation of neurons in layer 2/3 of mouse primary somatosensory cortex (S1) during a texture discrimination task to test for the presence of decision signals and probe their behavioral relevance. Small but distinct populations of neurons carried information about the stimulus irrespective of the behavioral outcome (stimulus neurons), or about the choice irrespective of the presented stimulus (decision neurons). Decision neurons show categorical coding that develops during learning, and lack a conclusive decision signal in Miss trials. All-optical photostimulation of decision neurons during behavior improves behavioral performance, establishing a causal role in driving behavior. The fact that stimulus and decision neurons are intermingled challenges the idea of S1 as a purely sensory area, and causal perturbation suggests a direct involvement of S1 decision neurons in the decision-making process.
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The primary sensory cortex not only retains relatively weak and sparse choice-related activity but also, in this naturalistic behavioral task, directly participates in categorical coding of an all-or-none decision variable into which the majority of neural populations are recruited.
“These observations indicate that the primary sensory cortex not only retains relatively weak and sparse choice-related activity, as it was shown previously (10, 11, 19–25), but also, in our naturalistic behavioral task, directly participates in a categorical coding of all-or-none DV into which the majority of neural populations are recruited.”
From Neural correlates of perceptual decision-making in the primary somatosensory cortex (2026) · cites this paper as [24]
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Experimental — accuracy not yet measuredNeural activity in subpopulations of layer 2/3 wS1 neurons is correlated with behavioral choice.
“More recent experiments (22–25) have shown neural activity in subpopulations of layer 2/3 of wS1 neurons is correlated with behavioral choice.”
From Neural correlates of perceptual decision-making in the primary somatosensory cortex (2026) · cites this paper as [24]
Sensory S1 and S2 cortices have strongly lateralized ipsilateral outputs and are densely and reciprocally interconnected, forming a top-down feedback loop that exhibits choice-related activity.
“In contrast, sensory S1 and S2 cortices have strongly lateralized ipsilateral outputs (43, 59) and are densely and reciprocally interconnected (60) forming a top–down feedback loop that exhibits choice-related activity (10, 11, 22–25).”
From Neural correlates of perceptual decision-making in the primary somatosensory cortex (2026) · cites this paper as [24]
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BackgroundGauld, O. M., Packer, A. M., Russell, L. E., Dalgleish, H. W. P., Iuga, M., Sacadura, F., Roth, A., Clark, B. A., & Häusser, M. (2024) · Neuron · doi:10.1016/j.neuron.2024.04.015 · Cited by 18 · see citing works
“111,112 In contrast with recent studies involving the selective optical stimulation of functionally tuned neurons during behavior, 43,45,46,100,109,113,114 we did not find a significant effect of target ensemble tuning on task performance.”
Self-citationComputational methods to study information processing in neural circuits
Koren, V., Bondanelli, G., & Panzeri, S. (2023) · Computational and Structural Biotechnology Journal · doi:10.1016/j.csbj.2023.01.009 · Cited by 21 · see citing works
“These synaptic currents decorrelate the activity of neurons with similar stimulus selectivity, which has been empirically observed in primary sensory cortices [55,56].”
Network influence determines the impact of cortical ensembles on stimulus detection
Bounds, H. A., & Adesnik, H. (2025) · Neuron · doi:10.1016/j.neuron.2025.04.023 · Cited by 19 · see citing works
“…cortex suggested that an ensem-ble’s information about the animal’s choice determines the impact of an ensemble on behavior in a two-choice task, 44 and many others have posited that a correlation with the choice of the animal suggests that a neuron is involved in downstream readout…”
InfluentialNetwork influence determines the impact of cortical ensembles on stimulus detection
Bounds, H. A., & Adesnik, H. (2024) · bioRxiv · doi:10.1101/2024.08.18.608496 · Cited by 11
“…cortex suggested that an ensemble’s information about the animal’s choice determines the impact of an ensemble on behavior in a two choice task 46 , and many others have posited that a correlation with choice of the animal suggests that a neuron is involved in downstream readout…”
Steinfeld, R., Tacão-Monteiro, A., & Renart, A. (2024) · Current Biology · doi:10.1016/j.cub.2024.04.040 · Cited by 17 · see citing works
“29 Neurons in the deep cortical layers of the sensory cortex have been shown to encode action preparation in non-deci-sion settings 30,31 and choice in go/no-go somatosensory 17,32 and visual detection 33 and forced choice 22 tasks.”
Wei, H.-R., Tang, L. F., Yang, X., Chen, C., Li, L.-X., Mao, Y., Wang, D., Kang, F., Qun, S., Zhang, Z., et al. (2024) · Cell Reports · doi:10.1016/j.celrep.2024.113804 · Cited by 10 · see citing works
Self-citationMulti-region coordination of evidence accumulation and decision commitment in the brainPreprint
Bondy, A., Charlton, J. A., Luo, T. Z., Kopec, C. D., Stagnaro, W. M., Venditto, S. J. C., Shi, K., Xie, E. Y., Beau, M., Lynch, L. A., et al. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.08.21.609044 · Cited by 22 · see citing works
“S1 is reciprocally connected to M1 60 , projects broadly within striatum 54 , and has recently been established to play a causal role in the formation of perceptual decisions 61 .”
An adaptive and flexible role for primary sensory cortex
Waiblinger, C., Reedy, A. R., & Stanley, G. B. (2025) · Nature Neuroscience · doi:10.1038/s41593-025-02124-9 · Cited by 8 · see citing works
“Empirical evidence for this model demonstrates that reward-based learning in response to prolonged stimulus exposure can modify stimulus selectivity and behavior-related representations in the auditory 70 , visual 71 and somatosensory 55,72 cortices.”
A fluid-walled microfluidic platform for human neuron microcircuits and directed axotomy
Nebuloni, F., Do, Q. B., Cook, P. R., Walsh, E. J., & Wade-Martins, R. (2023) · bioRxiv · doi:10.1101/2023.10.14.562004 · Cited by 8
Coexistence of state, choice, and sensory integration coding in barrel cortex LII/III
Gardères, P.-M., Le Gal, S., Rousseau, C., Mamane, A., Ganea, D. A., & Haiss, F. (2024) · Nature Communications · doi:10.1038/s41467-024-49129-9 · Cited by 7 · see citing works
The therapeutic potential of early exercise in Alzheimer's disease: Focus on the brain-spleen axis
Wang, T., Feng, L., & Tian, Z. (2025) · Ageing Research Reviews · doi:10.1016/j.arr.2025.102980 · Cited by 6 · see citing works
Decoding chronic pain: the glutamate-GABA tug of war in the cerebral cortex
Huang, D., Dong, Y., He, L., Zhou, R., Zhou, J., Yang, S., & Yu, S. (2025) · Frontiers in Molecular Neuroscience · doi:10.3389/fnmol.2025.1572775 · Cited by 8 · see citing works
“…across the brain, orchestrates responses to the multifaceted components of pain, and ultimately drives adaptive changes in behavior, cognition, and emotional state (Gan et al., 2022; Li et al., 2022; Martins et al., 2015; Falconi-Sobrinho et al., 2017; Singh et al., 2020; Buetfering et al., 2022).”
A real-time all-optical interface for dynamic perturbation of neural activity during behavior
Zhang, Z., Dzialecka, P., Russell, L. E., Ratto, R., Buetfering, C., Gauld, O. M., Selviah, D. R., & Häusser, M. (2025) · Cell Reports Methods · doi:10.1016/j.crmeth.2025.101180 · Cited by 5 · see citing works
“7 17 Interventions in the activity of the right neurons at precisely the right time point in the development of these neural dynamics are, therefore, key to testing the causal roles of specific activity patterns.”
Self-citationA cortical output channel for perceptual categorization
Schneider, N. A., Malina, M., & Williamson, R. S. (2025) · bioRxiv · doi:10.1101/2025.09.05.674544 · Cited by 4
“Re-310 sponses related to behavioral choice have been observed in multiple primary sensory cortices within both superficial and deep layers [44, 61–64, 66–68, 76, 89–91].”
Feature-specific inhibitory connectivity augments the accuracy of cortical representationsPreprint
Ogando, M. B., Abdeladim, L., Sit, K. K., Shin, H., Sridharan, S., Gopakumar, K., & Adesnik, H. (2025) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2025.08.02.668307 · Cited by 4 · see citing works
“Previous studies have yielded conflicting results as to whether photostimulating similarly tuned PC ensembles amplifies or suppresses local co-tuned representations 4,17,18,24,25,54 .”
Dai, J., & Sun, Q. (2024) · Cell Reports · doi:10.1016/j.celrep.2024.114672 · Cited by 3 · see citing works
Learning within a sensory-motor circuit links action to expected outcomePreprint
Zhou, W., & Schneider, D. M. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.02.08.579532 · Cited by 5 · see citing works
Do, J., Jung, M. W., & Lee, D. (2023) · Scientific Reports · doi:10.1038/s41598-023-49862-z · Cited by 8 · see citing works
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Precision cutaneous stimulation in freely moving mice
Parkes, I., Schorscher-Petcu, A., Gan, Q., & Browne, L. E. (2025) · bioRxiv · doi:10.1101/2024.08.06.606618 · Cited by 3
Context-dependent activation of V1 parvalbumin interneurons enhances visual discrimination
Kukovska, L., Wilmes, K. A., Homma, N. Y., Claudia Clopath, A., & Poort, J. (2025) · PLoS Biology · doi:10.1371/journal.pbio.3003518 · Cited by 5 · see citing works
“Future studies could determine contributions of specific subpopulations of neurons using two-photon photostimulation [80,81].”
Inácio, A. R. L., Lam, K. C., Zhao, Y., Pereira, F., Gerfen, C. R., & Lee, S. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.05.25.542329 · Cited by 6 · see citing works
Shin, S., Oh, J., Kim, S. K., Lee, Y., & Kim, S. J. (2025) · Experimental & Molecular Medicine · doi:10.1038/s12276-025-01456-7 · Cited by 4 · see citing works
“The S1 undergoes signi fi cant changes during the process of learning, as it not only represents simple low-level stimuli but also encompasses cognitive information necessary for learning, such as reward 17 , behavioral choice 18 and attention 19 .”
Panniello, M., Gillon, C. J., Maffulli, R., Celotto, M., Richards, B. A., Panzeri, S., & Kohl, M. M. (2024) · Cell Reports · doi:10.1016/j.celrep.2024.114244 · Cited by 6 · see citing works
“8 Insight into learning-related changes in cortical neuronal activity is key to understanding how the brain enables flexible behavior.”
BackgroundPanniello, M., Gillon, C. J., Maffulli, R., Celotto, M., Panzeri, S., Richards, B., & Kohl, M. (2022) · bioRxiv · doi:10.1101/2022.12.04.518156 · Cited by 2
Neural correlates of perceptual decision-making in the primary somatosensory cortex
Armstrong, A. G., & Vlasov, Y. (2026) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2514107123 · Cited by 1 · see citing works
“…21) sensory response (peak 2 in our case) is mostly perception-related, involving thalamo-cortico-thalamic loops (58), while “late” mostly decision-related response (19) (peak 4 in our experiments) reflect reverberation of neural activity (26, 30) in cortico-cortical S1 to S2 loops (10, 11, 22–24).”
Distinct Orbitofrontal Feedback Signals Shape Sensory Behavioral Strategies during Flexible LearningPreprint
Teutsch, J., Rao, R., Humphries, M. D., Maggi, S., & Banerjee, A. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.07.15.737076
“Consistent with this idea, population-level neural responses in S1 reorganize over learning 14,16 , suggesting that neuronal ensembles encode decision-related task variables that evolve over learning.”
InfluentialLi, H., Chen, J., Zhong, W., Lian, N., Huang, Y., Yao, L., Yin, P., Xu, Z., Qin, X., Tan, J., et al. (2026) · Nature Communications · doi:10.1038/s41467-026-70754-z · Cited by 3 · see citing works
Pandey, A., Kang, S., Pacchiarini, N., Wyszynska, H., Masseri, Z., O'Neill, J., Honey, R. C., & Fox, K. D. (2026) · European Journal of Neuroscience · doi:10.1111/ejn.70390 · Cited by 1 · see citing works
“However, while many neurons carry information about texture in S1, few appear specifically to code for it (Garion et al. 2014; Buetfering et al. 2022).”
Yang, H., Gui, Z., Gao, Y.-L., Yu, W., Dang, D., Zheng, N., Xu, S., & Wang, J. (2025) · Physiology and Behavior · doi:10.1016/j.physbeh.2025.115133 · Cited by 1 · see citing works
Neural Correlates of Perceptual Decision Making in Primary Somatosensory Cortex
Armstrong, A. G., & Vlasov, Y. D. (2025) · bioRxiv · doi:10.1101/2025.03.29.646003 · Cited by 1
“…that “early” 8,9 sensory response (peak 2 in our case) is mostly perception-related and may rely on thalamo-cortico-thalamic loops 56 , while “late” mostly decision-related response 7 (peak 4 in our experiments) reflect reverberation of neural activity 17,18 in cortico-cortical S1-S2 loops 10-12,”
Li, K., Liang, H., Qiu, J.-L., Zhang, X.-L., Cai, B.-B., Wang, D.-P., Zhang, D.-M., Lin, B., Han, H., Yang, G., et al. (2025) · Journal of NeuroEngineering and Rehabilitation · doi:10.1186/s12984-025-01655-3 · Cited by 1 · see citing works
“Additionally, by changing the frequency of optogenetic stimulation, the impact of different stimulus frequencies on mouse behavior was explored, finding that specific frequencies can effectively activate the target neural population and significantly affect mouse behavior [46].”
InfluentialBroschard, M. B., Kim, J., Halverson, H. E., Farley, S. J., & Freeman, J. H. (2025) · bioRxiv · doi:10.1101/2025.04.21.649785 · Cited by 1 · see citing works
“Buetfering et al., (2022) 44 found similar results in the mouse primary somatosensory cortex: some neurons correlated with the identity of the presented stimulus, whereas other neurons correlated with the mice’s decisions.”
Reach-to-Grasp and tactile discrimination task: A new task for the study of sensory-motor learning
Yoshinaga, Y., & Sato, N. (2024) · Behavioural Brain Research · doi:10.1016/j.bbr.2024.115007 · Cited by 2 · see citing works
Dai, J., & Sun, Q. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.08.30.555603 · Cited by 1 · see citing works
Learning-induced reorganization of neuronal subnetworks in the primary sensory cortexPreprint
Yang, Y., Shen, H., & Kwon, S. E. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.02.21.529414 · Cited by 1 · see citing works
Zhan, X., Tan, P., Tan, T., & Chen, C.-C. (2026) · Frontiers in Aging Neuroscience · doi:10.3389/fnagi.2026.1932616
“Texture discrimination in rodents depends on active whisking, tactile sampling, and cortical processing of texture-related information (von Heimendahl et al., 2007; Wu et al., 2013; Garion et al., 2014; Buetfering et al., 2022).”
Beyond single-cell sensory representations in cortex.
Willard, K. J., & Bruno, R. M. (2026) · Current Opinion in Neurobiology · doi:10.1016/j.conb.2026.103280
A stimulus–state geometry in somatosensory cortex reorganizes during inflammatory painPreprint
Schorscher-Petcu, A., Parkes, I., & Browne, L. E. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.08.11.744145
“In barrel cortex, activity carries decision-and outcome-related signals during tactile learning (Bale et al., 2021; Buetfering et al., 2022) and reflects body-state variables such as head and body configuration (Gantar et al., 2025; Palacio-Manzano et al., 2026).”
Raster photostimulation of large-scale neural populationsPreprint
LaFosse, P. K., Flickinger, D., Jaindl, G., Drinnenberg, A., Grødem, S., Lensjø, K. K., Ramakrishnan, C., Siverts, L., Zeng, H., Tasic, B., et al. (2026) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.64898/2026.04.21.719951
Lang, L., Zheng, C. Y., Blackwell, J. M., Camera, G. L., & Fontanini, A. (2025) · eLife · doi:10.7554/elife.109313
“Cortical circuits produce time-varying patterns of population and single-neuron activity that play a fundamental role in perceptual and behavioral processes ( Shuler and Bear, 2006 ; Guo et al., 2014b ; Buetfering et al., 2022 ).”
Pain as muse: How creative acts flourish in the shadow of struggle
Khalil, R., Frühholz, S., & Landry, M. (2026) · Neuroscience & Biobehavioral Reviews · doi:10.1016/j.neubiorev.2026.106650
Wang, H., Xiao, Y., Tang, W., Lai, K., Ma, Q., Zhao, X., Li, B., Shi, H., & He, H. (2026) · Nature Communications · doi:10.1038/s41467-026-69446-5
Lang, L., Zheng, C., Blackwell, J. M., Camera, G. L., & Fontanini, A. (2025) · bioRxiv · doi:10.1101/2025.10.06.680705
“Cortical circuits produce time-varying patterns of population and single neuron activity that play a fundamental role in perceptual and behavioral processes ( Shuler and Bear, 2006 ; Guo et al., 2014b ; Buetfering et al., 2022 ).”
Σεμελίδου, Ο., Gauvrit, T., Vandromme, C., Cornier, A., Saint-Jean, A., Feuvre, Y. L., Ginger, M. R., & Frick, A. (2026) · Advanced Science · doi:10.1002/advs.202519479 · Cited by 3 · see citing works
“Although decision-tuned neurons can emerge in L2/3 following extensive learning—such as during a whisker-based two-choice discrimination task [ 88 ]—this phenomenon appears task-dependent.”
Zhu, Z., Jiang, T., Jia, X., Wang, X., & Ren, M. (2025) · Neuroscience Research · doi:10.1016/j.neures.2025.104912
Perceptual choice and motor signals in mouse somatosensory cortexPreprint
Minamisawa, G., Dong, M., & O’Connor, D. H. (2024) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2024.12.06.627272
“Best-of-class evidence for identifying decision signals in cortex, however, requires precise gain-of-function manipulations combined with neural and behavioral analysis 24,25 .”
QuantFormer: Learning to Quantize for Neural Activity Forecasting in Mouse Visual Cortex
Calcagno, S., Kavasidis, I., Palazzo, S., Brondi, M., Sità, L., Turri, G., Giordano, D., Kostic, V. R., Fellin, T., Pontil, M., et al. (2024) · arXiv.org · doi:10.48550/arxiv.2412.07264
“For instance, studies have shown specific neuronal ensembles encoding stimulus features and behavior in the sensory cortex [23, 24], and in the motor cortex during motor programs [25].”
Discrete tactile feature comparison subprocess in human brain during a decision-making process
Lee, D. H., Kim, J. S., Ryun, S., & Chung, C. K. (2023) · Cortex · doi:10.1016/j.cortex.2023.11.004 · Cited by 1 · see citing works
Learning enhances representations of taste-guided decisions in the mouse gustatory insular cortex
Kogan, J. F., & Fontanini, A. (2023) · bioRxiv · doi:10.1101/2023.10.16.562605
“Much of the existing work on discrimination learning has been performed in the visual, auditory, somatosensory, and olfactory systems, 2,3,5,21,32–34 and comparatively little is known about discrimination learning in taste.”
Kimura, R. (2023) · Neuroscience Research · doi:10.1016/j.neures.2023.05.002
- Authors
- Buetfering, C., Zhang, Z., Pitsiani, M., Smallridge, J., Boven, E., McElligott, S., & Häusser, M.
- Affiliations
- University Medical Center of the Johannes Gutenberg University Mainz (DE)
- Stanford Medicine (US)
- University College London (GB)
- University of Zurich (CH)
- University of Bristol (GB)
- Corresponding author
- Buetfering, Christina
- Häusser, Michael
- Venue
- Nature Neuroscience
- Published
- 2022-08-30
- Volume
- 25
- Issue
- 9
- Pages
- 1225-1236
- ISSN
- 1097-6256 (ISSN-L)
- 1546-1726
- PMC
- PMC7613627
- OpenAlex
- W4293581938
- Cited by
- 59
- References
- 73
- Citations by year
- 2026: 13 · 2025: 18 · 2024: 15 · 2023: 13
- FWCI
- 5.93
- Type
- journal article
- Language
- en
- Update notices
- correction (2022-11-21)
- Open access
- green
- www.ncbi.nlm.nih.gov
- Locations
- Nature Neuroscience (publishedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- UCL Discovery (University College London) (submittedVersion)
- The HKU Scholars Hub (University of Hong Kong) (submittedVersion)
- Funders
- Wellcome Trust (201225/Z/16/Z)
- Topics
- Life Sciences › Neuroscience › Cognitive Neuroscience › Neural dynamics and brain function
- Life Sciences › Neuroscience › Cognitive Neuroscience › Visual perception and processing mechanisms
- Life Sciences › Neuroscience › Cognitive Neuroscience › Memory and Neural Mechanisms
- Keywords
- primary somatosensory cortex, texture discrimination, behavioral relevance, optogenetic perturbation, two-photon calcium imaging, layer 2/3 neurons
- MeSH terms
- Animals, Calcium, Learning, Neurons, Somatosensory Cortex, Mice, Optogenetics
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