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Cerebellar granule cells encode the expectation of reward

Overview of attention for article published in Nature, March 2017
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  • In the top 5% of all research outputs scored by Altmetric
  • High Attention Score compared to outputs of the same age (99th percentile)
  • High Attention Score compared to outputs of the same age and source (81st percentile)


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584 Mendeley
3 CiteULike
Cerebellar granule cells encode the expectation of reward
Published in
Nature, March 2017
DOI 10.1038/nature21726
Pubmed ID

Mark J. Wagner, Tony Hyun Kim, Joan Savall, Mark J. Schnitzer, Liqun Luo


The human brain contains approximately 60 billion cerebellar granule cells, which outnumber all other brain neurons combined. Classical theories posit that a large, diverse population of granule cells allows for highly detailed representations of sensorimotor context, enabling downstream Purkinje cells to sense fine contextual changes. Although evidence suggests a role for the cerebellum in cognition, granule cells are known to encode only sensory and motor context. Here, using two-photon calcium imaging in behaving mice, we show that granule cells convey information about the expectation of reward. Mice initiated voluntary forelimb movements for delayed sugar-water reward. Some granule cells responded preferentially to reward or reward omission, whereas others selectively encoded reward anticipation. Reward responses were not restricted to forelimb movement, as a Pavlovian task evoked similar responses. Compared to predictable rewards, unexpected rewards elicited markedly different granule cell activity despite identical stimuli and licking responses. In both tasks, reward signals were widespread throughout multiple cerebellar lobules. Tracking the same granule cells over several days of learning revealed that cells with reward-anticipating responses emerged from those that responded at the start of learning to reward delivery, whereas reward-omission responses grew stronger as learning progressed. The discovery of predictive, non-sensorimotor encoding in granule cells is a major departure from the current understanding of these neurons and markedly enriches the contextual information available to postsynaptic Purkinje cells, with important implications for cognitive processing in the cerebellum.

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Mendeley readers

The data shown below were compiled from readership statistics for 584 Mendeley readers of this research output. Click here to see the associated Mendeley record.

Geographical breakdown

Country Count As %
United States 12 2%
United Kingdom 5 <1%
France 3 <1%
Japan 2 <1%
Germany 2 <1%
India 1 <1%
Israel 1 <1%
Austria 1 <1%
Mexico 1 <1%
Other 5 <1%
Unknown 551 94%

Demographic breakdown

Readers by professional status Count As %
Student > Ph. D. Student 163 28%
Researcher 130 22%
Student > Master 68 12%
Student > Bachelor 51 9%
Student > Doctoral Student 36 6%
Other 90 15%
Unknown 46 8%
Readers by discipline Count As %
Neuroscience 224 38%
Agricultural and Biological Sciences 153 26%
Psychology 52 9%
Medicine and Dentistry 35 6%
Engineering 22 4%
Other 39 7%
Unknown 59 10%

Attention Score in Context

This research output has an Altmetric Attention Score of 345. This is our high-level measure of the quality and quantity of online attention that it has received. This Attention Score, as well as the ranking and number of research outputs shown below, was calculated when the research output was last mentioned on 06 September 2019.
All research outputs
of 13,982,802 outputs
Outputs from Nature
of 71,030 outputs
Outputs of similar age
of 263,704 outputs
Outputs of similar age from Nature
of 833 outputs
Altmetric has tracked 13,982,802 research outputs across all sources so far. Compared to these this one has done particularly well and is in the 99th percentile: it's in the top 5% of all research outputs ever tracked by Altmetric.
So far Altmetric has tracked 71,030 research outputs from this source. They typically receive a lot more attention than average, with a mean Attention Score of 78.3. This one has done particularly well, scoring higher than 94% of its peers.
Older research outputs will score higher simply because they've had more time to accumulate mentions. To account for age we can compare this Altmetric Attention Score to the 263,704 tracked outputs that were published within six weeks on either side of this one in any source. This one has done particularly well, scoring higher than 99% of its contemporaries.
We're also able to compare this research output to 833 others from the same source and published within six weeks on either side of this one. This one has done well, scoring higher than 81% of its contemporaries.