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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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In the citing paper
“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]
Experimental — accuracy not yet measuredThe source backs part of the claim, not all of it — or backs it for a different population, condition or comparison.
In the citing paper
“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]
Experimental — accuracy not yet measuredThe cited source says what the claim attributes to it.
In the citing paper
“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]
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In the citing paper
“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]
Experimental — accuracy not yet measuredIn the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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 the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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
In the citing paper
“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 the citing paper
“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]
In the citing paper
“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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Metadata
- 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
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