Probing the Neural Mechanism of Binocular Information Processing with VEPs Ryusuke HAYASHI, Yoichi MIYAWAKI, Taro MAEDA, and Susumu TACHI random-dot s

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1 Probing the Neural Mechanism of Binocular Information Processing with VEPs Ryusuke HAYASHI, Yoichi MIYAWAKI, Taro MAEDA, and Susumu TACHI random-dot stereogram RDS visual evoked potential VEP VEP RDS VEP RDS anti-rdsrds V1 random-dot stereogram 1. random-dot stereogram RDS [1] visual evoked potential VEP VEP RDS VEP [2] [5] RDS 200 ms VEP VEP [3] Graduate School of Engineering, The University of Tokyo, Tokyo, Japan VEP RDS D II Vol. J84 D II No. 3 pp

2 2001/3 Vol. J84 D II No. 3 [6] V1 V2 V3 V3A VP MT MST [7], [8] V1 V2 [6] MT [8] 2. 2 Marr Marr & Poggio [9] Marr Marr [10] 2. 3 interocularly unpaired region [11] [12] V [13] [14] V1 [15] [16] Stereo Optical cm SHARP XV-E ch 8ch 1ch 5 30 kω MME µv/v Hz A D 1 khz IBM PC/AT 560

3 Fig. 1 1 The experimental system. Fig. 2 2 The sequence of events in a single trial of stimulus-presentation. 50 SGI Indigo RDS 30 Hz dynamic RDS binocularly correlated random-dot stereopattern CRD dynamic 2 CRD 100 ms CRD 700 ms CRD 600 ms CRD 1 2s % 1 [pixel] pixel 160 pixel 41 cd/m RDS θ [rad] d scr [m] d eye [m] d [m] d d 2 scrθ d scrθ d eye (1) d RDS [5], [17] ( 36.3 ) RDS (O2) (Pz) 561

4 2001/3 Vol. J84 D II No. 3 (a) (b) 4 RDS Fig. 4 The latencies of the negative peaks to RDSs and monocular-cue stimuli. 3 RDS O2 Pz Fig. 3 VEPs to RDSs of a crossed disparity for subject RH (a) and SS (b), recorded at scalp sites O2 and Pz. 5 RDS Fig. 5 The latencies of the positive peaks to RDSs and monocular-cue stimuli. VEP 3 Pz ms VEP ms O2 562

5 4 RDS 5 [5], [17] RDS VEP RDS VEP dynamic CRD dynamic RDS V1 V [2], [4] V1 O RDS VEP (36.3 ) RDS ( 6 RDS O2 Fig. 6 VEPs to RDSs of crossed and uncrossed disparities for subject RH, recorded at site O2. 7 RDS Fig. 7 The latencies of the negative peaks to RDSs ) 12.1 ( ) RDS anti-rds RDS anticorrelated RDS anti-rdsanti-rds 9 RDS 563

2001/3 Vol. J84 D II No. 3 8 RDS Fig. 8 The latencies of the positive peaks to RDSs. 10 anti-rds O2 Fig. 10 VEPs to anti-rdss of crossed and uncrossed disparities for subject RH, recorded at site O2.

6 2001/3 Vol. J84 D II No. 3 8 RDS Fig. 8 The latencies of the positive peaks to RDSs. 10 anti-rds O2 Fig. 10 VEPs to anti-rdss of crossed and uncrossed disparities for subject RH, recorded at site O2. 9 anti-rds Fig. 9 An example of anti-rds. anti-rds anti-rds RDS anti-rds binocularly uncorrelated random-dot stereopattern, uncrd VEP anti-rds ( 36.3 ) (36.3 ) anti-rds O2 10 VEP RDS 2 3 RDS anti-rds VEP anti-rds anti-rds anti-rds RDS ( 36.3 ) RDS (36.3 ) RDS

7 11 anti-rds Fig. 11 The latencies of the negative peaks to anti-rdss and uncrds. 13 RDS O2 Fig. 13 VEPs to RDSs of crossed and uncrossed disparities for subject RH, recorded at site O2. The effect of stimulus size. 12 anti-rds Fig. 12 The latencies of the positive peaks to anti-rdss and uncrds. anti-rds VEP Richards [18] [19] ms ±36.3 Panum 565

8 2001/3 Vol. J84 D II No. 3 Panum (±12.1 ) ms V1 [20] V anti-rds RDS V1 anti-rds Cumming & Parker [6] anti-rds Cumming & Parker anti-rds anti-rds anti-rds VEP RDS V anti-rds VEP RDS V1 Ohzawa [21] 14 Fig. 14 A binocular energy neuron that prefers non-zero disparity. anti-rds 14 1 (2) x σ i [pixel] ω i [cycle/pixel] φ [rad] I l (x) I r(x) (E l E r) (3) x l x r [pixel] ( ) 1 g(x, φ, i)= 3 exp x2 2π 2 σ i 2σi 2 sin(2πω ix + φ) (2) E l (x l,φ,i)= I l (x)g(x x l,φ,i)dx E r(x r,φ,i)= I r(x)g(x x r,φ,i)dx (3) Simple Cell (4) Complex Cell 2 (5) 566

9 S(x l,x r,φ,i) C(x l,x r,φ,i) (x r x l ) S(x l,x r,φ,i)=e l (x l,φ,i)+e r(x r,φ,i) (4) C(x l,x r,φ,i)=s 2 (x l,x r,φ,i) +S 2 ( x l,x r,φ+ π 2,i ) (5) (6) RDS [22] Out(x l,x r)= π 2 φ=0 i C(x l,x r,φ,i) (6) 15 RDS Fig. 15 The responses of binocular energy neurons to an RDS RDS anti-rds 50 pixel 17 pixel 4 pixel 20 [0,π) 5 ((σ i,ω i)={(1, 1 ), (2, 1 ), (4, 1 1 ), (8, ), (16, )}) 5 (7) Out(x l,x r) Ôut(x l,x r)= max xl (Out(x l,x r)) Out(x l,x r) (7) max xr (Out(x l,x r)) RDS 15 x l x r Ôut(x l,x r) anti-rds RDS 16 anti-rds Fig. 16 The responses of binocular energy neurons to an anti-rds. anti-rds 567

10 2001/3 Vol. J84 D II No uncrd Fig. 17 The responses of binocular energy neurons to an uncrd [12] [15] [1] 18 Fig. 18 A model for the detection mechanism of interocularly unpaired regions. McLoughlin & Grossberg [11] anti-rds [15] anti-rds anti-rds RDS 5. 1 anti-rds anti-rds 568

11 Pz O2 19 Fig. 19 A model framework for stereoscopic depth perception and binocular rivalry. [17] 6. RDS VEP [1] B. Julesz, Binocular depth perception without familiarity cues, Science, vol.145, pp , [2] B. Julesz, W. Kropfl, and B. Petrig, Large evoked potentials to dynamic random-dot correlograms and stereograms permit quick determination of stereopsis, Proc. Natl. Acad. Sci. USA, vol.77, no.4, pp , [3] B. Fenelon, R.A. Neill, and C.T. White, Evoked potentials to dynamic random dot stereograms in upper, center and lower fields, Doc. Ophthalmol., vol.63, pp , [4] Dynamic random-dot pattern disparity VEP, vol.88, no.3, pp , [5] 2 D-II vol.j82-d-ii, no.5, pp , May [6] B.G. Cumming and A.J. Parker, Responses of primary visual cortical neurons to binocular disparity without depth perception, Nature, vol.389, pp , [7] G.F. Poggio, F. Gonzalez, and F. Krause, Stereoscopic mechanisms in monkey visual cortex: Binocular correlation and disparity selectivity, J. Neurosci., vol.8, no.12, pp , [8] G.C. DeAngelis, B.G. Cumming, and W.T. Newsome, Cortical area MT and the perception of stereoscopic depth, Nature, vol.394, pp , [9] [10] 1 pp.15 48, [11] N. McLoughlin and S. Grossberg, Cortical computation of stereo disparity, Vision Res., vol.38, no.1, pp.91 99, [12] K. Nakayama and S. Shimojo, Da Vinci stereop- 569

2001/3 Vol. J84 D II No. 3 sis: Depth and subjective occluding contours from unpaired image points, Vision Res., vol.30, no.11, pp.1811 1825, 1990. [13] R.

Nature, vol.380, pp.621 624, 1996. [15] F. Sengpiel, C. Blakemore, and R. Harrad, Interocular suppression in the primary visual cortex: A possible neural basis of binocular rivalry, Vision Res., vol.35, no.

Fenelon, Scalp response topography to dynamic random dot stereograms, Electroencephalogr. Clin. Neurophysiol., vol.69, pp.209 217, 1988. [18] W. Richards, Anomalous stereoscopic depth perception, J.

Day, Temporal integration differences between crossed and uncrossed stereoscopic mechanisms, Percept. Psychophys., vol.57, no.6, pp.891 897, 1995. [20] G.S. Masson, C. Busettini, and F.A.

12 2001/3 Vol. J84 D II No. 3 sis: Depth and subjective occluding contours from unpaired image points, Vision Res., vol.30, no.11, pp , [13] R. Blake, A neural theory of binocular rivalry, Psychol. Rev., vol.96, no.1, pp , [14] N.K. Logothetis, D.A. Leopold, and D.L. Sheinberg, What is rivalling during binocular rivalry? Nature, vol.380, pp , [15] F. Sengpiel, C. Blakemore, and R. Harrad, Interocular suppression in the primary visual cortex: A possible neural basis of binocular rivalry, Vision Res., vol.35, no.2, pp , [16] R. Blake, Y. Yang, and H.R. Wilson, On the coexistence of stereopsis and binocular rivalry, Vision Res., vol.31, nos.7/8, pp , [17] R.A. Neill and B. Fenelon, Scalp response topography to dynamic random dot stereograms, Electroencephalogr. Clin. Neurophysiol., vol.69, pp , [18] W. Richards, Anomalous stereoscopic depth perception, J. Opt. Soc. Am., vol.61, no.3, pp , [19] R. Patterson, R. Cayko, G.L. Short, R. Flanagan, L. Moe, E. Taylor, and P. Day, Temporal integration differences between crossed and uncrossed stereoscopic mechanisms, Percept. Psychophys., vol.57, no.6, pp , [20] G.S. Masson, C. Busettini, and F.A. Miles, Vergence eye movements in response to binocular disparity without depth perception, Nature, vol.389, pp , [21] I. Ohzawa, G.C. DeAngelis, and R.D. Freeman, Stereoscopic depth discrimination in the visual cortex: Neurons ideally suited as disparity detectors, Science, vol.249, pp , [22] D.J. Fleet, H. Wagner, and D.J. Heeger, Neural encoding of binocular disparity: Energy models, position shifts and phase shifts, Vision Res., vol.36, no.12, pp , SFN ARVO IEEE SFN MIT IEEE/EMBS IMEKO IMEKO SICE 9 11 ARVO, IEEE, 570

2 The Characteristics of Two Negative Peaks on Visual Evoked Potentials with Depth Perception Yoichi MIYAWAKI, Yasuyuki YANAGIDA, Taro MAEDA, and Susu

2 The Characteristics of Two Negative Peaks on Visual Evoked Potentials with Depth Perception Yoichi MIYAWAKI, Yasuyuki YANAGIDA, Taro MAEDA, and Susumu TACHI Random-Dot Stereogram 200 ms 40 180 ms 280