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Cerebellar granule cells encode the expectation of reward

机译：小脑颗粒细胞编码对奖励的期望

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The human brain contains approximately 60 billion cerebellar granule cells(1), 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(2-6). Although evidence suggests a role for the cerebellum in cognition(7-10), granule cells are known to encode only sensory(11-13) and motor(14) 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.
机译：人脑包含约600亿个小脑颗粒细胞（1），其数量超过所有其他脑神经元的总和。经典理论认为，大量多样的颗粒细胞可以高度详细地表示感觉运动情境，从而使下游的浦肯野细胞能够感知精细的情境变化（2-6）。尽管有证据表明小脑在认知中发挥作用（7-10），但已知颗粒细胞仅编码感觉（11-13）和运动（14）背景。在这里，在行为小鼠中使用双光子钙成像，我们显示颗粒细胞传达了有关奖励期望的信息。小鼠发起自愿的前肢运动以延迟糖水奖励。一些颗粒细胞优先响应奖励或奖励遗漏，而其他颗粒细胞选择性地编码奖励预期。奖励的反应不仅限于前肢运动，因为巴甫洛夫式的任务引起了类似的反应。与可预期的奖励相比，尽管奖励和舔responses反应相同，但意外奖励却引起了明显不同的颗粒细胞活动。在这两个任务中，奖赏信号遍布多个小脑小叶。在几天的学习中跟踪相同的颗粒细胞后发现，具有奖励预期反应的细胞从学习开始时就已经对奖励递送做出响应的细胞中出现了，而随着学习的进展，奖励遗漏的反应变得越来越强。在颗粒细胞中预测性，非感觉运动编码的发现是对这些神经元的当前理解的重大偏离，并且显着丰富了突触后浦肯野细胞可用的背景信息，对小脑的认知加工具有重要意义。

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  • 来源
    《Nature》 |2017年第7648期|96-100|共5页
  • 作者

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

  • 作者单位

    Stanford Univ, Dept Biol, Stanford, CA 94305 USA|Stanford Univ, Howard Hughes Med Inst, Stanford, CA 94305 USA;

    Stanford Univ, Dept Biol, Stanford, CA 94305 USA|Stanford Univ, Howard Hughes Med Inst, Stanford, CA 94305 USA|Stanford Univ, Dept Elect Engn, Stanford, CA 94305 USA;

    Stanford Univ, Dept Biol, Stanford, CA 94305 USA|Stanford Univ, Howard Hughes Med Inst, Stanford, CA 94305 USA;

    Stanford Univ, Dept Biol, Stanford, CA 94305 USA|Stanford Univ, Howard Hughes Med Inst, Stanford, CA 94305 USA|Stanford Univ, Dept Appl Phys, Stanford, CA 94305 USA;

    Stanford Univ, Dept Biol, Stanford, CA 94305 USA|Stanford Univ, Howard Hughes Med Inst, Stanford, CA 94305 USA;

  • 收录信息 美国《科学引文索引》(SCI);美国《工程索引》(EI);美国《生物学医学文摘》(MEDLINE);美国《化学文摘》(CA);
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