If another set of objects retains their values for a long time, neurons in the caudate tail retain the sensitivity to the objects and, when any of the objects appear, react to it automatically

regardless of the outcome; this occurs quickly to many objects. Behaviorally, our inactivation experiments indicate that the caudate head and tail guide saccades aiming at valuable objects in different manners. The caudate head guides controlled saccades based on the flexible values (with immediate feedbacks), whereas the caudate tail guides automatic saccades based on the stable values (with no feedback). GSK126 cost Consistent with these results, neurons in these caudate subregions showed value-differential presaccadic activity but in different contexts: caudate head neurons during controlled saccades versus caudate tail neurons during automatic saccades. Notably, the inactivation of the caudate head as well as tail appeared to decrease the suppression of saccades to low-valued objects, rather than decrease the facilitation Ixazomib price of saccades to high-valued objects. This may be determined by the balance between the direct and indirect pathways (Hikosaka et al., 2000). How the balance might be controlled remains to be studied. The controlled saccades guided by caudate head and the automatic saccades guided by the caudate tail may be equivalent to

a well-documented dichotomy of behavior, such as goal-directed behavior versus skill (or habit) (Balleine and Dickinson, 1998), controlled versus automatic behavior (Schneider and Shiffrin, 1977), and System 2 versus System 1 (Evans, 2008). Several lines of evidence in human neuroimaging, human clinical, animal lesion, and physiological studies suggest that different regions of the basal ganglia are involved in controlled

versus automatic behavior (Ashby and Maddox, 2005, Balleine and O’Doherty, 2010, Hikosaka et al., 1999, Redgrave et al., 2010 and Yin and Knowlton, 2006). Human neuroimaging data suggest that subregions of the basal ganglia become active differentially depending on planning, skill acquisition, reward prediction, and feedbacks (Balleine and O’Doherty, 2010, Seger, 2008 and Wunderlich et al., 2012). Human patients with Parkinson’s disease are impaired in cognitive tasks that require flexible adaptations to oxyclozanide environmental changes, such as set shifting and value reversal (Brown and Marsden, 1990, Cools et al., 1984, Kehagia et al., 2010 and Lees and Smith, 1983). On the other hand, Parkinson’s disease patients are also impaired in probabilistic category learning tasks that require visual skills (Ashby and Maddox, 2005, Knowlton et al., 1996 and Shohamy et al., 2004). Patients with Huntington’s disease may show profound impairments in visual recognition (Lawrence et al., 1998), even early in the disease when neurodegeneration is detected mainly in the caudate tail (Vonsattel et al., 1985).