2015/9 Vol. J98 D No. 9 Shidara [7] t s t V (s t)=e[r t+1 + γr t+2 + γ 2 r t+3 + ] (1) r t t E γ 0 1 V (s t) TD V new(s t 1) V

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1 a) b) Modeling the Function of the Ventral Striatum in Reinforcement Learning Based on the Analysis of Neuronal Activity Masanari SHINOTSUKA a), Masahiko MORITA b), and Munetaka SHIDARA TD striosome striosome 1. [1] Schultz TD Graduate School of Systems and Information Engineering, University of Tsukuba, Tennodai, Tsukuba-shi, Japan Faculty of Engineering, Information and Systems, University of Tsukuba, Tennodai, Tsukuba-shi, Japan Faculty of Medicine, University of Tsukuba, Tennodai, Tsukuba-shi, Japan a) m.shinotsuka2@gmail.com b) mor@bcl.esys.tsukuba.ac.jp DOI: /transinfj.2014JDP7137 [2] Barto [3] Doya [4] striosome V (s) striosome striosome [5], [6] D Vol. J98 D No. 9 pp c

2 2015/9 Vol. J98 D No. 9 Shidara [7] t s t V (s t)=e[r t+1 + γr t+2 + γ 2 r t+3 + ] (1) r t t E γ 0 1 V (s t) TD V new(s t 1) V old (s t 1)+αδ t 1 (2) δ t 1 δ t 1 = r t + γv (s t) V (s t 1) (3) TD temporal differencetd 1 Fig. 1 Neural circuits of the basal ganglia. (1) (3) 0 TD basal ganglia cerebral cortex 1 (striatum) striosome matrix striosome (DA cell) matrix (internal segment of globus pallidus, GPi) (substantia nigra pars reticulata, SNr) GPi/SNr (thalamus) Schultz [2] 1278

3 TD striosome matrix striosome matrix striosome [8] [9] Hebb TD matrix striosome actor critic Barto [3] matrix Q(s, a) Doya [4] striosome 2 striosome s t V (s t) striosome TD striosome TD striosome 2 Fig. 2 Structure common to conventional reinforcement learning models of the basal ganglia. striosome striosome [10], [11] Cromwell [11] Shidara [7] [12] Goldstein [5] Kim [6] 1279

4 2015/9 Vol. J98 D No. 9 striosome 2. 2 Shidara [7] A Wait Go OK B / / B 3C 1/1, 1/2, 2/2, 1/3, 2/3, 3/3 1/1, 2/2, 3/3 1 1/2 1/2, 1/3, 2/3 1/6 1/ Shidara [7] Fig. 3 Multiple trial reward schedule task (adapted from Shidara et al. [7]). 1 Shidara [7] Table 1 Response in the cue condition (adapted from Shidara et al. [7]). 1/3 1/2 2/3 3/3 2/2 1/1 n (1) 16 (2) 13 (3) 6 (4) 3 (5)

5 [12] 3. 1 (1) (2) (1) (2) 2/3, 3/3, 2/2 1/3, 1/2, 1/1 Shidara σ = ms 200 ms 1000 ms 0 90% Fig. 4 Response period. 5 Fig. 5 Histogram of the response onset time /26 [13], [14] 100 ms ms

6 2015/9 Vol. J98 D No. 9 6 Fig. 6 Classification diagram of history dependence for the ventral striatum neurons. 1 1/2 1/3 2/ % n = n 1 (1) (5) (1) (2) (1) 8 (2) 7 (3) (5) ms Shidara striosome

7 Fig. 7 7 Structure of the proposed model. 8 Fig. 8 Network output to the test sequence. Elman [15] t cue t cue t r t+1 1 1/2 1/3 2/ Elman TD δ t 1 = r t + γo t O t 1 (4) 0 O t t r t 0 1 TD r t O t γ /2 2/2 1/3 2/3 3/

8 2015/9 Vol. J98 D No (a) 2 F (1, 190) = 34.1 p <0.01; 2 F (1, 190) = 19.1 p < Fig. 9 Classification diagram of history dependence for the middle elements of the model (b) F (1, 190) = 4.99 p< (a): F (1, 145) = 15.9 p<0.01; 2 F (1, 145) = 4.21 p <0.05, (b): F (1, 227) = 4.36 p < Fig Example of the response of middle elements to a random sequence. Fig Example of the response of ventral striatum neurons in the random condition. 1284

9 Fig Correspondence of the proposed model to the brain structure. 13 Fig. 13 State values estimated from the internal state F (5, 193) = 3.53 p <0.01 2/2 3/3 2/3 3/3 vs 2/3 t(70) = 4.41 p <0.01 2/2 vs 2/3 t(60) = 2.6 p<0.011/1, 1/2, 1/3 [10], [11] 1 1 V V

10 2015/9 Vol. J98 D No. 9 [12] TD V V Q matrix [16] TD 1 1 TD (B) , 1286

11 , [1] R.S. Sutton and A.G. Barto, Reinforcement Learning, MIT Press, [2] W. Schultz, P. Dayan, and P.R. Montague, A neural substrate of prediction and reward, Science, vol.275, pp , 1997 [3] A.G. Barto, Adaptive critics and the basal ganglia, in Models of Information Processing in the Basal Ganglia, ed. J.C. Houk, J.L. Davis, and D.G. Beiser, pp , MIT Press, [4] K. Doya, Complementary roles of basal ganglia and cerebellum in learning and motor control, Current Opinion in Neurobiology, vol.10, no.6, pp , [5] B.L. Goldstein, B.R. Barnett, G. Vasquez, S.C. Tobia, V. Kashtelyan, A.C. Burton, D.W. Bryden, and M.R. Roesch, Ventral striatum encodes past and predicted value independent of motor contingencies, Journal of Neuroscience, vol.32, pp , [6] Y.B. Kim, N. Huh, H. Lee, E.H. Baeg, D. Lee, and M.W. Jung, Encoding of action history in the rat ventral striatum, J. Neurophysiology, vol.98, pp , [7] M. Shidara, T.G. Aiger, and B.J. Richmond, Neuronal signals in the monkey ventral striatum related to progress through a predictable series of trials, J. Neuroscience, vol.18, pp , [8] C.R. Gerfen, The neostriatal mosaic: Multiple levels of compartmemtal organization in the basal ganglia, Annual Review of Neuroscience, vol.15, pp , [9] J.N.J. Reynolds, B.I. Hyland, and J.R. Wickens, A cellular mechanism of reward-related learning, Nature, vol.413, pp.67 70, [10] W. Schultz, P. Apicella, E. Scarnati, and T. Ljungberg, Neuronal activity in monkey ventral striatum related to the expectation of reward, Journal of Neuroscience, vol.12, pp , [11] H.C. Cromwell and W. Schultz, Effects of expectations for different reward magnitudes on neuronal activity in primate striatum, J. Neurophysiology, vol.89, pp , [12] vol.25, no.4, pp , [13] Z. Liu and B.J. Richmond, Response differences in monkey TE and perirhinal cortex: Stimulus association related to reward schedules, J. Neurophysiology, vol.83, pp , [14] Y. Naya, M. Yoshida, and Y. Miyashita, Forward processing of long-term associative memory in monkey inferotemporal cortex, J. Neuroscience, vol.23, pp , [15] J.L. Elman, Finding structure in time, Cognitive Science, vol.14, pp , [16] Y. Sawatsubashi, M.F.B. Samusudin, and K. Shibata, Emergence of discrete and abstract state representation in continuous input task through reinforcement learning, Advances in Intelligent Systems and Computing, vol.208, pp.13 22,

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