第 4 章 考察
4.2. 結論
本論文では,運動の制御とは独立した空間で行う対象の選択と,実際に行う
運動の空間で行う運動選択に,複数の大脳皮質-基底核ループがどのように関わ
っているのかを調べることを最終目標として,サルを用いた実験(第2章)と,
ラットを用いた実験1(第3章)を行った.サルを用いた実験では,運動の制御 とは独立した空間で行う対象の選択に線条体がどのように関わるのかを電気生
理学的に検証した.報酬を最大化するための行動を学習する課題の中で,運動
要素を含まない対象の選択と,意思決定を実行するための運動の選択を時間的
に分離した試行を考案し,課題遂行中の線条体から細胞外記録を行った.その
結果,線条体の中でも,吻側尾状核で対象の選択に関わる情報が強く表現され
ていることが明らかになった.ラットを用いた実験1では,意思決定の最終過程,
すなわち運動実行を行う際に,一次運動野とその出力先である背外側線条体の
スパイク情報の性質を調べるために,頭部固定下でレバー運動中のラットを用
いてホールセル記録を行い,その時の膜電位変動とスパイクとの関係を解析し
た.その結果,一次運動野・線条体,どちらの領域においても,スパイクは膜
電位の揺らぎよりはむしろ膜電位そのものが安定的に変化することによって生
じる性質があることが明らかになった.
これら神経細胞レベルでの意思決定研究で得た結果は,さまざまなレベルで
80
の意思決定に関わる皮質-基底核ループの役割を明らかにするための重要な基礎
的所見であり,将来的には,多様な症状を呈する大脳基底核疾患の神経メカニ
ズムの解明にもつながると考えられる.
81
謝辞
本研究を遂行するにあたり,鮫島和行博士,礒村宜和博士には,実験の手順 から論文の執筆にいたるまで懇切丁寧な指導をしていただきました.丹治順博 士には,日頃より多くのご高配を賜りました.木村實博士には,議論を通じて 多くの知識や示唆を頂きました.藤山文乃博士には,論文執筆にあたり暖かい ご支援を賜りました.加藤康広博士,塚本葉子博士,梶原隆文博士には,有用 なご意見とデータをいただきました.銅谷賢治博士にはデータ解析についての 貴重なアドバイスを頂きました.濵谷京子さん,川島伊久乃さん,硤合千恵さ んには,事務補助をしていただきました.星英司博士をはじめとする星研究室 の皆様,礒村研究室,木村研究室の皆様には,たくさんのご厚情に預かりまし た.日頃より暖かいご支援ご協力をしていただいた玉川大学脳科学研究所の皆 様に厚く御礼申し上げます.また,ご多忙のところ本論文の審査をして頂いた 坂上雅道博士,酒井裕博士,松元健二博士に深く御礼申し上げます.
最後に,これまで常に支え続けてくれた妻・あゆみと息子・晟に心より感謝 いたします.
82
引用文献
第 1 章
[1] J. Neumann, O. Morgenstern, Theory of Games and Economic Behavior, Princeton Univ Press, Princeton, NJ (1944).
[2] R.S. Sutton, A.G. Barto, Reinforcement learning: An introduction, MIT Press, Cambridge MA USA (1998).
[3] A. Rangel, C. Camerer, P.R. Montague, A framework for studying the neurobiology of value-based decision making, Nat Rev Neurosci, 9 (2008) 545-556.
[4] E.V. Evarts, Relation of pyramidal tract activity to force exerted during voluntary movement, J Neurophysiol, 31 (1968) 14-27.
[5] O. Hikosaka, H.F. Kim, M. Yasuda, S. Yamamoto, Basal ganglia circuits for reward value-guided behavior, Annu Rev Neurosci, 37 (2014) 289-306.
[6] Graybiel, A. M. Habits, rituals, and the evaluative brain. Annu Rev Neurosci 31, 359-387, doi:10.1146/annurev.neuro.29.051605.112851 (2008).
[7] B.J. Knowlton, J.A. Mangels, L.R. Squire, A neostriatal habit learning system in humans, Science, 273 (1996) 1399-1402.
[8] M. Kobayakawa, S. Koyama, M. Mimura, M. Kawamura, Decision making in Parkinson's disease: Analysis of behavioral and physiological patterns in the Iowa gambling task, Mov Disord, 23 (2008) 547-552.
[9] G.E. Alexander, M.R. DeLong, P.L. Strick, Parallel organization of functionally segregated circuits linking basal ganglia and cortex, Annu Rev Neurosci, 9 (1986) 357-381.
83
[10] S.N. Haber, B. Knutson, The reward circuit: linking primate anatomy and human imaging, Neuropsychopharmacology, 35 (2010) 4-26.
[11] M. Kobayakawa, N. Tsuruya, M. Kawamura, Decision-making performance in Parkinson's disease correlates with lateral orbitofrontal volume, J Neurol Sci, 372 (2017) 232-238.
[12] W. Schultz, P. Dayan, P.R. Montague, A neural substrate of prediction and reward, Science, 275 (1997) 1593-1599.
[13] Y. Takikawa, R. Kawagoe, H. Itoh, H. Nakahara, O. Hikosaka, Modulation of saccadic eye movements by predicted reward outcome, Exp Brain Res, 142 (2002) 284-291.
[14] K. Nakamura, O. Hikosaka, Role of dopamine in the primate caudate nucleus in reward modulation of saccades, J Neurosci, 26 (2006) 5360-5369.
[15] K. Samejima, Y. Ueda, K. Doya, M. Kimura, Representation of action-specific reward values in the striatum, Science, 310 (2005) 1337-1340.
[16] B. Lau, P.W. Glimcher, Value representations in the primate striatum during matching behavior, Neuron, 58 (2008) 451-463.
[17] L.H. Tai, A.M. Lee, N. Benavidez, A. Bonci, L. Wilbrecht, Transient stimulation of distinct subpopulations of striatal neurons mimics changes in action value, Nat Neurosci, 15 (2012) 1281-1289.
[18] DA. Norman, T. Shallice, Attention to action: willed an automatic control of behavior. Center for Human Information Processing Technical Report no. 99(1980).
[19] A. Baddeley, G. Hitch, Working memory. In The Psychology of Learning and Motivation. Edited by Bower IGA. New York:Academic Press, (1974):47-90.
[20] A. Baddeley, S.D. Sala, Working memory and executive control. Philos Trans R
84
Soc Lond B (1996), 351:1397-1404.
[21] T. Shallice, Specific impairments of planning. Philos Trans R Soc Lond B Biol Sci (1982) 298:199-209.
[22] C. Padoa-Schioppa, X. Cai, The orbitofrontal cortex and the computation of subjective value: consolidated concepts and new perspectives, Ann N Y Acad Sci, 1239 (2011) 130-137.
[23] C. Padoa-Schioppa, J.A. Assad, Neurons in the orbitofrontal cortex encode economic value, Nature, 441 (2006) 223-226.
[24] X. Cai, C. Padoa-Schioppa, Neuronal encoding of subjective value in dorsal and ventral anterior cingulate cortex, J Neurosci, 32 (2012) 3791-3808.
[25] K. Samejima, K. Doya, Multiple representations of belief states and action values in corticobasal ganglia loops, Ann N Y Acad Sci, 1104 (2007) 213-228.
[26] S. Nonomura, Y. Fujiwara-Tsukamoto, T. Kajihara, F. Fujiyama, Y. Isomura, Continuous membrane potential fluctuations in motor cortex and striatum neurons during voluntary forelimb movements and pauses, Neurosci Res, 120 (2017) 53-59.
第2章
[1] J. Neumann, O. Morgenstern, Theory of Games and Economic Behavior, Princeton Univ Press, Princeton, NJ (1944).
[2] R.S. Sutton, A.G. Barto, Reinforcement learning: An introduction, MIT Press, Cambridge MA USA (1998).
[3] A. Rangel, C. Camerer, P.R. Montague, A framework for studying the neurobiology of value-based decision making, Nat Rev Neurosci, 9 (2008) 545-556.
85
[4] A. Baddeley, Working memory, Curr Biol, 20 (2010) R136-140.
[5] G.E. Alexander, M.R. DeLong, P.L. Strick, Parallel organization of functionally segregated circuits linking basal ganglia and cortex, Annu Rev Neurosci, 9 (1986) 357-381.
[6] A.M. Graybiel, S.L. Rauch, Toward a neurobiology of obsessive-compulsive disorder, Neuron, 28 (2000) 343-347.
[7] J.W. Mink, The Basal Ganglia and involuntary movements: impaired inhibition of competing motor patterns, Arch Neurol, 60 (2003) 1365-1368.
[8] C. Beste, C. Saft, J. Andrich, R. Gold, M. Falkenstein, Stimulus-response compatibility in Huntington's disease: a cognitive-neurophysiological analysis, J Neurophysiol, 99 (2008) 1213-1223.
[9] E.H. Yeterian, G.W. Van Hoesen, Cortico-striate projections in the rhesus monkey:
the organization of certain cortico-caudate connections, Brain Res, 139 (1978) 43-63.
[10] L.D. Selemon, P.S. Goldman-Rakic, Longitudinal topography and interdigitation of corticostriatal projections in the rhesus monkey, J Neurosci, 5 (1985) 776-794.
[11] A.W. Flaherty, A.M. Graybiel, Two input systems for body representations in the primate striatal matrix: experimental evidence in the squirrel monkey, J Neurosci, 13 (1993) 1120-1137.
[12] S.N. Haber, Corticostriatal circuitry, Dialogues Clin Neurosci, 18 (2016) 7-21.
[13] C. Padoa-Schioppa, Neurobiology of economic choice: a good-based model, Annu Rev Neurosci, 34 (2011) 333-359.
[14] P. Cisek, Making decisions through a distributed consensus, Curr Opin Neurobiol, 22 (2012) 927-936.
[15] M.L. Platt, P.W. Glimcher, Neural correlates of decision variables in parietal
86
cortex, Nature, 400 (1999) 233-238.
[16] L.P. Sugrue, G.S. Corrado, W.T. Newsome, Matching behavior and the representation of value in the parietal cortex, Science, 304 (2004) 1782-1787.
[17] M.R. Roesch, C.R. Olson, Impact of expected reward on neuronal activity in prefrontal cortex, frontal and supplementary eye fields and premotor cortex, J
[18] K. Matsumoto, W. Suzuki, K. Tanaka, Neuronal correlates of goal-based motor selection in the prefrontal cortex, Science, 301 (2003) 229-232.
Neurophysiol, 90 (2003) 1766-1789.
[19] Y. Takikawa, R. Kawagoe, H. Itoh, H. Nakahara, O. Hikosaka, Modulation of saccadic eye movements by predicted reward outcome, Exp Brain Res, 142 (2002) 284-291.
[20] W. Schultz, P. Dayan, P.R. Montague, A neural substrate of prediction and reward, Science, 275 (1997) 1593-1599.
[21] K. Samejima, Y. Ueda, K. Doya, M. Kimura, Representation of action-specific reward values in the striatum, Science, 310 (2005) 1337-1340.
[22] B. Lau, P.W. Glimcher, Value representations in the primate striatum during matching behavior, Neuron, 58 (2008) 451-463.
[23] L.H. Tai, A.M. Lee, N. Benavidez, A. Bonci, L. Wilbrecht, Transient stimulation of distinct subpopulations of striatal neurons mimics changes in action value, Nat Neurosci, 15 (2012) 1281-1289.
[24] C. Padoa-Schioppa, J.A. Assad, Neurons in the orbitofrontal cortex encode economic value, Nature, 441 (2006) 223-226.
[25] C. Padoa-Schioppa, Range-adapting representation of economic value in the orbitofrontal cortex, J Neurosci, 29 (2009) 14004-14014.
87
[26] X. Cai, C. Padoa-Schioppa, Contributions of orbitofrontal and lateral prefrontal cortices to economic choice and the good-to-action transformation, Neuron, 81 (2014) 1140-1151.
[27] J. Lauwereyns, Y. Takikawa, R. Kawagoe, S. Kobayashi, M. Koizumi, B. Coe, M.
Sakagami, O. Hikosaka, Feature-based anticipation of cues that predict reward in monkey caudate nucleus, Neuron, 33 (2002) 463-473.
[28] H.C. Cromwell, W. Schultz, Effects of expectations for different reward magnitudes on neuronal activity in primate striatum, J Neurophysiol, 89 (2003) 2823-2838.
[29] T. Aosaki, H. Tsubokawa, A. Ishida, K. Watanabe, A.M. Graybiel, M. Kimura, Responses of tonically active neurons in the primate's striatum undergo systematic changes during behavioral sensorimotor conditioning, J Neurosci, 14 (1994) 3969-3984.
[30] O. Hikosaka, M. Sakamoto, S. Usui, Functional properties of monkey caudate neurons. I. Activities related to saccadic eye movements, J Neurophysiol, 61 (1989) 780-798.
[31] S.N. Haber, B. Knutson, The reward circuit: linking primate anatomy and human imaging, Neuropsychopharmacology, 35 (2010) 4-26.
[32] X. Cai, S. Kim, D. Lee, Heterogeneous coding of temporally discounted values in the dorsal and ventral striatum during intertemporal choice, Neuron, 69 (2011) 170-182.
[33] H. Yamada, N. Matsumoto, M. Kimura, History- and current instruction-based coding of forthcoming behavioral outcomes in the striatum, J Neurophysiol, 98 [34] K. Samejima, K. Doya, Multiple representations of belief states and action values
88
in corticobasal ganglia loops, Ann N Y Acad Sci, 1104 (2007) 213-228.(2007) 3557-3567.
[35] I. Divac, H.E. Rosvold, M.K. Szwarcbart, Behavioral effects of selective ablation of the caudate nucleus, J Comp Physiol Psychol, 63 (1967) 184-190.
[36] R. Levy, H.R. Friedman, L. Davachi, P.S. Goldman-Rakic, Differential activation of the caudate nucleus in primates performing spatial and nonspatial working memory tasks, J Neurosci, 17 (1997) 3870-3882.
[37] M. Sakagami, K. Tsutsui, The hierarchical organization of decision making in the primate prefrontal cortex, Neurosci Res, 34 (1999) 79-89.
[38] J. Lauwereyns, M. Sakagami, K. Tsutsui, S. Kobayashi, M. Koizumi, O. Hikosaka, Responses to task-irrelevant visual features by primate prefrontal neurons, J
Neurophysiol, 86 (2001) 2001-2010.
[39] M.H. Schieber, Inactivation of the ventral premotor cortex biases the laterality of motoric choices, Exp Brain Res, 130 (2000) 497-507.
[40] Y. Nakayama, T. Yamagata, J. Tanji, E. Hoshi, Transformation of a virtual action plan into a motor plan in the premotor cortex, J Neurosci, 28 (2008) 10287-10297.
[41] D. Thura, P. Cisek, Deliberation and commitment in the premotor and primary motor cortex during dynamic decision making, Neuron, 81 (2014) 1401-1416.
[42] H.F. Kim, O. Hikosaka, Distinct basal ganglia circuits controlling behaviors guided by flexible and stable values, Neuron, 79 (2013) 1001-1010.
第 3 章
[1] C.B. Boulay, X.Y. Chen, J.R. Wolpaw, Electrocorticographic activity over
sensorimotor cortex and motor function in awake behaving rats, J Neurophysiol, 113 (2015) 2232-2241.
89
[2] F.S. Chance, L.F. Abbott, A.D. Reyes, Gain modulation from background synaptic input, Neuron, 35 (2002) 773-782.
[3] D. Chen, E.E. Fetz, Characteristic membrane potential trajectories in primate sensorimotor cortex neurons recorded in vivo, J Neurophysiol, 94 (2005) 2713-2725.
[4] E.V. Evarts, Relation of pyramidal tract activity to force exerted during voluntary movement, J Neurophysiol, 31 (1968) 14-27.
[5] Y. Fujiwara-Tsukamoto, Y. Isomura, M. Imanishi, T. Ninomiya, M. Tsukada, Y.
Yanagawa, T. Fukai, M. Takada, Prototypic seizure activity driven by mature hippocampal fast-spiking interneurons, J Neurosci, 30 (2010) 13679-13689.
[6] P. Gao, A.M. Hattox, L.M. Jones, A. Keller, H.P. Zeigler, Whisker motor cortex ablation and whisker movement patterns, Somatosens Mot Res, 20 (2003) 191-198.
[7] A.P. Georgopoulos, J.F. Kalaska, R. Caminiti, J.T. Massey, On the relations between the direction of two-dimensional arm movements and cell discharge in primate motor cortex, J Neurosci, 2 (1982) 1527-1537.
[8] N. Ho, A. Destexhe, Synaptic background activity enhances the responsiveness of neocortical pyramidal neurons, J Neurophysiol, 84 (2000) 1488-1496.
[9] J. Igarashi, Y. Isomura, K. Arai, R. Harukuni, T. Fukai, A theta-gamma oscillation code for neuronal coordination during motor behavior, J Neurosci, 33 (2013) 18515-18530.
[10] Y. Isomura, R. Harukuni, T. Takekawa, H. Aizawa, T. Fukai, Microcircuitry coordination of cortical motor information in self-initiation of voluntary movements, Nat Neurosci, 12 (2009) 1586-1593.
[11] Y. Isomura, T. Takekawa, R. Harukuni, T. Handa, H. Aizawa, M. Takada, T.Fukai, Reward-modulated motor information in identified striatum neurons,
90
J Neurosci, 33 (2013) 10209-10220.
[12] A.N. Iwaniuk, I.Q. Whishaw, On the origin of skilled forelimb movements, Trends Neurosci, 23 (2000) 372-376.
[13] T. Kajihara, M.N. Anwar, M. Kawasaki, Y. Mizuno, K. Nakazawa, K. Kitajo, Neural dynamics in motor preparation: From phase-mediated global computation to amplitude-mediated local computation, Neuroimage, 118 (2015) 445-455.
[14] A. Khorasani, N. Heydari Beni, V. Shalchyan, M.R. Daliri, Continuous Force Decoding from Local Field Potentials of the Primary Motor Cortex in Freely Moving Rats, Sci Rep, 6 (2016) 35238.
[15] R. Kimura, A. Saiki, Y. Fujiwara-Tsukamoto, F. Ohkubo, K. Kitamura, M.
Matsuzaki, Y. Sakai, Y. Isomura, Reinforcing operandum: rapid and reliable learning of skilled forelimb movements by head-fixed rodents, J Neurophysiol, 108 (2012) 1781-1792.
[16] R. Kimura, A. Saiki, Y. Fujiwara-Tsukamoto, Y. Sakai, Y. Isomura, Large-scale analysis reveals populational contributions of cortical spike rate and synchrony to behavioural functions, J Physiol, 595 (2017) 385-413.
[17] O. Kjaerulff, O. Kiehn, Distribution of networks generating and coordinating locomotor activity in the neonatal rat spinal cord in vitro: a lesion study, J Neurosci, 16 (1996) 5777-5794.
[18] Y. Koshimizu, S.X. Wu, T. Unzai, H. Hioki, T. Sonomura, K.C. Nakamura, F.
Fujiyama, T. Kaneko, Paucity of enkephalin production in neostriatal striosomal neurons: analysis with preproenkephalin-green fluorescent protein transgenic mice, Eur J Neurosci, 28 (2008) 2053-2064.
[19] V. Navarro, M. Le Van Quyen, J. Martinerie, D. Rudrauf, M. Baulac, C. Menini,
91
Loss of phase synchrony in an animal model of partial status epilepticus, Neuroscience, 148 (2007) 304-313.
[20] M. Matsumura, Intracellular synaptic potentials of primate motor cortex neurons during voluntary movement, Brain Res, 163 (1979) 33-48.
[21] B.K. Murphy, K.D. Miller, Multiplicative gain changes are induced by excitation or inhibition alone, J Neurosci, 23 (2003) 10040-10051.
[22] G. Paxinos., C. Watson., The Rat Brain in Stereotaxic Coordinates, 6th ed.
Academic Press, London (2007)
[23] J.F. Poulet, C.C. Petersen, Internal brain state regulates membrane potential synchrony in barrel cortex of behaving mice, Nature, 454 (2008) 881-885.
[24] A. Saiki, R. Kimura, T. Samura, Y. Fujiwara-Tsukamoto, Y. Sakai, Y. Isomura, Different modulation of common motor information in rat primary and secondary motor cortices, PLoS One, 9 (2014) e98662.
[25] J. Schiemann, P. Puggioni, J. Dacre, M. Pelko, A. Domanski, M.C. van Rossum, I.
Duguid, Cellular mechanisms underlying behavioral state-dependent bidirectional modulation of motor cortex output, Cell Rep, 11 (2015) 1319-1330.
[26] T. Sippy, D. Lapray, S. Crochet, C.C. Petersen, Cell-Type-Specific Sensorimotor Processing in Striatal Projection Neurons during Goal-Directed Behavior, Neuron, 88 (2015) 298-305.
[27] J.B. 6ers, L.A. Dinardo, H. Karimnamazi, Motor and premotor mechanisms of licking, Neurosci Biobehav Rev, 21 (1997) 631-647.
[28] T. Unzai, E. Kuramoto, T. Kaneko, F. Fujiyama, Quantitative Analyses of the Projection of Individual Neurons from the Midline Thalamic Nuclei to the Striosome and Matrix Compartments of the Rat Striatum, Cereb Cortex, 27 (2017) 1164-1181.
92
[29] C. von Nicolai, G. Engler, A. Sharott, A.K. Engel, C.K. Moll, M. Siegel,
Corticostriatal coordination through coherent phase-amplitude coupling, J Neurosci, 34 (2014) 5938-5948.
[30] T. Yamashita, A. Pala, L. Pedrido, Y. Kremer, E. Welker, C.C. Petersen, Membrane potential dynamics of neocortical projection neurons driving target-specific signals, Neuron, 80 (2013) 1477-1490.
[31] E. Zagha, X. Ge, D.A. McCormick, Competing Neural Ensembles in Motor Cortex Gate Goal-Directed Motor Output, Neuron, 88 (2015) 565-577.
第 4 章
[1] S. Nonomura, Y. Fujiwara-Tsukamoto, T. Kajihara, F. Fujiyama, Y. Isomura, Continuous membrane potential fluctuations in motor cortex and striatum neurons during voluntary forelimb movements and pauses, Neurosci Res, 120 (2017) 53-59.
[2] X. Wan, H. Nakatani, K. Ueno, T. Asamizuya, K. Cheng, K. Tanaka, The neural basis of intuitive best next-move generation in board game experts, Science, 331 (2011) 341-346.
[3] X. Wan, D. Takano, T. Asamizuya, C. Suzuki, K. Ueno, K. Cheng, T. Ito, K. Tanaka, Developing intuition: neural correlates of cognitive-skill learning in caudate nucleus, J Neurosci, 32 (2012) 17492-17501.
[4] H.F. Kim, O. Hikosaka, Distinct basal ganglia circuits controlling behaviors guided by flexible and stable values, Neuron, 79 (2013) 1001-1010.
[5] E.H. Yeterian, G.W. Van Hoesen, Cortico-striate projections in the rhesus monkey:
the organization of certain cortico-caudate connections, Brain Res, 139 (1978) 43-63.
[6] L.D. Selemon, P.S. Goldman-Rakic, Longitudinal topography and interdigitation of corticostriatal projections in the rhesus monkey, J Neurosci, 5 (1985) 776-794.
93
[7] A.W. Flaherty, A.M. Graybiel, Two input systems for body representations in the primate striatal matrix: experimental evidence in the squirrel monkey, J Neurosci, 13 (1993) 1120-1137.
[8] S.N. Haber, Corticostriatal circuitry, Dialogues Clin Neurosci, 18 (2016) 7-21.
[9] K. Wunderlich, P. Dayan, R.J. Dolan, Mapping value based planning and extensively trained choice in the human brain, Nat Neurosci, 15 (2012) 786-791.
[10] K. Samejima, K. Doya, Multiple representations of belief states and action values in corticobasal ganglia loops, Ann N Y Acad Sci, 1104 (2007) 213-228.
[11] P. Cisek, Making decisions through a distributed consensus, Curr Opin Neurobiol, 22 (2012) 927-936.
94
研究実績
原著論文(学位審査の条件となる論文に*印を付けること)
[1] 野々村聡,意思決定の多様性とその神経基盤(総説),脳科学とリハビリテ
ーション, 第13巻: 29-39,2013年
[2] *Satoshi Nonomura, Yoko Fujiwara-Tsukamoto, Takafumi Kajihara, Fumino
Fujiyama and Yoshikazu Isomura, Continuous membrane potential in motor cortex
and striatum neurons during voluntary forelimb movements and pauses,
Neuroscience Research, 2017
[3] Satoshi Nonomura, Kenji Doya, Jun Tanji and Kazuyuki Samejima, Neural activity
in the striatum during cognitive-decision-making (投稿準備中)
国際会議(ポスター発表)
[4] Satoshi Nonomura, Yasuhiro. X Kato, Kazuyuki Samejima, Kenji Doya and Jun
Tanji, Poster session, Neural activity in dorsal striatum during cognitive decision
making, Tohoku University International synposium, Miyagi, 2011/1/22-24
[5] Satoshi Nonomura, Yasuhiro Kato, Kenji Doya, Jun Tanji and Kazuyuki Samejima,
Poster session, Neural activity in rostral striatum during cognitive decision making,
Society for Neuroscience, Washington DC, USA, 2011/11/12-16
[6] Satoshi Nonomura, Yasuhiro Kato, Kenji Doya, Jun Tanji and Kazuyuki Samejima,
95
Poster session, The anterior caudate nucleus contribute to comparison of reward
values before motor choice in macaque monkey, Society for Neuroeconomics,
Miami, USA, 2012/9/28-30
[7] Satoshi Nonomura, Yasuhiro Kato, Kazuyuki Samejima, Kenji Doya and Jun Tanji,
Poster session, Neural activities in the rostral striatum during comparison of option
values in the object and action space, Joint Tamagawa Caltech Lecture
Course ,Hawaii, USA, 2013/3/4-10
[8] Satoshi Nonomura, Yasuhiro Kato, Kenji Doya, Jun Tanji and Kazuyuki Samejima,
Poster session, The neuronal activity in the rostral striatum during comparing action
values of objects and movements, International symposium on Prediction and
Decision Making, Kyoto Japan, 2013/10/13-14
[9] Satoshi Nonomura, Yasuhiro Kato, Kazuyuki Samejima, Kenji Doya and Jun Tanji,
Poster session, The neuronal activity in the rostral striatum during the action value
comparison for object and movement aspect, Society for Neuroscience, California,
USA, 2013/11/9-13
国内学会・シンポジウム等(口頭発表)
[10] 野々村聡、山口良哉、鮫島和行、丹治順、口頭発表、大脳基底核の認知機
能への関与、第17回脳機能とリハビリテーション研究会学術集会、東京、
96
2010/4/18
[11] 野々村聡、加藤康広、鮫島和行、銅谷賢治、丹治順、口頭発表、認知的意
思決定における吻側線条体の神経活動、第18回脳機能とリハビリテーション 研究会学術集会、東京、2011/4/29
[12] 野々村聡、加藤康広、鮫島和行、丹治順、口頭発表、選択肢の価値の比較
に関わる大脳基底核・線条体の神経活動、第19回脳機能とリハビリテーシ ョン研究会学術集会、東京、2012/4/29
国内学会・シンポジウム等(ポスター発表)
[13] 野々村聡、加藤康広、鮫島和行、銅谷賢治、丹治順、ポスター発表、認知
的意思決定における背側線条体の神経活動、第11回夏のワークショップ、北
海道、2010/7/29
[14] Satoshi Nonomura, Kazuyuki Samejima, Kenji Doya and Jun Tanji, Poster session,
Neural Activities in the Dorsal Striatum during Cognitive Decision Making,
Neuro2010, Hyogo, September2-4, 2010
[15] Satoshi Nonomura, Yasuhiro. X Kato, Kazuyuki Samejima, Kenji Doya and Jun
Tanji, Poster session, Neural activity in dorsal striatum during cognitive
decision-making, Joint Tamagawa-Keio- Caltech Lecture Course on
Neuroeconomics,
97
Tokyo, 2010/9/8-10
[16] 鮫島和行,野々村聡,加藤康広,銅谷賢治、丹治順、ポスター発表、複数
属性情報に基づく意思決定における線条体神経活動、第11回冬のワークショ
プ、北海道、2011/1/11-13
[17] 野々村聡、加藤康広、鮫島和行、銅谷賢治、丹治順、ポスター発表、認知
的意思決定における吻側線条体の神経活動、第18回脳機能とリハビリテーシ ョン研究会学術集会、東京、2011/4/29
[18] Satoshi Nonomura, Yasuhiro. X Kato, Kazuyuki Samejima, Kenji Doya and Jun
Tanji, Poster session, Neural activity in rostral striatum during cognitive decision
making, Joint Tamagawa Caltech Lecture Course, Kyoto, 2011/6/7
[19] 野々村聡、加藤康広、鮫島和行、銅谷賢治、丹治順、ポスター発表、 Neural
activity in Dorsal Striatum during Comparing the Reward Values、2011年度包括脳
ネットワーク夏のワークショップ、兵庫、2011/8/21-24
[20] Satoshi Nonomura, Yasuhiro Kato, Kazuyuki Samejima, Kenji Doya and
Jun Tanji, Poster session, Neural activity in the anterior striatum during comparison
of reward values, 日本神経回路学会第21回全国大会(jnns2011), Okinawa,
2011/12/15-17
[21] 野々村聡、加藤康広、鮫島和行、銅谷賢治、丹治順、ポスター発表、報酬