21. Wang X, Chen G, Gao W, Ebner T (2009) Long-term potentiation of the responses to parallel fiber stimulation in mouse cerebellar cortex in vivo. Neuroscience 162: 713–722. 22. Cheron G, Gall D, Servais L, Dan B, Maex R, et al. (2004) Inactivation of calcium-binding protein genes induces 160 Hz oscillations in the cerebellar cortex of alert mice. J Neurosci 24: 434–441. 23. Eccles JC, Sasaki K, Strata P (1967) Interpretation of the potential fields generated in the cerebellar cortex by a mossy fibre volley. Exp Brain Res 3: 58–80. 24. Chen G, Popa LS, Wang X, Gao W, Barnes J, et al. (2009) Low-frequency oscillations in the cerebellar cortex of the tottering mouse. J Neurophysiol 101: 234–245. 25. Cheron G, Sausbier M, Sausbier U, Neuhuber W, Ruth P, et al. (2009) BK channels control cerebellar Purkinje and Golgi cell rhythmicity in vivo. PLoS One 4: e7991. 26. Bower JM, Woolston DC (1983) Congruence of spatial organization of tactile projections to granule cell and Purkinje cell layers of cerebellar hemispheres of the albino rat: vertical organization of cerebellar cortex. J Neurophysiol 49: 745–766. 27. Morissette J, Bower JM (1996) Contribution of somatosensory cortex to responses in the rat cerebellar granule cell layer following peripheral tactile stimulation. Exp Brain Res 109: 240–250. 28. Eccles JC, Llina´s RR, Sasaki K (1966) Parallel fibre stimulation and the responses induced thereby in the Purkinje cells of the cerebellum. Exp Brain Res 1: 17–39. 29. Llina´s RR (1982) General discussion: radial connectivity in the cerebellar cortex: a novel view regarding the functional organization of the molecular layer. In: Palay SL, Chan-Palay V, eds. The Cerebellum: New Vistas New York: Springer Verlag. pp 189–194. 30. Mapelli J, D’Angelo E (2007) The spatial organization of long-term synaptic plasticity at the input stage of cerebellum. J Neurosci 27: 1285–1296. 31. Eccles JC, Ito M, Szenta´gothai J (1967) The Cerebellum as a Neuronal Machine. New York: Springer-Verlag. 32. Armstrong DM, Drew T (1980) Responses in the posterior lobe of the rat cerebellum to electrical stimulation of cutaneous afferents to the snout. J Physiol 309: 357–374. 33. Sims RE, Hartell (2006) Differential susceptibility to synaptic plasticity reveals a functional specialization of ascending axon and parallel fiber synapses to cerebellar Purkinje cells. J Neurosci 26: 5153–5159. 34. Isope P, Barbour B (2002) Properties of unitary granule cell – Purkinje cell synapses in adult rat cerebellar slices. J Neurosci 22: 9668–9678. 35. Lu H, Esquivel AV, Bower JM (2009) 3D electron microscopic reconstruction of segments of rat cerebellar Purkinje cell dendrites receiving ascending and parallel fiber granule cell synaptic inputs. J Comp Neurol 514: 583–594. 36. Daniel H, Levenes C, Cre´pel F (1998) Cellular mechanisms of cerebellar LTD. Trends Neurosci 21: 401–407. 37. Gilbert PF, Thach WT (1977) Purkinje cell activity during motor learning. Brain Res 128: 309–328. Cerebellar LTD in Alert Mice 38. Ojakangas CL, Ebner TJ (1992) Purkinje cell complex and simple spike changes during a voluntary arm movement learning task in the monkey. J Neurophysiol 68: 2222–2236. 39. Medina JF, Lisberger SG (2008) Links from complex spikes to local plasticity and motor learning in the cerebellum of awake-behaving monkeys. Nat Neurosci 11: 1185–1192. 40. Llina´s RR, Lang EJ, Welsh JP (1997) The cerebellum, LTD, and memory: alternative views. Learn Mem 3: 445–455. 41. Llina´s RR, Sugimori M (1980) Electrophysiological properties of in vitro Purkinje cell dendrites in mammalian cerebellar slices. J Physiol 305: 197–213. 42. Eilers J, Augustine GJ, Konnerth A (1995) Subthreshold synaptic Ca2+ signalling in fine dendrites and spines of cerebellar Purkinje neurons. Nature 373: 155–158. 43. Hartell (1996) Strong activation of parallel fibers produces localized calcium transients and a form of LTD that spreads to distant synapses. Neuron 16: 601–610. 44. Carvell GE, Simons DJ (1990) Biometric analyses of vibrissal tactile discrimination in the rat. J Neurosci 10: 2638–2648. 45. Me´gevand P, Troncoso E, Quairiaux C, Muller D, Michel CM, et al. (2009) Long-term plasticity in mouse sensorimotor circuits after rhythmic whisker stimulation. J Neurosci 29: 5326–5335. 46. Bosman LW, Koekkoek SK, Shapiro J, Rijken BF, Zandstra F, et al. (2010) Encoding of whisker input by cerebellar Purkinje cells. J Physiol 588: 3757–3783. 47. Ebner TJ, Yu QX, Bloedel JR (1983) Increase in Purkinje cell gain associated with naturally activated climbing fiber input. J Neurophysiol 50: 205–219. 48. Holtzman T, Rajapaksa T, Mostofi A, Edgley SA (2006) Different responses of rat cerebellar Purkinje cells and Golgi cells evoked by widespread convergent sensory inputs. J Physiol (Lond) 574: 491–507. 49. Brunel N, Hakim V, Isope P, Nadal JP, Barbour B (2004) Optimal information storage and the distribution of synaptic weights: perceptron versus Purkinje cell. Neuron 43: 745–757. 50. Mittmann W, Ha¨usser M (2007) Linking synaptic plasticity and spike output at excitatory and inhibitory synapses onto cerebellar Purkinje cells. J Neurosci 27: 5559–5570. 51. Chen C, Thompson RF (1995) Temporal specificity of long-term depression in parallel fiber-Purkinje synapses in rat cerebellar slice. Learn Mem 2: 185–198. 52. Ito M (2002) The molecular organization of cerebellar long-term depression. Nat Rev Neurosci 3: 896–902. 53. Poulet JF, Petersen CC (2008) Internal brain state regulates membrane potential synchrony in barrel cortex of behaving mice. Nature 454: 881–885. 54. Wang SS, Denk W, Ha¨usser M (2000) Coincidence detection in single dendritic spines mediated by calcium release. Nat Neurosci 3: 1266–1273. 55. Steuber V, Mittmann W, Hoebeek FE, Silver RA, De Zeeuw CI (2007) Cerebellar LTD and pattern recognition by Purkinje cells. Neuron 54: 121–136. 56. D’Angelo E, De Zeeuw CI (2009) Timing and plasticity in the cerebellum: focus on the granular layer. Trends Neurosci 32: 30–40. PLoS ONE | www.plosone.org 13 April 2012 | Volume 7 | Issue 4 | e36184