Neurons and Perception March 17, 2009

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1 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:

2 Monkey visual cortex visual cortex streams areas modules 30, ,000 columns 3,000, subcolumns 30,000, neurons 300,000,000

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7 Periphery: neuronal sampling

8 Starlight Relative photon flux Flux density (#/s/mm 2 ) 18,000 total 4,000 visible infrared visible ultraviolet Frequency (THz) Wavelength (nm)

9 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

10 Sinusoidal Gratings

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12 !"" #$%$&'(#),+-"" #$%$&'(#) *!+!"" #$%$&'(#) *-"+""" #$%$&'(#)

13 Periphery: neuronal sampling neuronal sensitivity

14 Detection of weak flashes Hecht et al., subject 1 subject 2 subject 3 flashes seen (percent) photons per flash

15 Response of Limulus optic fiber to weak flashes Hartline et al., frequency of response impulse 2 impulses impulses quanta per flash

16 !"#$%&'()'&*)+",'-.%$'()*"/"0/.%&)%1)23.&).&*"&/'/.%& 4'((+%)5)6%,'&22%&)789: 7== 2;.3")1.$"* +",'-.%$'()*"/"0/.%& ;"$0"&/)%1.&*"&/'/.%&2 *"/"0/"* = = B= <= 23.&).&*"&/'/.%&)>?@A

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20 Dynamic random dot motion Motion coherence

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

23 A&B%";&$C$D+/EF$8GHH 9&%,"2$:&;,.,&-! <"2#/"-$:&;,.,&-! 9&%,"2$:&;,.,&-! ;"#,"2$3":&/&23&$(I* '/& '"%# ;"#,"2$3":&/&23&$(I* '/& '"%# 3"2#/+%#$(I* 86 8 '/& '"%# !"#$%#&' (!&)*!"#$%#&' (!&)* %'+#,+-$./&01&234 (3435!&)*

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

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26 !"#$%"&'()'"#*'+,,-

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

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

29 #! E2>,7F/5(/2,"/=GH= '()*+,+-./0/1+23( / 8-6)95-:-; $! %! &!!!"!!"&!"% <)*5/8-;!"$!"#

30 #! E2>,7F/5(/2,"/=GH= '()*+,+-./0/1+23( / 8-6)95-:-; $! %! &!!!"!!"&!"% <)*5/8-;!"$!"#

31 Mean 12 Mean 12 Mean 7 Mean 7 Orientation Orientation Spikes Per Stimulus Spikes Per Stimulus

32 Variance of cat V1 neuron response Dean vary: spatial frequency contrast response variance (spikes 2 ) response mean (spikes)

33 Relationship between response variance and mean Britten et al MT neurons 100 response variance response mean

34 Poisson distributions probability mean spikes response (spikes)

35 56+,43/#D34E/,84#F6//# #+8#G63:#H/30I60# J3>/8K#6+#3/%#!LM!# *+,-./.01#"#2.34+3# &$#!$$#>676+,+,840# $%)# $#2.34+3# "#2.34+3# ## 907,:60;0<# '$# ($# )$# 8A#B## 07,:60#,4## )$$#-0# $%!# $# $%$# $%)# $%(# C,-6#90<# $%'# $%&# $# $# "#!$# B#!"#

36 !"#"$%"&'()"&*+$,-'#.*&*#+"&$/+$#'0!123'*,*45/$/' 6*&4(7'"+'*48'9:;9' )&()(&+$(,'<'=>*,+*'?'#&$+"&$(,' 9' <' 9' <' <' A&(B*B$4$+5'(C'D'' /)$E"/'$,'F<<'G/' <'=>*,+*' <' <89' <8F' )&()(&+$(,'<'=>*,+*'?'#&$+"&$(,' 9' <' <' <'=>*,+*' <' <89' <8F' )&()(&+$(,'<'=>*,+*'?'#&$+"&$(,' 9' <' <' <'=>*,+*' <' <89' <8F' )&()(&+$(,'<'=>*,+*'?'#&$+"&$(,' 9' <' <' <'=>*,+*' <' <89' <8F'

37 56+,43/#D34E/,84#F6//# #+8#G63:#H/30I60# J3>/8K#6+#3/%#!LM!# *+,-./.01#"#2.34+3# &$#!$$#>676+,+,840# $%)# $#2.34+3# "#2.34+3# ## 907,:60;0<# '$# ($# )$# 8A#B## 07,:60#,4## )$$#-0# $%!# $# $%$# $%)# $%(# C,-6#90<# $%'# $%&# $# $# "#!$# B#!"# 56O6,P6>#Q76>3+,4E#FI3>3O+6>,0+,O#95QF<# 7>878>+,84# "#2.34+3#N#O>,+6>,84# $# $#!# 7>878>+,84#$#2.34+3#N#O>,+6>,84#

38 Receiver Operating Characteristic (ROC) Analysis Britten et al p (preferred > criterion) p (null > criterion) response (spikes) motion strength 1.0 proportion correct motion strength 100

39 !"##$%&"'()'*+,$-./)"#.0+.'(+%)'*0$+1')2+&$#3"#4.'5$+6789 :#)22$'+$2+.0;+6<==>9 <;? A? C#"&"#2)"' 5"##$52?;E?;A '$1#"' 4"'F$G B14,$# "3 '$1#"'% 7"2)"'+5"-$#$'5$??;< < <? C$#3"#4.'5$+#.2)" 6'$1#"'D,$-./)"#9

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

42 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

43 C#*..$(7$.70127DEE '9$#$(8$!678'9$#$(8$ '9$#$(8$ &#'&'#.*'( '/.#*01%!2"!2! 3!2"! "!! "!! "! #$%&'(%$ )%&*+$%,%- &#$/$##$:7:$8*%*'(% (;117:$8*%*'(%

44 !"#$%&'()#*+*$,$-.'/$-"'0-$12,20'#33'-"&')&%&(-$4&'3$&,5 6)$--&7'&-'+,8'9::; <8==9 ()#(#)-$#7 #3 7&2)#70 #7')&%&(-$4&'3$&,5 <8> <8= <8? #33')&%&(-$4&'3$&,5 ()#(#)-$#7 #3 7&2)#70 <8> <8= <8? %"#$%&'()#*+*$,$-.

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46 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

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

48 8.19,:1.17,;7<=7:.>-,:7/,70,/$,:7,:1./!"! +,&.-.:/70,/$,: #$%& '!"()!* <,/$,: 1/-.:%/& <.2$>0 '!"A()* 0$11.2 +,--.+/ 6)!!! )!! /$0.7'01*

49 !"#$%&#"#'%(')*'+&,'-./'&"01%&#'2%11"3+4"'5647'1"+246%&'468" =>C =>= 2%7"1"&2" )* -./ 8",608 2%7"1"&2" =>C =>= 2%7"1"&2" 8",608 2%7"1"&2" 2%11"3+46%& 5647'!* =>= ;=>C ;=>B =>= ;=>C ;=>B =>D ;=>A &EFA ;=>A &EC=G ;<== = <== 468"'98#: ;<== = <== 468"'98#:

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51 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

52 Binocular rivalry

53 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) V1/V2 V4 IT time (ms)

54 !"#$%&!%#'()*+,%'-%./0)-%12 3&*&-45,6%7*)586%7+)/-6%.8898+%:;;;

55 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 time (s) 10 0 time (s) subject 4 false alarms V1 V2 visual areas V3 V1 V2 visual areas V3 noise

56 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

57 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

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

59

60 Psychometric function for detecting microstimulation at a site in monkey V1 100 percent correct 50! = 5.2 µa " = 7.8 p = pulse amplitude (µa) 12

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

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

63 Thresholds for detecting electrical microstimulation 30 V3A median µa 30 V1 median 5.3 V1 7a DP V4 V2 LIP VIP MST MT µa VP V4 TEO PIT CIT AIT V2 median µa

64 Thresholds for detecting electrical microstimulation 30 V3A median MT median µa µa 30 V1 median 5.3 V1 7a DP V4 V2 LIP VIP MST MT µa VP V4 TEO PIT CIT AIT V2 median µa

65 Thresholds for detecting electrical microstimulation 30 V3A median MT median µa µa 30 V1 median 5.3 V1 7a DP V4 V2 LIP VIP MST MT µa VP V4 TEO PIT CIT AIT V2 median 6.4 IT median µa µa

66 Thresholds for detecting electrical microstimulation 30 V3A median MT median 9.2 FEF median µa µa 30 V1 median 5.3 V1 7a DP V4 V2 LIP VIP MST MT FEF µa µa VP V4 TEO PIT CIT AIT V2 median 6.4 IT median µa µa

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Signal detection theory. Direct comparison of perception and neural activity. ? Choice probability. Visual stimulus. Behavioral judgement

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