Neurons and Perception March 17, 2009

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Neurons and Perception March 17, 2009 John Maunsell Maier, A., WIlke, M., Aura, C., Zhu, C., Ye, F.Q., Leopold, D.A. (2008) Divergence of fmri and neural signals in V1 during perceptual suppression in the awake monkey. Nature Neuroscience 11:1193-1200.

Monkey visual cortex visual cortex 1 3-10 streams 3 10-30 areas 30 100-1000 modules 30,000 10-10,000 columns 3,000,000 10 subcolumns 30,000,000 10-100 neurons 300,000,000

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Purves 2002 www.purveslab.net

Purves 2002 www.purveslab.net

Periphery: neuronal sampling

Starlight Relative photon flux Flux density (#/s/mm 2 ) 18,000 total 4,000 visible infrared visible ultraviolet 0 435 750 1000 Frequency (THz) 690 400 Wavelength (nm)

Midget and Parasol Retinal Ganglion Cells Watanabe & Rodieck, 1989 Midget 1.0 mm 3.2 mm 5.6 mm Parasol 1.0 mm 3.1 mm 5.8 mm

Sinusoidal Gratings

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Periphery: neuronal sampling neuronal sensitivity

Detection of weak flashes Hecht et al., 1942 100 subject 1 subject 2 subject 3 flashes seen (percent) 0 4.5 7.3 12.1 4.5 7.3 12.1 4.5 7.3 12.1 photons per flash

Response of Limulus optic fiber to weak flashes Hartline et al., 1947 1.0 0.8 frequency of response 0.6 0.4 1 impulse 2 impulses 0.2 3 impulses 0.0 0.6 1.0 1.6 2.5 4.0 6.3 10 16 quanta per flash

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Dynamic random dot motion Motion coherence 0.0 0.5 1.0

Periphery: correlation with neuronal sampling correlation with neuronal sensitivity Central: selective effects of ablation

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Periphery: correlation with neuronal sampling correlation with neuronal sensitivity Central: selective effects of ablation selective effects of microstimulation

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Effects of microstimulation of inferotemporal cortex on reports of seeing a face Afraz et al. 2006-80% -50% 0% 20% 50% 100% face responses 50% 0% -80% 0% 80% visual signal

Periphery: correlation with neuronal sampling correlation with neuronal sensitivity Central: selective effects of ablation selective effects of microstimulation correlation with neuronal performance

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Mean 12 Mean 12 Mean 7 Mean 7 Orientation Orientation 0 10 20 Spikes Per Stimulus 0 10 20 Spikes Per Stimulus

Variance of cat V1 neuron response Dean 1981 100 10 vary: spatial frequency contrast response variance (spikes 2 ) 1 0.1 0.01 0.01 0.1 1 10 response mean (spikes)

Relationship between response variance and mean Britten et al. 1993 1000 216 MT neurons 100 response variance 10 1 0.1 0.01 0.01 0.1 1 10 100 1000 response mean

Poisson distributions 0.4 1.25 0.3 2.5 probability 0.2 5 mean spikes 10 0.1 20 40 0.0 0 10 20 30 40 50 60 response (spikes)

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Receiver Operating Characteristic (ROC) Analysis Britten et al. 1993 1 12.8 12.8 p (preferred > criterion) 6.4 3.2 1.6 0.8 6.4 0 0 p (null > criterion) 1 40 0 0 50 100 response (spikes) 0.8 1.6 3.2 motion strength 1.0 proportion correct 0.5 1 10 motion strength 100

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Temperature change threshold for monkey thermoreceptors Johnston et al., 1973 effect of number of neurons effect of correlation 0.3 0.3 threshold temperature change (deg) ρ=0.0 n=50 n=1 4 ρ=1.0 0.5 0.0 0 4 response duration (s) 64 0.0 0 4 response duration (s) 0.0

Periphery: correlation with neuronal sampling correlation with neuronal sensitivity Central: selective effects of ablation selective effects of microstimulation correlation with neuronal performance trial-by-trial correlation with choice

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Periphery: correlation with neuronal sampling correlation with neuronal sensitivity Central: selective effects of ablation selective effects of microstimulation correlation with neuronal performance trial-by-trial correlation with choice trial-by-trial correlation with reaction time

Motion detection task fixation point Cue receptive field 0.0 coherent (500-8000 ms) Time Coherent motion (750 ms)

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Periphery: correlation with neuronal sampling correlation with neuronal sensitivity Central: selective effects of ablation selective effects of microstimulation correlation with neuronal performance trial-by-trial correlation with choice trial-by-trial correlation with reaction time correlation with perception during rivalry

Binocular rivalry

Binocular rivalry in early monkey visual cortex Logothetis et al. 1996, 1997 left eye right eye 100 animal reports null animal reports preferred percent modulated average response (spikes/s) 50 0-500 0 500-500 0 500 0 V1/V2 V4 IT time (ms)

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BOLD signals in early visual cortex match perception rather than the physical stimulus Ress & Heeger 2003 criterion trial frequency correct rejects no misses false alarms yes hits average neuronal response 0.3 plaid background noise background subject 1 hits subject 2 hits plaid background background plus target fmri response ( % BOLD) 0.0 subject 3 misses false alarms hits false alarms misses misses -0.3 0 time (s) 10 0 time (s) 10 0.15 subject 4 false alarms 0.00-0.05 V1 V2 visual areas V3 V1 V2 visual areas V3 noise

Periphery: correlation with neuronal sampling correlation with neuronal sensitivity Central: selective effects of ablation selective effects of microstimulation correlation with neuronal performance trial-by-trial correlation with choice trial-by-trial correlation with reaction time correlation with perception during rivalry generation of percepts through microstimulation

Human cortical stimulation sites causing effects Penfield and Rasmussen, 1950 Somatosensory responses were elicited primarily from the cortex adjacent to the central fissure.... Rarely, however, were sensory responses encountered at a distance greater than 1 cm from the central fissure. visual auditory equilibratory sensation in eye sensation in abdomen arrest of speech mastication neck movements head contraversion eyelid movements eye movements

Detection of Microstimulation of Human Visual Cortex Murphey et al. (2009)

Psychometric function for detecting microstimulation at a site in monkey V1 100 percent correct 50! = 5.2 µa " = 7.8 p = 1.0-0.5 0 3 6 pulse amplitude (µa) 12

Thresholds for detecting electrical microstimulation 30 V1 median 5.3 V1 7a DP V4 V2 LIP VIP MST MT 0 1 10 100 µa VP V4 TEO PIT CIT AIT

Thresholds for detecting electrical microstimulation 30 V1 median 5.3 V1 7a DP V4 V2 LIP VIP MST MT 0 1 10 100 µa VP V4 TEO PIT CIT AIT V2 median 6.4 1 10 100 µa

Thresholds for detecting electrical microstimulation 30 V3A median 8.1 0 1 10 100 µa 30 V1 median 5.3 V1 7a DP V4 V2 LIP VIP MST MT 0 1 10 100 µa VP V4 TEO PIT CIT AIT V2 median 6.4 1 10 100 µa

Thresholds for detecting electrical microstimulation 30 V3A median 8.1 25 MT median 9.2 0 1 10 100 µa 0 1 10 100 µa 30 V1 median 5.3 V1 7a DP V4 V2 LIP VIP MST MT 0 1 10 100 µa VP V4 TEO PIT CIT AIT V2 median 6.4 1 10 100 µa

Thresholds for detecting electrical microstimulation 30 V3A median 8.1 25 MT median 9.2 0 1 10 100 µa 0 1 10 100 µa 30 V1 median 5.3 V1 7a DP V4 V2 LIP VIP MST MT 0 1 10 100 µa VP V4 TEO PIT CIT AIT V2 median 6.4 IT median 10.3 1 10 100 µa 1 10 100 µa

Thresholds for detecting electrical microstimulation 30 V3A median 8.1 25 MT median 9.2 FEF median 14.4 0 1 10 100 µa 0 1 10 100 µa 30 V1 median 5.3 V1 7a DP V4 V2 LIP VIP MST MT FEF 1 10 100 µa 0 1 10 100 µa VP V4 TEO PIT CIT AIT V2 median 6.4 IT median 10.3 1 10 100 µa 1 10 100 µa