< Curriculum Vitae > Okihide Hikosaka, M.D., Ph.D. Senior Researcher Chief, Section of Neuronal Networks Laboratory of Sensorimotor Research National Eye Institute, National Institutes of Health Bldg. 49, Rm. 2A50 Bethesda, MD 20892, U.S.A. TEL: 301-402-7959 FAX: 301-402-0511 e-mail: oh@lsr.nei.nih.gov Joint appointment: Cellular Neurobiology Branch Division of Intramural Research National Institute on Drug Abuse (NIDA), National Institutes of Health 1973 M.D. at University of Tokyo 1978 Ph.D. (Basic Medicine) at University of Tokyo 1978 Lecturer at Toho University School of Medicine 1979 Assistant professor at Toho University School of Medicine 1979-1982 Visiting Scientist at National Eye Institute 1983 Associate professor at Toho University School of Medicine 1988 Professor at National Institute of Physiological Sciences in Okazaki 1993-2002 Professor at Juntendo University School of Medicine 2002- Senior Investigator at Laboratory of Sensorimotor Research, NEI, NIH Awards and Honors: Tsukahara Memorial Award, 1989 Symposium Lecture, Society for Neuroscience, 19th Annual Meeting, 1989 Special Lecture, Society for Neuroscience, 24th Annual Meeting, 1994 Opening Lecture, International Neurootological & Equilibriometric Society, XXIInd Congress, 1995 Plenary Lecture, The Fifth International Conference on Neural Information Processing, 1998 Tokizane Toshihiko Memorial Award, 1999 Keynote Lecture, Organization for Human Brain Mapping, 9th annual meeting, 2003 Symposium Lecture, Organizer, Society for Neuroscience, 38th Annual Meeting, 2008 Symposium Lecture, International Union of Physiological Sciences, 2009 Presidential Special Lecture, Society for Neuroscience, 40th Annual Meeting, 2010 Keynote Lecture, Janelia Farm Conference, 2011 Member, American Academy of Arts & Sciences, 2011 Inaugural Phillip A. Sharp Lecture in Neural Circuits. 2012 Keynote Lecture, 11th Triennial meeting of the International Basal Ganglia Society, 2013 Symposium Lecture, Society for Neuroscience, Annual Meeting, 2014 Plenary lecture, French Neuroscience Society Meeting, 2015 The Torsten Wiesel Lecture (Plenary lecture), IBRO World Congress, 2015 Plenary lecture, Japan Neuroscience Society Meeting, 2015 Golden Brain Award, 2015

Research areas: Control of eye movements Functions of the basal ganglia Neural mechanisms of motivation Neural mechanisms of procedural learning Neural mechanisms of skill Neural mechanisms of attention Professional Service Editorial board Trends in Cognitive Sciences (2011-) Current Opinion in Neurobiology (2008-) Neuroscience Research Neural Networks Journal of Neuroscience

< Bibliography > < Original papers > Kim HF, Ghazizadeh A, Hikosaka O (2015) Dopamine neurons encoding long-term memory of object value for habitual behavior. Cell 163: 1165-1175. Monosov I, Leopold DA, Hikosaka O (2015) Neurons in the primate medial basal forebrain signal combined information about reward uncertainty, value, and punishment anticipation. J Neurosci 35: 7443-7459. Yasuda M, Hikosaka O (2015) Functional territories in primate substantia nigra pars reticulata separately signaling stable and flexible values. J Neurophysiol 113: 1681–1696. Kim HF, Ghazizadeh A, Hikosaka O (2014) Separate groups of dopamine neurons innervate caudate head and tail encoding flexible and stable value memories. Front Neuroanat 8:120. Hong S, Hikosaka O (2014) Pedunculopontine tegmental nucleus neurons provide reward, sensorimotor, and alerting signals to midbrain dopamine neurons. Neuroscience 282C: 139-155. Hong S, Hikosaka O (2013) Diverse sources of reward value signals in the basal ganglia nuclei transmitted to the lateral habenula in the monkey. Front Hum Neurosci 7: 778. Kim HF, Hikosaka O (2013) Distinct basal ganglia circuits controlling behaviors guided by flexible and stable values. Neuron 79: 1001-1010. Yamamoto S, Kim HF, Hikosaka O (2013) Reward value-contingent changes of visual responses in the primate caudate tail associated with a visuomotor skill. J Neurosci 33: 11227-11238. Monosov I, Hikosaka O (2013) Selective and graded coding of reward uncertainty by neurons in the primate anterodorsal septal region. Nat Neurosci 16:756-762. Daye PM, Monosov I, Hikosaka O, Leopold DA, Optican LM (2013) pyElectrode: An open-source tool using structural MRI for electrode positioning and neuron mapping. J Neurosci Methods 213: 123-131. Tachibana Y, Hikosaka O (2012) The primate ventral pallidum encodes expected reward value and regulates motor action. Neuron 76: 826-837. Yasuda M, Yamamoto S, Hikosaka O (2012) Robust representation of stable object values in the oculomotor basal ganglia. J Neurosci 32: 16917-16932.

Yamamoto S, Monosov IE, Yasuda M, Hikosaka, O (2012) What and where information in the caudate tail guides saccades to visual objects. J Neurosci 32: 11005-11016 Monosov IE, Hikosaka O (2012) Regionally distinct processing of rewards and punishments by the primate ventromedial prefrontal cortex. J Neurosci 32: 10318-10330. Matsumoto M, Hikosaka O (2011) Electrical Stimulation of the Primate Lateral Habenula Suppresses Saccadic Eye Movement through a Learning Mechanism. PLoS One 6:e26701. Bromberg-Martin ES, Hikosaka O (2011) Lateral habenula neurons signal errors in the prediction of reward information. Nat Neurosci 14:1209-1216. Hong S, Jhou TC, Smith M, Saleem KS, Hikosaka O (2011) Negative reward signals from the lateral habenula to dopamine neurons are mediated by rostromedial tegmental nucleus in primates. J Neurosci 31:11457-11471. Hong S, Hikosaka O (2011) Dopamine-mediated learning and switching in cortico-striatal circuit explain behavioral changes in reinforcement learning. Front Behav Neurosci 5:15. Terao Y, Fukuda H, Yugeta A, Hikosaka O, Nomura Y, Segawa M, Hanajima R, Tsuji S, Ugawa Y (2011) Initiation and inhibitory control of saccades with the progression of Parkinson's disease - Changes in three major drives converging on the superior colliculus. Neuropsychologia 49:1794-1806. Bromberg-Martin ES, Matsumoto M, Nakahara H, Hikosaka O (2010d) Multiple timescales of memory in lateral habenula and dopamine neurons. Neuron 67:499-510. Bromberg-Martin ES, Matsumoto M, Hikosaka O (2010b) Distinct tonic and phasic anticipatory activity in lateral habenula and dopamine neurons. Neuron 67:144-155. Bromberg-Martin ES, Matsumoto M, Hong S, Hikosaka O (2010c) A pallidus-habenula-dopamine pathway signals inferred stimulus values. J Neurophysiol 104:1068-1076. Bromberg-Martin ES, Hikosaka O, Nakamura K (2010a) Coding of task reward value in the dorsal raphe nucleus. J Neurosci 30:6262-6272. Yugeta A, Terao Y, Fukuda H, Hikosaka O, Yokochi F, Okiyama R, Taniguchi M, Takahashi H, Hamada I, Hanajima R, Ugawa Y (2010) Effects of STN stimulation on the initiation and inhibition of saccade in Parkinson disease. Neurology 74:743-748. Bromberg-Martin ES, Hikosaka O (2009) Midbrain dopamine neurons signal preference for advance information about upcoming rewards. Neuron 63:119-126.

Matsumoto M, Hikosaka O (2009) Two types of dopamine neuron distinctly convey positive and negative motivational signals. Nature 459:837-841. Matsumoto M, Hikosaka O (2009) Representation of negative motivational value in the primate lateral habenula. Nat Neurosci 12:77-84. Hong S, Hikosaka O (2008) The globus pallidus sends reward-related signals to the lateral habenula. Neuron 60:720-729. Bissmarck F, Nakahara H, Doya K, Hikosaka O (2008) Combining Modalities with Different Latencies for Optimal Motor Control. J Cogn Neurosci 20:1966-1979. Isoda M, Hikosaka O (2008) A neural correlate of motivational conflict in the superior colliculus of the macaque. J Neurophysiol 100:1332-1342. Hikosaka O, Isoda M (2008) Brain mechanisms for switching from automatic to controlled eye movements. Prog Brain Res 171:375-382. Matsumoto M, Hikosaka O (2008) Negative motivational control of saccadic eye movement by the lateral habenula. Prog Brain Res 171:399-402. Isoda M, Hikosaka O (2008) Role for subthalamic nucleus neurons in switching from automatic to controlled eye movement. J Neurosci 28:7209-7218. Nakamura K, Matsumoto M, Hikosaka O (2008) Reward-dependent modulation of neuronal activity in the primate dorsal raphe nucleus. J Neurosci 28:5331-5343. Ikeda T, Hikosaka O (2007) Positive and negative modulation of motor response in primate superior colliculus by reward expectation. J Neurophysiol 98: 3163–3170 Matsumoto M, Hikosaka O (2007) Lateral habenula as a source of negative reward signals in dopamine neurons. Nature 447:1111-1115. Isoda M, Hikosaka O (2007) Switching from automatic to controlled action by monkey medial frontal cortex. Nat Neurosci 10:240-248. Kobayashi S, Kawagoe R, Takikawa Y, Koizumi M, Sakagami M, Hikosaka O (2007) Functional differences between macaque prefrontal cortex and caudate nucleus during eye movements with and without reward. Exp Brain Res 176:341-355. Ding L, Hikosaka O (2007) Temporal Development of Asymmetric Reward-Induced Bias in Macaques. J Neurophysiol 97:57-61. Nakamura K, Hikosaka O (2006) Facilitation of saccadic eye movements by postsaccadic

electrical stimulation in the primate caudate. J Neurosci 26:12885-12895. Nakahara H, Nakamura K, Hikosaka O (2006) Extended LATER model can account for trial-by-trial variability of both pre- and post-processes. Neural Netw 19:1027-1046. Kobayashi S, Nomoto K, Watanabe M, Hikosaka O, Schultz W, Sakagami M (2006) Influences of rewarding and aversive outcomes on activity in macaque lateral prefrontal cortex. Neuron 51:861-870. Watanabe K, Ikeda H, Hikosaka O (2006) Effects of explicit knowledge of workspace rotation in visuomotor sequence learning. Exp Brain Res 174:673-678. Ding L, Hikosaka O (2006) Comparison of reward modulation in the frontal eye field and caudate of the macaque. J Neurosci 26:6695-6703. Nakamura K, Hikosaka O (2006) Role of dopamine in the primate caudate nucleus in reward modulation of saccades. J Neurosci 26:5360-5369. Watanabe K, Hikosaka O (2005) Immediate changes in anticipatory activity of caudate neurons associated with reversal of position-reward contingency. J Neurophysiol 94:1879-1887. Takikawa Y, Kawagoe R, Hikosaka O (2004) A possible role of midbrain dopamine neurons in short- and long-term adaptation of saccades to position-reward mapping. J Neurophysiol 92:2520-2529. Kawagoe R, Takikawa Y, Hikosaka O (2004) Reward-predicting activity of dopamine and caudate neurons - a possible mechanism of motivational control of saccadic eye movement. J Neurophysiol 91:1013-1024. Nakahara H, Itoh H, Kawagoe R, Takikawa Y, Hikosaka O (2004) Dopamine neurons can represent context-dependent prediction error. Neuron 41:269-280. Nomura Y, Fukuda H, Terao Y, Hikosaka O, Segawa M (2003) Abnormalities of voluntary saccades in Gilles de la Tourette's syndrome: pathophysiological consideration. Brain Dev 25 Suppl 1:S48-54. Watanabe K, Lauwereyns J, Hikosaka O (2003) Neural correlates of rewarded and unrewarded eye movements in the primate caudate nucleus. J Neurosci 23:10052-10057. Ikeda T, Hikosaka O (2003). Reward-dependent gain and bias of visual responses in primate superior colliculus. Neuron 39: 693-700. Sakai K, Kitaguchi K, Hikosaka O (2003) Chunking during human visuomotor sequence

learning. Exp Brain Res 152:229-242. Watanabe K, Lauwereyns J, Hikosaka O (2003) Effects of motivational conflicts on visually elicited saccades in monkeys. Exp Brain Res 152:361-367. Itoh H, Nakahara H, Hikosaka O, Kawagoe R, Takikawa Y, Aihara K (2003) Correlation of Primate Caudate Neural Activity and Saccade Parameters in Reward-Oriented Behavior. J Neurophysiol 89:1774-1783. Hikosaka O, Rand MK, Nakamura K, Miyachi S, Kitaguchi K, Sakai K, Lu X, Shimo Y (2002b) Long-term retention of motor skill in macaque monkeys and humans. Exp Brain Res 147:494-504. Miyachi S, Hikosaka O, Lu X (2002) Differential activation of monkey striatal neurons in the early and late stages of procedural learning. Exp Brain Res 146:122-126. Lauwereyns J, Watanabe K, Coe B, Hikosaka O (2002) A neural correlate of response bias in monkey caudate nucleus. Nature 418:413-417. Coe B, Tomihara K, Matsuzawa M, Hikosaka O (2002) Visual and Anticipatory Bias in Three Cortical Eye Fields of the Monkey during an Adaptive Decision-Making Task. J Neurosci 22:5081-5090. Lu X, Matsuzawa M, Hikosaka O (2002) A neural correlate of oculomotor sequences in supplementary eye field. Neuron 34:317-325. Nakahara H, Amari Si S, Hikosaka O (2002) Self-organization in the basal ganglia with modulation of reinforcement signals. Neural Comput 14:819-844. Sato M, Hikosaka O (2002) Role of primate substantia nigra pars reticulata in reward-oriented saccadic eye movement. J Neurosci 22:2363-2373. Kobayashi S, Lauwereyns J, Koizumi M, Sakagami M, Hikosaka O (2002) Influence of reward expectation on visuospatial processing in macaque lateral prefrontal cortex. J Neurophysiol 87:1488-1498. Takikawa Y, Kawagoe R, Hikosaka O (2002a) Reward-dependent spatial selectivity of anticipatory activity in monkey caudate neurons. Journal of Neurophysiology 87:508-515. Lauwereyns J, Takikawa Y, Kawagoe R, Kobayashi S, Koizumi M, Coe B, Sakagami M, Hikosaka O (2002) Feature-based anticipation of cues that predict reward in monkey caudate nucleus. Neuron 33:463-473. Takikawa Y, Kawagoe R, Itoh H, Nakahara H, Hikosaka O (2002b) Modulation of saccadic

eye movements by predicted reward outcome. Experimental Brain Research 142:284-291. Nakahara H, Doya K, Hikosaka O (2001) Parallel cortico-bsal ganglia mechanisms for acquisition and execution of visuomotor seqeunces - A computational approach. Journal of Cognitive Neuroscience 13: 1-22. Sakagami M, Tsutsui K-I, Lauwereyns J, Koizumi M, Kobayashi S, Hikosaka O (2001) A code for behavioral inhibition on the basis of color, but not motion, in ventrolateral prefrontal cortex of macaque monkey. J Neurosci 21: 4801-4808 Lauwereyns J, Sakagami M, Tsutsui K-I, Kobayashi S, Koizumi M, Hikosaka O (2001) Responses to task-irrelevant visual features by primate prefrontal neurons. J Neurophysiol 86: 2001-2010. Shimo Y, Hikosaka O (2001) Role of tonically active neurons in primate caudate in reward-oriented saccadic eye movement. J Neurosci 21: 7804-7814. Lauwereyns J, Koizumi M, Sakagami M, Hikosaka O, Kobayashi S, Tsutsui K (2000) Interference from irrelevant features on visual discrimination by macaques (Macaca fuscata) : a behavioral analogue of human stroop effect. Journal of Experimental Psychology: Animal Behavior Processes 26: 352-357. Rand MK, Hikosaka O, Miyachi S, Lu X, Nakamura K, Kitaguchi K, Shimo Y (2000) Characteristics of sequential movements during early learning period in monkeys. Exp Brain Res 131: 293-304. Sakai K, Hikosaka O, Takino R, Miyauchi S, Nielsen M, Tamada T (2000) What and when: parallel and convergent processing in motor control. J Neurosci 20: 2691-2700. Sakai K, Hikosaka O, Miyauchi S, Sasaki Y, Fujimaki N, Pütz B (1999) Pre-SMA activation during sequence learning reflects visuo-motor association. J Neurosci 19:RC1: 1-6. Nakamura K, Sakai K, Hikosaka O (1999) Effects of local inactivation of monkey medial frontal cortex in learning of sequential procedures. J Neurophysiol 82: 1063-1068. Sakai K, Hikosaka O, Miyauchi S, Takino R, Tamada T, Iwata NK, Nielsen M (1999) Neural representation of a rhythm depends on its interval ratio. J Neurosci 19: 10074-10081. Lu X, Hikosaka O, Miyachi S (1998) Role of monkey cerebellar nuclei in skill for sequential movement. J Neurophysiol 79: 2245-2254. Nakamura K, Sakai K, Hikosaka O (1998) Neuronal activity in medial frontal cortex during learning of sequential procedures. J Neurophysiol 80: 2671-2687.

Kawagoe R, Takikawa Y, Hikosaka O (1998) Expectation of reward modulates cognitive signals in the basal ganglia. Nat Neurosci 5: 411-416. Rand MK, Hikosaka O, Miyachi S, Lu X, Miyashita K (1998) Characteristics of a long-term procedural skill in the monkey. Exp Brain Res 118: 293-297. Sakai K, Hikosaka O, Miyauchi S, Takino R, Sasaki Y, Pütz B (1998a) Transition of brain activation from frontal to parietal areas in visuo-motor sequence learning. J Neurosci 18: 1827-1840. Sakai K, Takino R, Hikosaka O, Miyauchi S, Sasaki Y, Pütz B, Fujimaki N (1998b) Separate cerebellar areas for motor control. NeuroReport 9: 2359-2363. Takikawa Y, Kawagoe R, Miyashita N, Hikosaka O (1998) Presaccadic omnidirectional burst activity in the basal interstitial nucleus in the monkey cerebellum. Exp Brain Res 121: 442-450. Terao Y, Fukuda H, Ugawa Y, Hikosaka O, Hanajima R, Furubayashi T, Sakai K, Miyauchi S, Sasaki Y, Kanazawa I (1998) Visualization of the information flow through human oculomotor cortical regions by transcranial magnetic stimulation. J Neurophysiol 80: 936-946. Horikawa HPM, Nakazato T, Hikosaka O (1997) Duration of catalepsy correlates with increased intrastriatal sulpiride. European Journal of Pharmacology 326: 15-21. Miyachi S, Hikosaka O, Miyashita K, Karádi Z, Rand MK (1997b) Differential roles of monkey striatum in learning of sequential hand movement. Exp Brain Res 115: 1-5. Nakahara H, Doya K, Hikosaka O, Nagano S (1997b) Multiple representations in the basal ganglia loops for sequential decision making. Technical Report of IEICE NC97-24: 97-104. Shimojo S, Miyauchi S, Hikosaka O (1997) Visual motion sensation yielded by non-visually driven attention. Vision Res 37: 1575-1580. Hikosaka O, Sakai K, Miyauchi S, Takino R, Sasaki Y, Pütz B (1996) Activation of human presupplementary motor area in learning of sequential procedures: A functional MRI study. J Neurophysiol 76: 617-621. Kojima J, Matsumura M, Togawa M, Hikosaka O (1996) Tonic activity during visuo-oculomotor behavior in the monkey superior colliculus. Neurosci Res 26: 17-28. Miyashita K, Rand MK, Miyachi S, Hikosaka O (1996a) Anticipatory saccades in sequential procedural learning in monkeys. J Neurophysiol 76: 1361-1366.

Miyashita N, Hikosaka O (1996) Minimal synaptic delay in the saccadic output pathway of the superior colliculus studied in awake monkey. Exp Brain Res 112: 187-196. Hikosaka O, Rand MK, Miyachi S, Miyashita K (1995b) Learning of sequential movements in the monkey - Process of learning and retention of memory. J Neurophysiol 74: 1652-1661. Iwamura Y, Tanaka M, Hikosaka O, Sakamoto M (1995) Postcentral neurons of alert monkeys activated by the contact of the hand with objects other than the monkey's own body. Neurosci Lett 186: 127-130. Kato M, Miyashita N, Hikosaka O, Matsumura M, Usui S, Kori A (1995) Eye movements in monkeys with local dopamine depletion in the caudate nucleus. I. Deficits in spontaneous saccades. J Neurosci 15: 912-927. Kori A, Miyashita N, Kato M, Hikosaka O, Usui S, Matsumura M (1995) Eye movements in monkeys with local dopamine depletion in the caudate nucleus. II. Deficits in voluntary saccades. J Neurosci 15: 928-941. Miyashita N, Hikosaka O, Kato M (1995b) Visual hemineglect induced by unilateral striatal dopamine deficiency in monkeys. NeuroReport 6: 1257-1260. Miyashita N, Hikosaka O, Kato M (1995c) Visual hemineglect induced by unilateral striatal dopamine dificiency in monkeys. NeuroReport 6: 1257-1260. Hikosaka O, Miyauchi S, Shimojo S (1993c) Focal visual attention produces illusory temporal order and motion sensation. Vision Res 33: 1219-1240. Hikosaka O, Miyauchi S, Shimojo S (1993e) Voluntary and stimulus-induced attention detected as motion sensation. Perception 22: 517-526. Hikosaka O, Sakamoto M, Miyashita N (1993f) Effects of caudate nucleus stimulation on substantia nigra cell activity in monkey. Exp Brain Res 95: 457-472. Iwamura Y, Tanaka M, Sakamoto M, Hikosaka O (1993) Rostrocaudal gradients in the neuronal receptive field complexity in the finger region of the alert monkey's postcentral gyrus. Exp Brain Res 92: 360-368. Matsumura M, Kojima J, Gardiner TW, Hikosaka O (1992) Visual and oculomotor functions of monkey subthalamic nucleus. J Neurophysiol 67: 1615-1632. Usui S, Fukuda H, Hikosaka O (1990a) Spontaneous saccades in MPTP-induced hemi-parkinsonian monkeys. Clin Neurol 30: 1181-1189.

Hikosaka O, Sakamoto M, Usui S (1989a) Functional properties of monkey caudate neurons. I. Activities related to saccadic eye movements. J Neurophysiol 61: 780-798. Hikosaka O, Sakamoto M, Usui S (1989b) Functional properties of monkey caudate neurons. II. Visual and auditory responses. J Neurophysiol 61: 799-813. Hikosaka O, Sakamoto M, Usui S (1989c) Functional properties of monkey caudate neurons. III. Activities related to expectation of target and reward. J Neurophysiol 61: 814-832. Sakamoto M, Hikosaka O (1989) Eye movements induced by microinjection of GABA agonist in the rat substantia nigra pars reticulata. Neurosci Res 6: 216-233. Hikosaka O, Sakamoto M (1987) Dynamic characteristics of saccadic eye movements in the albino rat. Neurosci Res 4: 304-308. Sakamoto M, Hikosaka O (1987) Dynamic characteristics of saccadic eye movements in the albino rat. Neurosci Res 4: 304-308. Hikosaka O, Wurtz RH (1986) Saccadic eye movements following injection of lidocaine into the superior colliculus. Exp Brain Res 61: 531-539. Hikosaka O, Sakamoto M (1986a) Cell activity in monkey caudate nucleus preceding saccadic eye movements. Exp Brain Res 63: 659-662. Hikosaka O, Sakamoto M (1986b) Neural activities in the monkey basal ganglia related to attention, memory and anticipation. Brain Dev 8: 454-462. Hikosaka O, Wurtz RH (1985a) Modification of saccadic eye movements by GABA-related substances. I. Effect of muscimol and bicuculline in the monkey superior colliculus. J Neurophysiol 53: 266-291. Hikosaka O, Wurtz RH (1985b) Modification of saccadic eye movements by GABA-related substances. II. Effects of muscimol in monkey substantia nigra pars reticulata. J Neurophysiol 53: 292-308. Hikosaka O, Tanaka M, Sakamoto M, Iwamura Y (1985) Deficits in manipulative behaviors induced by local injections of muscimol in the first somatosensory cortex of the conscious monkey. Brain Res 325: 1-2. Iwamura Y, Tanaka M, Sakamoto M, Hikosaka O (1985b) Diversity in receptive field properties of vertical neuronal arrays in the crown of the postcentral gyrus of the conscious monkey. Exp Brain Res 58: 400-411.

Iwamura Y, Tanaka M, Sakamoto M, Hikosaka O (1985d) Vertical neuronal arrays in the postcentral gyrus signaling active touch: A receptive field study in the conscious monkey. Exp Brain Res 58: 412-420. Hikosaka O, Wurtz RH (1983a) Effects on eye movements of a GABA agonist and antagonist injected into monkey superior colliculus. Brain Res 272: 368-372. Hikosaka O, Wurtz RH (1983b) Visual and oculomotor functions of monkey substantia nigra pars reticulata. I. Relation of visual and auditory responses to saccades. J Neurophysiol 49: 1230-1253. Hikosaka O, Wurtz RH (1983c) Visual and oculomotor functions of monkey substantia nigra pars reticulata. II. Visual responses related to fixation of gaze. J Neurophysiol 49: 1254-1267. Hikosaka O, Wurtz RH (1983d) Visual and oculomotor functions of monkey substantia nigra pars reticulata. III. Memory-contingent visual and saccade responses. J Neurophysiol 49: 1268-1284. Hikosaka O, Wurtz RH (1983e) Visual and oculomotor functions of monkey substantia nigra pars reticulata. IV. Relation of substantia nigra to superior colliculus. J Neurophysiol 49: 1285-1301. Iwamura Y, Tanaka M, Sakamoto M, Hikosaka O (1983a) Converging patterns of finger representation and complex response properties of neurons in area 1 of the first somatosensory cortex of the conscious monkey. Exp Brain Res 51: 327-337. Iwamura Y, Tanaka M, Sakamoto M, Hikosaka O (1983b) Functional subdivisions representing different finger regions in area 3 of the first somatosensory cortex of the conscious monkey. Exp Brain Res 51: 315-326. Hikosaka O, Igusa Y (1980) Axonal projection of prepositus hypoglossi and reticular neurons in the brainstem of the cat. Exp Brain Res 39: 441-451. Hikosaka O, Igusa Y, Imai H (1980a) Inhibitory connections of nystagmus-related reticular burst neurons with neurons in the abducens, prepositus hypoglossi and vestibular nuclei in the cat. Exp Brain Res 39: 301-311. Hikosaka O, Nakao S, Shimazu H (1980b) Postsynaptic inhibition underlying spike suppression of secondary vestibular neurons during quick phases of vestibular nystagmus. Neurosci Lett 16: 21-26. Ishimine S, Hikosaka O, Nakamura Y (1980) Spatio-temporal patterns of pre- and

postsynaptic inhibition induced by primary afferent activation in the trigeminal sensory nucleus in cats. Exp Brain Res 40: 154-163. Iwamura Y, Tanaka M, Hikosaka O (1980) Overlapping representation of fingers in area 2 of the monkey somatosensory cortex of the conscious monkey. Brain Res 197: 516-520. Hikosaka O, Igusa Y, Imai H (1978a) Firing pattern of prepositus hypoglossi and adjacent reticular neurons related to vestibular nystagmus in the cat. Brain Res 144: 395-403. Hikosaka O, Igusa Y, Nakao S, Shimazu H (1978b) Direct inhibitory synaptic linkage of pontomedullary reticular burst neurons with abducens motoneurons in the cat. Exp Brain Res 33: 337-352. Hikosaka O, Maeda M, Nakao S, Shimazu H, Shinoda Y (1977) Presynaptic impulses in the abuducens nucleus and their relation to postsynaptic potentials in motoneurons during vestibular nystagmus. Exp Brain Res 27: 355-376. Hikosaka O, Kawakami T (1977) Inhibitory reticular neurons related to the quick phase of vestibular nystagmus - their location and projection. Exp Brain Res 27: 377-396. Hikosaka O, Kawakami T (1976) Inhibitory interneurons in the reticular formation and their relation to vestibular nystagmus. Brain Res 117: 513-518. Hikosaka O, Maeda M (1973) Cervical effects on abducens motoneurons and their interaction with vestibulo-ocular reflex. Exp Brain Res 18: 512-530. < Reviews > Kim HF, Hikosaka O (2015) Parallel basal ganglia circuits for voluntary and automatic behaviour to reach rewards. Brain 138: 1776-1800. Gottlieb J, Hayhoe M, Hikosaka O, Rangel A (2014) Attention, reward, and information seeking. J Neurosci 34: 15497–15504. Proulx CD, Hikosaka O, Malinow R (2014) Reward processing by the lateral habenula in normal and depressive behaviors. Nat Neurosci 17: 1146-1152. Hikosaka O, Kim HF, Yasuda M, Yamamoto S (2014) Basal Ganglia circuits for reward value-guided behavior. Annu Rev Neurosci 37: 289-306. Hikosaka O, Yamamoto S, Yasuda M, Kim, HF (2013) Why skill matters. Trends Cogn Sci 17: 434-441.

Terao Y, Fukuda H, Ugawa Y, Hikosaka O (2013) New perspectives on the pathophysiology of Parkinson's disease as assessed by saccade performance: a clinical review. Clin Neurophysiol 124: 1491-1506. Nakahara H, Hikosaka O (2012) Learning to represent reward structure: a key to adapting to complex environments. Neurosci Res 74: 177-183. Isoda M, Hikosaka O (2011) Cortico-basal ganglia mechanisms for overcoming innate, habitual and motivational behaviors. Eur J Neurosci 33:2058-2069. Bromberg-Martin ES, Matsumoto M, Hikosaka O (2010) Dopamine in motivational control: rewarding, aversive, and alerting. Neuron 68:815-834. Hikosaka O (2010) The habenula: from stress evasion to value-based decision-making. Nat Rev Neurosci 11:503-513. Hikosaka O, Isoda M (2010) Switching from automatic to controlled behavior: cortico-basal ganglia mechanisms. Trends Cogn Sci 14: 154-161. Hikosaka O, Sesack SR, Lecourtier L, Shepard PD (2008) Habenula: crossroad between the basal ganglia and the limbic system. J Neurosci 28:11825-11829. Hikosaka O, Bromberg-Martin E, Hong S, Matsumoto M (2008) New insights on the subcortical representation of reward. Curr Opin Neurobiol 18:203-208. Hikosaka O (2008) [Decision-making and learning by cortico-basal ganglia network]. Brain Nerve 60:799-813. Balleine BW, Delgado MR, Hikosaka O (2007) The role of the dorsal striatum in reward and decision-making. J Neurosci 27:8161-8165. Hikosaka O (2007) Basal Ganglia Mechanisms of Reward-oriented Eye Movement. Ann N Y Acad Sci 1104:229-249. Hikosaka O (2007) Habenula. Scholarpedia 2:2703. http://www.scholarpedia.org/article/Habenula Hikosaka O (2007) GABAergic output of the basal ganglia. Prog Brain Res 160:209-226. Hikosaka O, Nakamura K, Nakahara H (2006) Basal Ganglia orient eyes to reward. J Neurophysiol 95:567-584. Sakai K, Hikosaka O, Nakamura K (2004) Emergence of rhythm during motor learning. Trends Cogn Sci 8:547-553.

Donoghue JP, Hikosaka O (2003) Motor systems. Curr Opin Neurobiol 13:643-646. Hikosaka O (2002) A new approach to the functional systems of the brain. Epilepsia 43 Suppl 9:9-15. Hikosaka O, Nakamura K, Sakai K, Nakahara H (2002) Central mechanisms of motor skill learning. Curr Opin Neurobiol 12: 217–222. Hikosaka O, Takikawa Y, Kawagoe R (2000) Role of the basal ganglia in the control of purposive saccadic eye movements. Physiol. Rev. 80: 953-978. Hikosaka O, Sakai K, Nakahara H, Lu X, Miyachi S, Nakamura K, Rand MK (1999) Neural mechanisms for learning of sequential procedures. In: The New Cognitive Neurosciences (Gazzaniga MS, eds), pp. 553-572, Cambridge, MA: The MIT Press. Hikosaka O, Nakahara H, Rand MK, Sakai K, Lu X, Nakamura K, Miyachi S, Doya K (1999) Parallel neural networks for learning sequential procedures. Trends Neurosci 22: 464-471. Hikosaka O (1999) Probing consciousness with an electrode. Brain Res Bull 50: 337-338. Hikosaka O (1998b) Neural systems for control of voluntary action - a hypothesis. Advances in Biophysics 35: 81-102. Hikosaka O, Miyashita K, Miyachi S, Sakai K, Lu X (1998) Differential roles of the frontal cortex, basal ganglia, and cerebellum in visuomotor sequence learning. Neurobiology of Learning and Memory 70: 137-149. Hikosaka O, Miyauchi S, Shimojo S (1996e) Orienting of spatial attention - its reflexive, compensatory, and voluntary mechanisms. Cognitive Brain Research 5: 1-9. Hikosaka O, Miyauchi S, Takeichi H, Shimojo S (1996f) Multimodal spatial attention visualized by motion illusion. In: Attention and Performance XVI : Information Integration in Perception and Communication (Inui T, McClelland J, eds), pp. 237-261. Cambridge, MA: MIT Press. Hikosaka O, Miyauchi S, Shimojo S (1993d) Visual attention revealed by an illusion of motion. Neurosci Res 18: 11-18. Hikosaka O (1991a) Basal ganglia - possible role in motor coordination and learning. Curr Opin Neurobiol 1: 638-643. Hikosaka O (1989c) Role of basal ganglia in saccades. Rev Neurol (Paris) 145: 580-586.

Evarts EV, Kimura M, Wurtz RH, Hikosaka O (1984) Behavioral correlates of activity in basal ganglia neurons. Trends Neurosci 7: 447-453. < Book sections > Hikosaka O (2009) Basal Ganglia and Oculomotor Control. In: Squire LR (ed.) Encyclopedia of Neuroscience, volume 2, pp. 53-61. Oxford: Academic Press. Shimojo S, Hikosaka O, Miyauchi S (In press) Automatic and controlled processes of attention revealed in the line motion effect. In: Attention & Performance XV (Gopher D, Koirat A, eds), pp. Cambridge, MA: MIT Press. Hikosaka O, Sakai K, Nakahara H, Lu X, Miyachi S, Nakamura K, Rand MK (1999) Neural mechanisms for learning of sequential procedures. In: The Cognitive Neurosciences (Gazzaniga MS, eds), pp. 553-572. Cambridge, MA: The MIT Press. Hikosaka O, Miyachi S, Miyashita K, Rand MK (1996a) Learning of sequential procedures in monkeys. In: The Acquisition of Motor Behavior in Vertebrates (Bloedel JR, Ebner TJ, Wise SP, eds), pp. 303-317. Cambridge, MA: The MIT Press. Hikosaka O, Miyachi S, Miyashita K, Rand MK, Karadi Z (1996b) Procedural learning in monkeys - possible roles of the basal ganglia. In: Perception, Memory, and Emotion: Frontier in Neuroscience (Ono T, McNaughton BL, Molotchnikoff S, Rolls ET, Nishijo H, eds), pp. 403-420. Oxford, UK: Elsevier. Hikosaka O, Miyachi S, Miyashita K, Sakai K, Miyauchi S, Takino R, Sasaki Y, Pütz B (1996c) Neural mechanisms of sequential procedural learning. In: Integrative and Molecular Approach to Brain Function (Ito M, Miyashita Y, eds), pp. 285-292. Amsterdam: Elsevier. Hikosaka O, Miyashita K, Sakai K, Miyauchi S, Takino R, Sasaki Y, Pütz B (1996d) Role of the medial frontal cortex in sequential procedural learning. In: Brain Processes and Memory (Ishikawa K, McGaugh JL, Sakata H, eds), pp. 299-306. Amsterdam: Elsevier. Hikosaka O, Rand MK, Miyashita K, Miyachi S (1996g) Learning of sequential procedures in monkeys. In: Recent Advances in Clinical Neurophysiology (Kimura J, Shibasaki H, eds), pp. 296-303. Amsterdam: Elsevier. Hikosaka O, Sakai K, Miyauchi S, Takino R, Sasaki Y, Pütz B (1996j) Brain activation during learning of sequential procedures. In: Visualization of Information Processing in the Human Brain: Recent Advances in MEG and Functional MRI (Hashimoto I, Okada YC, Ogawa S, eds), pp. 245-252. Amsterdam: Elsevier.

Segawa M, Tanaka R, Nomura Y, Fukuda H, Hikosaka O (1996) Saccadic eye movements in basal ganglia disorders. In: Recent Advances in Clinical Neurophysiology (Kimura J, Shibasaki H, eds), pp. 170-176. Amsterdam: Elsevier. Shimojo S, Tanaka Y, Hikosaka O, Miyauchi S (1996) Vision, attention, and action: Inhibition and facilitation in sensory-motor links revealed by the reaction time and the line motion. In: Attention and Performance XVI : Information Integration in Perception and Communication (Inui T, McClelland J, eds), pp. 597-630. Cambridge, MA: MIT Press. Hikosaka O (1995) Neural mechanisms of saccadic eye movement. In: Vertigo, Nausea, tinnitus and Hearing Loss in Central and Peripheral Vestibular Diseases. Proceedings of the XXIInd Annual Meeting of the International Neurootological and Equilibriometric Society. (Claussen C-F, Sakata E, Itoh A, eds), pp. 3-18. Amsterdam: Elsevier. Hikosaka O, Fukuda H, Segawa M, Nomura Y (1995a) Voluntary saccades in normal and in dopamine deficient subjects. In: Age-Related Dopamine-Deficient Disorders (Segawa M, Nomura Y, eds), pp. 59-68. Basal: Karger. Hikosaka O, Rand MK, Miyachi S, Miyashita K (1995c) Procedural learning in the monkey. In: Functions of the Cortico-Basal Ganglia Loop (Kimura M, Graybiel AM, eds), pp. 18-30. Tokyo: Springer. Kato M, Hikosaka O (1995) Function of the indirect pathway in the basal ganglia oculomotor system: visuo-oculomotor activities of external pallidum neurons. In: Age-Related Dopamine-Deficient Disorders (Segawa M, Nomura Y, eds), pp. 178-187. Basal: Karger. Hikosaka O (1994) Role of basal ganglia in control of innate movements, learned behavior and cognition - a hypothesis. In: The Basal Ganglia IV: New Ideas and Data on Structure and Function (Percheron G, McKenzie JS, Féger J, eds), pp. 589-596. New York: Plenum Press. Segawa M, Hikosaka O, Fukuda H, Uetake K (1994) Deficits in saccadic eye movements in basal ganglia disorders. In: The Basal Ganglia IV: New Ideas and Data on Structure and Function (Percheron G, McKenzie JS, Feger J, eds), pp. 525-531. New York: Plenum Press. Hikosaka O, Matsumura M, Kojima J, Gardiner TW (1993b) Role of basal ganglia in initiation and suppression of saccadic eye movements. In: Role of the Cerebellum and Basal Ganglia in Voluntary Movement (Mano N, Hamada I, DeLong MR, eds), pp. 213-219. Amsterdam: Elsevier. Hikosaka O (1993) Role of the basal ganglia in motor learning: a hypothesis. In: Brain Mechanisms of Perception and Memory (Ono T, Squire LR, Raichle ME, PerrettD.I., Fukuda M, eds), pp. 497-513. New York: Oxford University Press.

Hikosaka O, Fukuda H, Kato M, Uetake K, Nomura Y, Segawa M (1993a) Deficits in saccadic eye movements in hereditary progressive dystonia with marked diurnal fluctuation. In: Hereditary Progressive Dystonia with Marked Diurnal Fluctuation (Segawa M, eds), pp. 159-177. Lancs & New York: The Parthenon Publishing Group. Hikosaka O, Imai H, Segawa M (1992) Saccadic eye movements in parkinsonism. In: Vestibular and Brain Stem Control of Eye, Head and Body Movements (Shimazu H, Shinoda Y, eds), pp. 405-414. Tokyo: Japan Scientific Society Press. Hikosaka O (1991b) Role of the forebrain in oculomotor function. In: Role of the Forebrain in Sensation and Behavior (Holstege G, eds), pp. 101-107. Amsterdam: Elsevier. Hikosaka O (1989a) Motor programming in basal ganglia. In: Neural Programming (Ito M, eds), pp. 101-109. Tokyo: Japan Scientific Societies Press. Hikosaka O (1989b) Role of basal ganglia in initiation of voluntary movements. In: Dynamic Interactions in Neural Networks: Models and Data (Arbib MA, Amari S, eds), pp. 153-167. New York: Springer-Verlag. Hikosaka O, Wurtz RH (1989) The basal ganglia. In: The Neurobiology of Saccadic Eye Movements (Wurtz RH, Goldberg ME, eds), pp. 257-281. Amsterdam: Elsevier. Hikosaka O, Segawa M, Imai H (1987) Voluntary saccadic eye movement: application to analyze basal ganglia disease. In: Highlights in Neuro-Ophthalmology (Ishikawa S, eds), pp. 133-138. Amsterdam: Aeolus Press. Nomura Y, Segawa M, Soda M, Hikosaka O (1987) Voluntary saccadic eye movements in basal ganglia disorders. In: Highlights in Neuro-Ophthalmology (Ishikawa S, eds), pp. 139-145. Amsterdam: Aeolus Press. Segawa M, Nomura Y, Hikosaka O, Soda M, Usui S, Kase M (1987) Roles of the basal ganglia and related structure in symptoms of dystonia. In: The Basal Ganglia II. Structure and Function-Current Concepts (Carpenter MB, Jayaraman A, eds), pp. 489-504. New York: Plenum. Wurtz RH, Hikosaka O (1986) Role of the basal ganglia in the initiation of saccadic eye movements. In: Progress in Brain Research (Freund H-J, Büttner U, Cohen B, Noth J, eds), pp. 175-190. Amsterdam: Elsevier. Iwamura Y, Tanaka M, Sakamoto M, Hikosaka O (1985a) Comparison of the hand and finger representation in areas 3,1, and 2 of the monkey somatosensory cortex. In: Development, Organization, and Processing in Somatosensory Pathways (Rowe M, Willis D, eds), pp. 239-245. New York: Liss Inc.

Hikosaka O, Wurtz RH (1981) The role of substantia nigra in the initiation of saccadic eye movements. In: Progress in Oculomotor Research (Fuchs A, Becker W, eds), pp. 145-152. North Holland: Elsevier. Iwamura Y, Tanaka M, Hikosaka O (1981) Cortical neuronal mechanisms of tactile perception studied in the conscious monkey. In: Brain Mechanisms of Sensation (Katsuki, Sato M, Norgren R, eds), pp. 61-70. New York: J. Wiley. Wurtz RH, Albano JE, Hikosaka O (1980) Relation of superior colliculus to the initiation of eye movements. In: Adv. Physiol. Sci. (Szentágothai J, Palkovits M, Hámori J, eds), pp. 251-257. < Invited Lectures > “Parallel basal ganglia circuits for reward value-guided behavior” Plenary lecture Japan Neuroscience Society Meeting Kobe, Japan Jul. 29, 2015 “Parallel basal ganglia circuits for voluntary and automatic behavior” The Torsten Wiesel Lecture IBRO World Congress Rio de Janeiro, Brazil Jul. 10, 2015 “Basal ganglia mechanisms for value-guided behavior” Plenary lecture French Neuroscience Society Meeting Montpellier, France May 20, 2015 “Multiple mechanisms for choosing good objects” NIDA Conference: 40 years of dopamine and reward: a celebration of the career of Roy Wise National Institute on Drug Abuse, Baltimore, MD May 13, 2015 “Voluntary and automatic gaze orienting controlled by basal ganglia circuits” Janelia Farm Conference: Motivational Circuits in Natural and Learned Behaviors Janelia Farm Research Campus, VA Mar. 29 – Apr. 1, 2015

“Basal ganglia mechanisms for automatic and voluntary choice of good objects” Cornell University Dec. 4, 2014 “Gaze Orienting based on Long-term Value Memory – Basal Ganglia mechanism” Symposium: “Attention, Reward, and Information Seeking” Society for Neuroscience Annual Meeting Washington DC, USA Nov. 17, 2014 “Parallel basal ganglia circuits for choosing good objects” Columbia University May 29, 2014 “Parallel basal ganglia mechanisms for choosing good objects” Harvard University, The Center for Brain Science Apr. 29, 2014 “Basal ganglia decision mechanisms based on short-term and long-term memories” Carlson Symposium: “Cortical and Subcortical Mechanisms of Decision” University of Chicago Apr. 2, 2014 “Basal ganglia mechanisms for choosing good objects” Stony Brook University Mar. 13, 2014 “Basal ganglia circuits for reward value-guided behavior” Yerkes National Primate Center, Emory University Atlanta, GA Sep. 30, 2013 “Distinct basal ganglia circuits controlling behaviors aiming at flexible and stable values” University of Pittsburgh Jul. 25, 2013 “Long-term object-value memories in the basal ganglia underlying visuomotor skill” University of Colorado at Boulder Boulder, CO Apr. 23, 2013 “Differential neuronal encoding of value, salience, and uncertainty” Sixth Annual Swartz Symposium: Neural Circuits for Decision Making and Reinforcement Learning

Yale University Apr. 12, 2013 “Parallel control of volition and skill by the basal ganglia” Keynote Lecture IBAGS-XI 11th Triennial meeting of the International Basal Ganglia Society Eilat, Israel March 3-7, 2013 “Role of basal ganglia in valuation, memory, and skill” University of California, San Diego San Diego, CA Feb. 12, 2013 “Choosing valuable objects automatically – a basal ganglia mechanism” Workshop 2: Cognitive Neuroscience Mathematical Biosciences Institute (MBI) Columbus, Ohio Dec. 11, 2012 “Basal ganglia mechanisms for choosing valuable objects automatically” University of Parma Parma, Italy Nov. 16, 2012 “Automatic valuation by the basal ganglia” ESF-FENS Conferences on The Dynamic Brain The Neurobiology of Emotion Stresa, Italy Nov. 13, 2012 “Basal ganglia mechanisms for choosing valuable objects automatically” Champalimaud Neuroscience Programme (CNP), Lisbon, Portugal Nov. 8, 2012 “Long-term reward experiences bias eye movements: a role of the basal ganglia” David Robinson Lecture Johns Hopkins University, School of Medicine Apr. 5, 2012 “Choosing good objects – a basal ganglia mechanism” Inaugural Phillip A. Sharp Lecture in Neural Circuits McGovern Institute, MIT

Mar. 1, 2012 “Some issues in decision-making research” University of Texas, Austin, TX Dec. 13, 2011 “Long-term reward experiences bias eye movements: a role of the basal ganglia” University of Texas, Austin, TX Dec. 12, 2011 “Long-term object-reward memories in the basal ganglia underlying visuomotor habit” University of Minnesota, MN Oct. 14, 2011 “How long-term reward experience influences behavior: a possible role of the posterior ganglia” Keynote Lecture Janelia Farm Conference: Neural Circuits of Decision-Making Janelia Farm Research Campus, VA Mar. 6-9, 2011 “Subcortical mechanisms of motivational behavior” Carnegie Institution Baltimore, MD Jan. 17, 2011 “Motivational Neuronal Circuits for Value, Salience, and Information” Presidential Special Lecture Society for Neuroscience, 40th Annual Meeting San Diego, CA Nov. 14, 2010 “Decision-making by cortico-basal ganglia networks” NIH Research Festival Symposia Session III: The Brain and the Construction of Complex Behaviors Oct. 6, 2010 “Lateral habenula sends motivational information to the basal ganglia” The International Basal Ganglia Society, 10th Triennial Meeting Long Branch, NJ Jun. 20-24, 2010 “Role of the lateral habenula in value-based decision-making” Symposium: The Lateral Habenula: Its Role in Behavior and Psychiatric Disorders

Feb. 28 – Mar. 3, 2010 Cold Spring Harbor, NY “Subcortical control of motivational behavior” RIKEN, BSI Jan. 26, 2010 “Subcortical mechanisms of motivational behavior” New York University Dec. 7, 2009 “Subcortical mechanisms of motivational behavior - Beyond dopamine neurons” University of Pennsylvania Nov. 23, 2009 “Subcortical neurons representing motivational value and salience” Yale University Sep. 17, 2009 “Rethinking dopamine neurons” Tamagawa University August 4, 2009 “Biasing, switching, and learning of behavior by the basal ganglia” Symposium: Multidisciplinary Approaches to Basal Ganglia Functions International Union of Physiological Sciences 2009 Kyoto, Japan July 29, 2009 “New concepts of dopamine neurons” Hokkaido University July 21, 2009 “Subcortical mechanisms of reward-based learning” Gordon Research Conferences: Neural Circuits & Plasticity Salve Regina University, Newport, RI June 7-12, 2009 “Motor skill learning and decision-making: cortico-basal ganglia mechanisms” Georgetown University May 12, 2009 “How the brain encodes reward” McGovern Institute (MIT) Symposium on “The Basal Ganglia in Health and Disease”

MIT, Boston May 7, 2009 “Subcortical networks for motivational decision making” Eye Movements and Vision: A Symposium in Honor of Fred Miles Oxford University Apr. 1-3, 2009 “Neural mechanisms of motor skill learning, execution, and switching” NINDS Mar. 3, 2009 “Role of the subcortical reward system in decision-making” Baylor College of Medicine Houston, Texas Feb. 6, 2009 “Multiple neural mechanisms of motor learning, automaticity, and switching” Netherlands Institute for Neuroscience (NIN) conference Amsterdam, Netherlands Dec. 9-12, 2008 “Role of the Primate Lateral Habenula in Negative Motivational Control of Oculomotor Behavior” Symposium: “Habenula - Crossroad between the Basal Ganglia and the Limbic System” Society for Neuroscience Annual Meeting Washington DC, USA Nov. 19, 2008. “Basal ganglia mechanisms of reward-based and instruction-based decision-making” University of Texas, Austin Oct. 6, 2008 “Reward-based control of eye movements by the basal ganglia and surrounding structures” National Institute on Drug Abuse (NIDA) Sep. 23, 2008 “Motivational and cognitive control of behavior by the basal ganglia” Workshop on Mechanism of Brain and Mind Sapporo, Japan Aug. 10, 2008 “Neural mechanisms of reward-based and instruction-based decision-making” Columbia University

Jun. 19, 2008 “Integration of cognition and motivation in the basal ganglia and surrounding structures” Albert Einstein College of Medicine of Yeshiva University Apr. 23, 2008 “Neural mechanisms of reward-oriented behavior: Beyond dopamine neurons” Harvard University Feb. 5, 2008 “Role of the medial frontal cortex in switching from automatic to controlled behavior” Symposium on “Using eye movements as an experimental probe of brain function” London, UK Dec 5-6, 2007 “Neural mechanisms of reward-oriented behavior: Beyond dopamine neurons” OIST Workshop on Cognitive Neurobiology Okinawa, Japan March 11-15, 2007 “Role of the medial frontal cortex in switching from automatic to controlled behavior” The 10th Tamagawa-Riken Dynamic Brain Forum (DBF’07) Hakuba, Japan March 5-9, 2007 “Mechanisms of reward-oriented eye movement: basal ganglia and beyond” Rutgers University Dec. 13, 2006 “Basal ganglia mechanisms of reward-oriented eye movement” Symposium on “Reward and decision making in cortico-basal ganglia networks” UCLA Lake Arrowhead Conference Center June 1-4, 2006 “Motivational control of saccadic eye movement: Role of the basal ganglia” Stanford University May 18, 2006 “Motivational control of saccadic eye movement: Role of the basal ganglia” Columbia University Mar 23, 2006 “Role of the basal ganglia in reward-oriented eye movement” University of Maryland

Dec. 2, 2005 “Representation and achievement of motivational goal by the basal ganglia” Duke University Nov. 29, 2005 “Basal ganglia mechanisms of reward-oriented eye movement” Symposium on “Neurobiology of Decision-Making” Cold Spring Harbor Laboratory May 24 (22-25), 2005 “Neural mechanisms of reward-oriented saccadic eye movement” University of Rochester Nov. 8, 2004 “Role of the basal ganglia in reward-oriented saccadic eye movement” University of Pittsburgh Jun. 4, 2004 “Motivational modulation of eye movements by the basal ganglia” Brandeis University Apr. 26, 2004 “Does the conscious mind control movement? ” Neural Control of Movement (NCM) 14th Annual Meeting Sitges, Spain Apr. 2, 2004 “Temporal organization of motor skill” Neural Control of Movement (NCM) Satellite Meeting “Motor Learning” Barcelona, Spain Mar. 26, 2004 “Motivational Control of Saccadic Eye Movement” Symposium: "Multidisciplinary Approaches to Sensorimotor Integration --- Old Questions Meet New Concepts ---" The National Institute for Physiological Sciences, Okazaki, Japan Mar. 18, 2004 “Role of the basal ganglia in reward-oriented saccadic eye movement” Tamagawa University Mar. 12, 2004

“Role of the basal ganglia in reward-oriented saccadic eye movement” University of California, San Francisco Feb. 5, 2004 “Motivational control of saccadic eye movement: Role of the basal ganglia” BME seminar series, Johns Hopkins University Oct. 24, 2003 “Neural networks for learning sequential procedures” 9th annual meeting of the Organization for Human Brain Mapping New York Jun. 20, 2003 “Neural circuits for motivational bias: Role of the basal ganglia” Oxford University Jun. 18, 2003 “Learning and memory of visuo-motor skills” Symposium: Neural Control of Skilled Hand Movements: Cognitive and Computational Aspects Stockholm, Sweden Jun. 12, 2003 “Role of the basal ganglia in the motivational control of saccadic eye movement” Emory University Dec. 11, 2002 “Neural control of voluntary action - Role of the basal ganglia” Symposium: Genes and Minds Initiative Workshop on Ape Genomics Tokyo March 14-15, 2001 “Neural control of voluntary saccades - Role of the basal ganglia” Symposium: Higher Nervous Control of Posture and Locomotion: Parallel and Centralized Control Mechanisms Okazaki March 19-21, 2001 “A principle of basal ganglia function: why and how” Panel: Principles of Basal Ganglia Function: From Motivation to Decision Making Eleventh Annual Meeting for Neural Control of Movement. Seville, Spain March 25-30, 2001

“Progress in the neurophysiology underlying movement disorders” The 5th International Coference on Procegress iin Alzheimer's and Parkinson's disease Kyoto, Japan March 31 – April 4, 2001 “Integration of motivation and cognition in the basal ganglia for goal-directed eye movement” CREST Workshop on Metalearning and Neuromodulation Seika, Kyoto, Japan April 6-7, 2001 “Motivational control of saccadic eye movement - a role of the basal ganglia” The Pittsburgh Chapter of the Society for Neuroscience Grass Traveling Scientist Pittsburgh, USA Nov. 13, 2000 “Role of the basal ganglia in the cognitive and motivational control of action” NIH Neuroscience Seminar Series Bethesda, USA Oct. 16, 2000 “Role of the basal ganglia in the motivational control of saccadic eye movement” Neuroscience Seminar at University of Washington Seattle, USA Oct. 12, 2000 “Role of the basal ganglia in the reward-directed saccadic eye movement” Neuroscience Seminar at Vanderbilt University Nashville, USA Oct. 10, 2000 “The dopaminergic neurons and behavior” World Congress on Rett syndrome 2000 Satellite Symposium: Developmental Neurology for Understanding the Pathophysiology of Rett syndrome Karuizawa, Japan Jul. 24, 2000 “Role of the basal ganglia in the cognitive and motivational control of action” RIKEN BSI Summer Program 2000 Saitama, Japan Jul. 14, 2000

“Integration of cognition and motivation in the basal ganglia for oculomotor control” Seminar at National Institute of Health Oct. 29, 1999 “Learning and memory for sequential procedures” International Seminar on Neurobiology X Tokyo Jan. 20-21, 1998 “Neural mechanisms for learning of sequential procedures” 1998 James S. McDonnell Foundation Summer Institute in Cognitive Neuroscience Squaw Creek, California, U.S.A. Jul. 7, 1998 “Multiple neural systems for procedural learning” The Canada-Japan Neuroscience Partnership Meeting Fuji, Japan Aug. 21, 1998 “Reward-dependent modulation of caudate and dopaminergic neurons” 6th Triennial Meeting of International Basal Ganglia Society Brewster, MA, U.S.A. Oct. 18, 1998 “Translation of motivation into action in the basal ganglia” Plenary Lecture The Fifth International Conference on Neural Information Processing Kita-kyushu, Japan Oct. 22, 1998 “Orienting of spatial attention revealed by motion illusion” Stanford University March 5, 1997. “Neural mechanisms of sequential procedural learning” University of California, San Francisco March 6, 1997. “Differential neural activity for learned and new actions” New York University's 4th Center for Neural Science Symposium "From Attention to Action: Contemporary Perspectives on Movement Planning, Preparation and Action" New York June 9-12, 1997.

“Differential roles of the frontal cortex, basal ganglia, and cerebellum in visuo-motor sequence learning” Sixth Conference on the Neurobiology of Learning and Memory "Brain and Memory: From Genes to Behavior" University of California, Irvine Oct. 21-24, 1997. “Neural mechanisms of sequential procedural learning” Uehara Memorial Foundation Symposium: Integrative and Molecular Approach to Brain Function Tokyo June 25-27, 1996. “Role of the basal ganglia in initiation and suppression of movements” 1st FAONs Congress IBRO Regional Congress Pattaya, Thailand Oct. 2-23, 1996. “Neural Mechanism of Saccadic Eye Movement” XXIInd Congress of the International Neurootological & Equilibriometric Society Opening Lecture: May, 1995 “Learning of sequential procedures in monkey” Acquisition of Motor Behavior in Vertebrates Satellite symposium for Fifth Annual Meeting of the Society for the Neural Control of Movement Key West, Florida April 23-25, 1995. “Basal ganglia and motor control” Looking Ahead in Sensorimotor Control: Big Questions for 2001 Kingson, Canada June 23-24, 1995. “Procedural learning in monkeys - possible roles of the basal ganglia” Perception, Memory, and Emotion: Frontiers in Neuroscience Toyama, Japan July 3-7, 1995. “Possible roles of the basal ganglia in procedural learning” Fourth IBRO World Congress of Neuroscience Kyoto

July 9-14, 1995. “Brain activation during learning of sequential procedures” Visualization of Information Processing in the Human Brain: Recent Advances in MEG and Functional MRI (The Tenth Tokyo Institute of Psychiatry International Symposium) Tokyo Oct. 12-13, 1995. “Learning of sequential procedures in monkeys” Motor Learning and Plasticity Workshop for The Xth International Congress of Electromyography and Clinical Neurophysiology Tokyo Oct. 16, 1995. “Role of the medial frontal cortex in sequential procedural learning” Nihon University International Symposium: Brain Processes and Memory Shizuoka, Japan Dec. 2, 1995. “Orienting of spatial attention - Its reflexive, compensatory, and voluntary mechanisms” Brain & Mind (The Ninth Toyota Conference) Shizuoka, Japan Dec. 5-8, 1995. “Multimodal spatial attention visualized by motion illusion” Attention & Performance XVI : Information Integration in Perception and Communication Kyoto July 11-15, 1994. “Muscimol, Memory, and Motivation in the Basal Ganglia” Special lecture Society for Neuroscience, 24th Annual Meeting Miami, Florida, USA Nov. 17, 1994. “Learning of sequential movements in the monkey” International Symposium: Functional Linkages Between the Cerebral Cortex and Basal Ganglia in the Control of Voluntary Movement (50th Anniversary of Osaka University) Osaka Dec. 13-15, 1993.

“Inhibitory Control of Saccadic Eye Movement by the Basal Ganglia” APS Conference on the Physiology and Pharmacology of Motor Control. San Diego (USA) Oct. 2-5, 1993. “Voluntary saccade” Symposium on Age Related Monoamine Dependent Disorders and their Modulation by Gene and Gender. Tokyo Nov. 12-13, 1993. “Saccadic eye movements in parkinsonism” Satellite Symposium of the 16th Barany Society Meeting: Vestibular and Brain Stem Control of Eye, Head and Body Movements. Tokyo, Japan May 25-27, 1990 “Basal ganglia control of eye movements” Symposium: Basal Ganglia Mechanisms in Health and Disease. Santillana del Mar, Spain September 1, 1990 “Dopamine and basal ganglia” Satellite Symposium of the 17th Congress of Collegium Internationale Neuro-psychopharmacologicum (Biwako Symposium): Mechanism of dopamine transmission: an update. Nagahama, Shiga, Japan September 8, 1990 “Electro-oculography” Satellite Symposium of the Joint Convention of the 5th International Child Neurology and the 3rd Asian and Oceanian Congress of Child Neurology: Symposium on Hereditary Progressive Dystonia with Marked Diurnal Fluctuation. Tokyo, Japan Novemember 11, 1990 “Role of forebrain in oculomotor control” Symposium: Role of the forebrain in sensation and behavior. NASA/Ames Research Center, Moffet Field, CA. USA. May 26-27, 1989. “Basal ganglia control of saccadic eye movement” Symposium: Brainstem Control of Posture and Movements. Asahikawa, Japan.

September 12-14, 1989. “Role of basal ganglia in oculomotor function” Symposium: “The Basal Ganglia: Structure and Function” Society for Neuroscience, 19th Annual Meeting. Phoenix, Arizona, USA. November 2, 1989. “Role of basal ganglia in saccades” Internatinal Meeting of the French Neurological Society, Gaze and Vision. Paris, France. June 1988. “Motor programming in basal ganalia” The Twelfth International Symposium of Brain Sciences, The Taniguchi Foundation, Neural Programming. Ohtsu, Japan. November 21-24, 1988. “Role of basal ganglia in the initiation of voluntary movements” Symposium: Competition and Cooperation in Neural Nets 2. University of Southern California, CA, USA. May 18-22, 1987.