Lesions of prefrontal cortex reduce attentional modulation of neuronal responses. and synchrony in V4

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Lesions of prefrontl ortex reue ttentionl moultion of neuronl responses n synhrony in V4 Georgi G. Gregoriou,, Anrew F. Rossi, 3 Leslie G Ungerleier, 4 Roert Desimone 5 Deprtment of Bsi Sienes, Fulty of Meiine, University of Crete, Institute of Applie n Computtionl Mthemtis, Fountion for Reserh n Tehnology, Hells, Herklion, Crete, 73 Greee 3 Division of Neurosiene n Bsi Behviorl Siene, NIMH, NIH, Bethes, MD 89, USA. 4 Lortory of Brin n Cognition, NIMH, NIH, Bethes, MD 89, USA 5 MGovern Institute for Brin Reserh, Msshusetts Institute of Tehnology (MIT), Cmrige, MA 39, USA. Supplementry Informtion 6 Figures Nture Neurosiene: oi:.38/nn.374

Sensory intertion (pir stim) Sensory intertion (pir stim) CONTROL HEMISPHERE Atten wy Atten stim Atten stim monkey monkey Slope.53 r =.7 Seletivity (stim stim) Slope.7 r =.67 Seletivity (stim stim) Slope.38 r =.4 Seletivity (stim stim) LESION HEMISPHERE Atten wy e Atten stim f Atten stim Slope.55 r =.64 Seletivity (stim stim) Slope.7 r =.74 Seletivity (stim stim) Slope.5 r =.6 Seletivity (stim stim) Supplementry Figure. Reltionship etween seletivity n sensory intertions n effet of ttention on sensory intertions. () Reltionship etween sensory intertion inies (siss, SI) n seletivity inies (orinte, SE) with ttention wy from the reeptive fiel, in the intt hemisphere. () Sme neuronl popultion s in ut with ttention irete to stimulus. Responses to the pir show greter influene of stimulus, s reflete y the inrese in the slope of the regression line tht esries the SE, SI reltionship (Atten wy, slope:.53 Atten stim, slope:.7). () Sme neuronl popultion s in n ut with ttention irete to stimulus. In this se responses to the pir show greter influene of stimulus, s reflete y the erese in the slope of the regression line tht esries the SE, SI reltionship (Atten wy, slope:.53 Atten stim, slope:.38). () Reltionship etween sensory intertion inies n seletivity inies with ttention wy from the reeptive fiel, in the lesion-ffete hemisphere. (e) Sme neuronl popultion s in ut with ttention irete to stimulus. Responses to the pir show greter influene of stimulus, s reflete y the inrese in the slope of the regression line tht esries the SE, SI reltionship (Atten wy, slope:.55 Atten stim, slope:.7). (f) Sme neuronl popultion s in e n f ut with ttention irete to stimulus. In this se responses to the pir show greter influene of stimulus, s reflete y the erese in the slope of the regression line tht esries the SE, SI reltionship (Atten wy, slope:.55 Atten stim, slope:.5). All visully responsive neurons re inlue, eh ontriuting one t point. Green irles orrespon to t from monkey n lue irles to t from monkey. Slopes n orreltion oeffiients (r) re reporte for eh grph. Responses were ompute within winow from 5 to 3 ms following stimulus onset. Nture Neurosiene: oi:.38/nn.374

Atten in (sp/se) Atten in (sp/se) 8 CONTROL 8 LESION 6 6 4 4 4 6 8 Atten out (sp/se) monkey monkey 4 6 8 Atten out (sp/se) Supplementry Figure. Distriution of ttentionl effets on firing rte in V4. Distriution of men normlize firing rte verge in winow 5-3 ms following the rry onset with ttention outsie (y-xis) n ttention insie (x-xis) the RF of the reore neurons in the ontrol () n lesion () hemisphere. Points ove the igonl inite positive ttentionl effet (higher response with ttention) wheres points elow the igonl inite negtive ttentionl effet (lower responses with ttention). Eh point represents one neuron. Green irles orrespon to t from monkey n lue irles to t from monkey. Nture Neurosiene: oi:.38/nn.374

Power tten in (5-3 Hz) Power tten in (5-9 Hz) CONTROL 8 LESION 8 6 6 4 4 monkey monkey 4 6 8 Power tten out (5-9 Hz) 4 6 8 Power tten out (5-9 Hz) 4 3 3 3 4 Power tten out (5-3 Hz) 3 Power tten out (5-3 Hz) Supplementry Figure 3. Distriution of ttentionl effets on LFP power in V4. (,) Distriution of men LFP gmm power (verge etween 5-9 Hz) lulte within winow -3 ms following the rry onset with ttention outsie (y-xis) n ttention insie (x-xis) the RF in the ontrol () n lesion () hemisphere. (,) Distriution of men LFP et power (verge etween 5-3 Hz) lulte within winow -3 ms following the rry onset with ttention insie (y-xis) n ttention outsie (x-xis) the RF in the ontrol () n lesion () hemisphere. Points ove the igonl inite positive ttentionl effet (higher power with ttention) wheres points elow the igonl inite negtive ttentionl effet (lower power with ttention). Other onventions s in Supplementry Figure. Nture Neurosiene: oi:.38/nn.374

Coherene tten in (5-3 Hz) Coherene tten in (5-3Hz) Coherene tten in (5-9 Hz) Coherene tten in (5-9Hz) CONTROL LESION.4.5.3.4..3.....3.4. monkey monkey...3.4.5 Coherene tten out (5-9 Hz) Coherene tten out (5-9 Hz).3.3.......3 Coherene tten out (5-3 Hz)...3 Coherene tten out (5-3 Hz) Supplementry Figure 4. Distriution of ttentionl effets on spike-lfp oherene in V4. (,) Distriution of men gmm spike-lfp oherene (verge etween 5-9 Hz) lulte within winow -4 ms following the rry onset with ttention insie (y-xis) n ttention outsie (x-xis) the RF in the ontrol () n lesion () hemisphere. (,) Distriution of men et oherene (verge etween 5-3 Hz) lulte within winow -4 ms following the rry onset with ttention insie (y-xis) n ttention outsie (x-xis) the RF in the ontrol () n lesion () hemisphere. Points ove the igonl inite positive ttentionl effet (higher oherene with ttention) wheres points elow the igonl inite negtive ttentionl effet (lower oherene with ttention). Other onventions s in Supplementry Figure. Nture Neurosiene: oi:.38/nn.374

Proportion of signls Proportion of signls.8.8.8.6.6.6.4.4.4....4...4.6 AI FR Corret intt Corret lesion Inorret intt Inorret lesion.8.6.4.4.4 AIoh gmm.8.6.4... AIpower gmm e..4.4 AIoh et.... AIpower et Supplementry Figure 5. Distriution of ttentionl effets on firing rtes, LFP power n spike-lfp oherene in V4 in orret n error trils. () Cumultive istriutions of ttentionl inies ompute for firing rtes in the ontrol n lesion-ffete hemisphere for orret n error trils. (,) Cumultive istriutions of ttentionl inies for gmm SFC verge over 5-9 Hz () n et SFC verge over 5-3 Hz () for the ontrol n lesion-ffete hemisphere in orret n inorret trils. () Cumultive istriutions of ttentionl inies ompute for gmm power in the ontrol n lesion-ffete hemisphere in orret n inorret trils. (e) Cumultive istriutions of ttentionl inies ompute for et power in the ontrol n lesion-ffete hemisphere in orret n inorret trils. In ll plots re lines illustrte t from the ontrol hemisphere, lue lines t from the lesion-ffete hemisphere, soli lines t from orret trils n she lines t from inorret trils. Nture Neurosiene: oi:.38/nn.374

Coherene Proportion of signls Normlize power (5-3 Hz) Proportion of signls Normlize power (5-9 Hz) Proportion of signls Normlize firing rte Proportion of signls CONTROL n= 5....3.4.5.5 n= 78.5..4.6.4..8 n= 78..4.4.. g j LESION n= 6....3.4.5 Atten insie RF Atten outsie RF.5 e.5..4.6.4..8.4..4...4.6 Supplementry Figure 6. Effet of ttention on firing rtes, gmm n et LFP power n oherene in the ontrol n lesion-ffete hemispheres limiting nlysis to trils from onitions with similr performne in the two hemifiels. () Normlize popultion verge firing rtes from the ontrol hemisphere. At the popultion level, tivity ws enhne with ttention y 8.5%, 5-3 ms poststimuli onset (6.9%, -3 ms post-stimuli onset; Wiloxon sign-rnk test, p <. in oth ses). () Normlize popultion verge firing rtes from the lesion-ffete hemisphere. At the popultion level, tivity ws enhne with ttention y 9.7%, 5-3 ms, post-stimuli onset (.4%, -3 ms poststimuli, Wiloxon sign-rnk test, p <. in oth ses) () Cumultive istriutions of ttentionl inies ompute from firing rtes for the ontrol (soli line) n lesion-ffete (she line) hemisphere. Attentionl moultion of firing rtes ws signifintly lower in the lesion ffete sie (5-3 ms poststimuli: ontrol hemisphere: mein AIFR =.9, lesion-ffete hemisphere: mein AIFR =.5, Wiloxon rnk-sum test p <.; -3 ms post-stimuli: ontrol hemisphere: mein AIFR =.9, lesion-ffete hemisphere: mein AIFR =.4, Wiloxon rnk-sum test p <.). (,e) Normlize LFP n= 9 n= 9..4.8 n= 336 n= 58 4 6 8 4.8 4 6 8 4 Frequeny (Hz) Frequeny (Hz). h k.8.6.4..8.6.4 f AI FR Control Lesion. Control Lesion... AIpower gmm.8.6.4 i. Control Lesion.... AIpower et.8.6.4 l. Control Lesion.4...4.6 AIoh gmm Nture Neurosiene: oi:.38/nn.374

gmm power verge etween 5-9 Hz in the ontrol () n lesion-ffete (e) hemisphere. At the popultion level, we foun signifint ttentionl enhnement of LFP power in gmm frequenies etween 5 n 9 Hz in oth hemispheres (verge power -3 ms, 5-35 ms post-stimuli onset, pire t-test, p <. in ll four ses). Gmm power ws enhne y 4.5% in the intt hemisphere n y 7% in the lesion-ffete hemisphere (.6% n 7%, respetively, 5-35 ms post-stimuli onset). (f) Cumultive istriutions of ttentionl inies in gmm power for the ontrol (soli line) n lesionffete (she line) hemisphere. Attentionl inies (AIpowergmm) were signifintly lower in the lesion-ffete hemisphere (-3 ms post-stimuli: mein AIpowergmm ontrol:.7, lesion:.3; 5-35 ms post-stimuli: mein AIpowergmm ontrol:.6, lesion:.3; Wiloxon rnk-sum test p <. in oth ses). (g,h) Normlize LFP et power verge etween 5-3 Hz in the ontrol (g) n lesionffete (h) hemisphere. Bet power ws signifintly reue with ttention in the ontrol hemisphere (verge power -3 ms n 5-35 ms post stimuli onset, pire t-test, p <. in oth ses; - 3 ms post stimuli onset: 3.9% erese, 5-35 ms post stimuli onset: 3% erese with ttention t the popultion level) n ws not signifintly moulte y ttention in the lesion-ffete hemisphere (pire t-test, -3 ms post stimuli onset: p=.7,.4% inrese with ttention t the popultion level; 5-35 ms post stimuli onset: p=.8, % inrese with ttention). (i) Cumultive istriutions of ttentionl inies in et power for the ontrol (soli line) n lesion-ffete (she line) hemisphere. Attentionl inies (AIpoweret) were signifintly lower in the ontrol hemisphere (-3 ms post stimuli onset: mein AIpoweret ontrol: -., lesion:.; 5-35 ms post stimuli onset: mein AIpoweret ontrol: -.3, lesion:.6; Wiloxon rnk-sum test p <.). (j,k) Spike-LFP oherene (SFC) in the ontrol (j) n lesion-ffete (k) hemisphere. SFC ws signifintly enhne with ttention in the gmm frequeny rnge (5-9 Hz) in oth hemipsheres (ontrol hemisphere:.6% inrese with ttention, pire t-test, p <.; lesion-ffete hemisphere: 6% inrese with ttention, pire t-test, p <.). Moreover, et SFC ws signifintly reue with ttention in the ontrol hemisphere n the effet ws only mrginl in the lesion-ffete hemisphere (ontrol: % erese with ttention, pire t-test p <.; lesion-ffete hemisphere: 5.8% erese with ttention, pire t-test p =.65). (l) Cumultive istriution of ttentionl inies for gmm oherene (verge etween 5 n 9 Hz) in the ontrol (soli line) n lesion (she line) hemisphere. The ttentionl moultion of oherene etween 5 n 9 Hz ws signifintly lower in the lesion-ffete hemisphere ompre to the intt hemisphere (Wiloxon rnk-sum test, p <.5, mein AIohgmm, ontrol:.8, lesion:.). Responses in,,,e,g,h,j,k re ligne on the presenttion of the stimuli rry. Re lines orrespon to responses when the trget stimulus ppere insie the RF of the reore neurons n the lk line orrespons to responses when the trget stimulus ppere outsie the RF. She res represent men ± s.e.m. Nture Neurosiene: oi:.38/nn.374