Yet, does cortical processing play a role in this contact response? Progress on sensory control of motor programs is in need of sophisticated yet rapidly learned behavior paradigms,

perhaps involving object recogniton (Brecht et al., 1997). Finally, it is important to redress our focus on signaling in thalamocortical pathways to the exclusion of feedback through basal ganglia as well as subcortical loops formed by pontine-cerebellar and collicular pathways. The involvement of basal ganglia in whisking is largely uncharted, as only sensory responses in anesthetized animals have been reported (Pidoux et al., 2011). Cerebellar projection cells respond to vibrissa input (Bosman et al., 2010) and cerebellar output can affect the timing

in vM1 cortex (Lang et al., 2006), but again there is no composite Cabozantinib manufacturer understanding. The situation is more advanced for the case IWR-1 chemical structure of the superior colliculus, which receives direct vibrissa input via a trigeminotectal pathway (Killackey and Erzurumlu, 1981; Figure 3), indirect input via a corticotectal pathway through vS1 and vM1 cortices (Alloway et al., 2010, Miyashita et al., 1994 and Wise and Jones, 1977) and can drive whisking as well (Hemelt and Keller, 2008). Recording in awake free ranging animals show that cells in the colliculus respond to vibrissa touch (Cohen and Castro-Alamancos, 2010), while experiments that used fictive whisking Oxalosuccinic acid with anesthetized animals show that cells can respond to movement in the absence of contact (Bezdudnaya and Castro-Alamancos, 2011). It remains to be determined if the colliculus contains neurons that report touch conditioned on the position of the vibrissae and, if so, how these interact with the computation of touch in cortex. The vibrissa system is a particularly powerful proving ground to establish basic circuitry for sensorimotor control. The relatively stereotyped whisking

motion, the separation of sensory and motor signals on different nerves, and the accessibility of the system for electrophysiological study allow for fine experimental control. How general are these results? Essential aspects of sensation, such as balance with the vestibular system, seeing through the visual system, or touch through the somatosensory system, all make use of moving sensors and must solve an analogous problem to that discussed for the case of the rodent vibrissa sensorimotor system. This problem has been well studied for the case of vestibular control (Cullen et al., 2011 and Green and Angelaki, 2010), but has gained accelerating interest for the cases of other sensory modalities, in part from the advent of automated behavioral procedures (Dombeck et al., 2007 and Perkon et al., 2011), new tools to record intracellular (Lee et al., 2006) and multicellular (Sawinski et al., 2009) activity from behaving animals, and tools for targeted optical stimulation (Gradinaru et al., 2007).