Arousal and Locomotion Make Distinct Contributions to Cortical Activity Patterns and Visual Encoding
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Abstract
Spontaneous and sensory-evoked cortical activity is highly state-dependent, yet relatively little is known about transitions between distinct waking states. Patterns of activity in mouse V1 differ dramatically between quiescence and locomotion, but this difference could be explained by either motor feedback or a change in arousal levels. We recorded single cells and local field potentials from area V1 in mice head-fixed on a running wheel and monitored pupil diameter to assay arousal. Using naturally occurring and induced state transitions, we dissociated arousal and locomotion effects in V1. Arousal suppressed spontaneous firing and strongly altered the temporal patterning of population activity. Moreover, heightened arousal increased the signal-to-noise ratio of visual responses and reduced noise correlations. In contrast, increased firing in anticipation of and during movement was attributable to locomotion effects. Our findings suggest complementary roles of arousal and locomotion in promoting functional flexibility in cortical circuits.
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Locomotion is a homogeneous behavioral state tightly coupled to arousal.
“This is because locomotion—where place cell activity has been classically characterized—is a homogeneous behavioral state, tightly coupled to arousal32–35.”
From Place and behavioral modulation of hippocampal neurons during immobility (2026) · cites this paper as [34]
In Results ('Dissociating locomotion and arousal'; page not provided), pupil diameter was consistently correlated with locomotion transitions and remained elevated during running, supporting a close association with arousal. The same passage reports elevated arousal without locomotion, and the paper does not establish that locomotion is a homogeneous behavioral state. (LLM, conf=0.84)
Experimental — accuracy not yet measuredThe onset of locomotion, as well as vestibular activation, are correlated with pupil dilations and brain state transitions.
“Indeed, the onset of locomotion, as well as vestibular activation, are correlated with pupil dilations and brain state transitions32,34,36.”
From Place and behavioral modulation of hippocampal neurons during immobility (2026) · cites this paper as [34]
The Results sections ‘Dissociating locomotion and arousal’ and ‘LFP modulation by behavioral state transition’ report pupil dilation around locomotion onset and associated changes in cortical activity. The paper does not report vestibular activation or link it to pupil dilation or brain-state transitions; searches for vestibular activation and stimulation found no such passage. Page numbers were unavailable in our copy. (LLM, conf=0.88)
Experimental — accuracy not yet measuredDuring running, the behavioral state is more homogeneous and likely corresponds to a high-arousal state compared to immobility, where the internal state is more variable and spontaneously fluctuating.
“These data indicate that during running, the behavioral state is more homogeneous (and likely corresponding to a high-arousal state34–36) compared to immobility, where the internal state is more variable and spontaneously fluctuating.”
From Place and behavioral modulation of hippocampal neurons during immobility (2026) · cites this paper as [34]
Elman RNNs can be viewed as an approximation to continuous time firing rate models of neural circuits.
“We examined Elman RNNs (“vanilla” RNNs), which are perhaps the simplest RNN architecture capable of theoretically representing any nonlinear dynamical system, 52 and which can be viewed as an approximation to continuous time firing rate models of neural circuits.53”
From Remapping in a recurrent neural network model of navigation and context inference (2023) · cites this paper as [53]
Spontaneous and sensory-evoked activity varies continuously and rapidly during wakefulness.
“This work has revealed that spontaneous and sensory-evoked activity varies continuously and rapidly during wakefulness.”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Even without movement, increases in pupil diameter are associated with increased cortical activation and suppression of low-frequency rhythms.
“Even in the absence of movement, increases in pupil diameter (dilation) were found to be associated with increases in cortical activation and suppression of low frequency rhythms.”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Initiation of locomotion is preceded by a substantial increase in pupil diameter, concurrent with reduced low-frequency cortical LFP synchronization and reduced hippocampal sharp wave and ripple oscillations.
“Interestingly, the initiation of locomotion is preceded by a substantial increase in pupil diameter (Figures 2–4), concurrent with the reduction of low-frequency cortical local field potential synchronization and reduced hippocampal sharp wave and ripple oscillations.”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
The transition from sitting quietly to exploratory behaviors such as whisking and locomotion is intertwined with changes in arousal tracked by pupillometry.
“The transition from sitting quietly to exploratory behaviors (whisking and locomotion) is intertwined with changes in arousal tracked by pupillometry.”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Low-frequency cortical activity is enhanced during pupillary constriction, especially below a critical pupil diameter.
“Likewise, low-frequency cortical activity is enhanced during pupillary constriction, especially below a critical level of pupil diameter.”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
The relationship between pupil diameter and cortical network activity has been observed in visual, somatosensory, and auditory cortical areas.
“has been observed in visual, somatosensory, and auditory cortical areas”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Simultaneous intracellular membrane potential and LFP recordings with pupil diameter in head-fixed mice revealed a marked relationship between pupil size, low-frequency (2-10 Hz) membrane potential/LFP fluctuations, and exploratory behaviors such as whisking and locomotion.
“Recently, intracellular membrane potential (Vm) and local field potential (LFP) recordings from cortical neurons collected simultaneously with pupil diameter in head-fixed, spontaneously locomoting or whisking mice revealed a marked relationship between 1) pupil size, 2) low frequency (2–10 Hz) fluctuations in membrane potential/LFP, and 3) exploratory behaviors (whisking and locomotion)”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Effects of locomotion on spontaneous discharge rates of presumed pyramidal neurons in other layers are more diverse.
“However, the effects of locomotion on the spontaneous discharge rates of presumed pyramidal neurons recorded in other layers is more diverse”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
A gradual decrease in pupil diameter after locomotion is associated with increased prevalence of low-frequency oscillations.
“This gradual decrease in pupil diameter is associated with an increase in the prevalence of low-frequency oscillations”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Eliciting arousal by air puff in a quiescent mouse significantly increases pupil diameter and strongly suppresses slow rhythmic activity.
“eliciting an increase in arousal through the delivery of an air puff to a quiescent mouse results in a significant increase in pupil diameter and strong suppression of slow rhythmic activity”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Applying the same arousing stimulus during locomotion, when average neuronal activity is elevated, does not suppress firing, whereas a transition to locomotion following induced arousal increases neuronal discharge.
“Application of the same arousing stimulus during locomotion, when average neuronal activity is elevated, does not result in a suppression of firing, whereas a transition to locomotion that occurs following induced arousal results in an increase in neuronal discharge”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Air puff-induced arousal elicits an average decrease in discharge rate of many cortical neurons, unlike the increase in neuronal activity prior to movement.
“However, in contrast with the increase in neuronal activity period prior to movement, air puff-induced arousal elicits an average decrease in the discharge rate of many cortical neurons.”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Following cessation of locomotion, head-fixed mice typically transition from high to moderate or low arousal over seconds or longer, as measured by pupil diameter.
“Following the cessation of locomotion, head-fixed mice typically exhibit a transition from high arousal to moderate or low arousal, as measured by pupil diameter, over a period of seconds or longer.”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Even in the quiescent state without overt movement, pupil dilation is associated with suppression of low-frequency and enhancement of gamma-frequency cortical network activities, occurring as rapid switching or graded continuous change.
“As described above, even in the quiescent state devoid of overt movements, pupil dilation is associated with suppression of low-frequency, and enhancement of gamma-frequency, cortical network activities that can occur as either a rapid switching (Figure 2A) or as a graded and continuous change”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
A subset of neurons shows a strong increase in evoked response with causally induced transition from low to high arousal.
“and a subset of neurons show a strong increase in evoked response with causally (i.e. airpuff) induced transition from low to high arousal”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Behavioral state is an important determinant of both the temporal precision of sensory responses and gamma-band synchronization.
“Behavioral state is an important determinant of both the temporal precision of sensory responses and gamma-band synchronization.”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Spontaneous or causally induced increases in arousal are associated with a monotonic increase in gamma-band synchronization in LFP and relative membrane potential power, and in spike-LFP phase locking.
“Spontaneous or causally induced increases in arousal are associated with a monotonic increase in gamma-band synchronization both in terms of local field potential and relative membrane potential power (Lee et al., 2014; Lima et al., 2011; McGinley et al., 2015; Reimer et al., 2014; Vinck et al., 2015), and spike-local field potential phase locking”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
In mouse visual cortex, the signal-to-noise ratio is small at low arousal and increases at high arousal and during locomotion.
“In mouse visual cortex, the S/N is small at low arousal levels, and increases at high arousal levels and during locomotion”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Pupil dilations during quiet wakefulness correspond to increased visual responsiveness and enhanced selectivity.
“Pupil dilations during quiet wakefulness also correspond to an increase in visual responsiveness along with an enhancement of selectivity”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Noise correlations within wakefulness are, to a significant degree, explained by fluctuations in arousal as indexed by pupil diameter.
“Recent work also shows that noise correlations within wakefulness are, to a significant degree, explained by fluctuations in arousal as indexed by pupil diameter (Reimer et al., 2014; Vinck et al., 2015).”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Pooling different behavioral states together in the analysis of noise correlations introduces shared variability in neuronal responses, over-estimating their magnitude.
“Moreover, pooling different behavioral states together in the analysis of noise correlations introduces shared variability in neuronal responses, over-estimating their magnitude (Vinck et al., 2015).”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Within wakefulness, noise correlations are lower during locomotion than quiescence.
“Within wakefulness, noise correlations are lower during locomotion than quiescence (Figure 6B) (Erisken et al., 2014; Vinck et al., 2015).”
From Waking state: rapid variations modulate neural and behavioral responses (2015) · cites this paper as Vinck et al., 2015
Arousal and locomotion are associated with reduced and enhanced spontaneous firing rates in mouse primary visual cortex (V1), respectively, indicating that mechanisms by which gain is enhanced during these two behavioural states may differ.
“Arousal and locomotion are associated with reduced and enhanced spontaneous firing rates in the mouse primary visual cortex (V1)33, respectively, indicating that mechanisms by which gain is enhanced during these two behavioural states may differ.”
From Mechanisms underlying gain modulation in the cortex (2020) · cites this paper as [33]
In visual cortex, the onset of arousal (as measured by pupil diameter) and locomotion increase neuronal gain.
“In the visual cortex, the onset of arousal (as measured by pupil diameter) and locomotion increase neuronal gain32,33,36,37.”
From Mechanisms underlying gain modulation in the cortex (2020) · cites this paper as [33]
Gain modulation mechanisms are robustly engaged by changes in internal state.
“Gain modulation mechanisms are also robustly engaged by changes in internal state32–35.”
From Mechanisms underlying gain modulation in the cortex (2020) · cites this paper as [33]
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- Authors
- Vinck, M., Batista-Brito, R., Knoblich, U., & Cardin, J. A.
- Affiliations
- Yale University (US)
- Corresponding author
- Cardin, Jessica A.
- Venue
- Neuron
- Publisher
- Elsevier BV
- Published
- 2015-05
- Volume
- 86
- Issue
- 3
- Pages
- 740-754
- ISSN
- 0896-6273 (ISSN-L)
- PMC
- PMC4425590
- OpenAlex
- W2086338669
- Cited by
- 819
- References
- 63
- Citations by year
- 2026: 71 · 2025: 78 · 2024: 108 · 2023: 92 · 2022: 89 · 2021: 76 · 2020: 71 · 2019: 72 · 2018: 57 · 2017: 52 · 2016: 41 · 2015: 12
- FWCI
- 32.49
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Licence
- Locations
- Neuron (publishedVersion)
- bioRxiv (Cold Spring Harbor Laboratory) (acceptedVersion)
- PubMed (publishedVersion)
- PubMed Central (acceptedVersion)
- Funders
- McKnight Foundation
- National Alliance for Research on Schizophrenia and Depression
- National Institutes of Health
- National Eye Institute (EY022951, R00EY018407, R01EY022951, P30EY026878)
- 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 › Sleep and Wakefulness Research
- Keywords
- arousal, locomotion, primary visual cortex, spontaneous cortical activity, population activity, signal-to-noise ratio, noise correlation, pupil size, running wheel, motor feedback, visual encoding, mice
- MeSH terms
- Action Potentials, Animals, Arousal, Computer Simulation, Fourier Analysis, Locomotion, Male, Pupil, Visual Cortex, Visual Pathways, Wakefulness, Signal Detection, Psychological, Mice
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
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