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Umbono wesisombululo esiphakamileyo ufuna isampulu yeretina ecolekileyo kunye nodityaniso lokwakha ngokutsha iipropati zento.Kubalulekile ukuba uqaphele ukuba xa udibanisa iisampuli zendawo ezivela kwizinto ezahlukeneyo, ukuchaneka kulahlekile.Ke ngoko, ukwahlulahlula, ukwahlulahlula iindawo zomfanekiso ukuze kuqwalaselwe ngokwahlukeneyo, kubalulekile ekubonweni.Kumsebenzi wangaphambili, izakhiwo ze-bistable lattice, ezinokuthi zithathwe njengenye okanye ngaphezulu kwendawo ehambayo, yayisetyenziselwa ukufunda le nkqubo.Apha, sixela ubudlelwane phakathi komsebenzi kunye nezigwebo zezahlulo kwimimandla ephakathi kwendlela yokubonwa kweprimate.Ngokukodwa, siye safumanisa ukuba ngokukhetha ukuqhelanisa i-neuron zexesha eliphakathi kwakunovakalelo kwiindlela zokuthungwa ezisetyenziselwa ukugqwetha imbono ye-bistable gratings kwaye ibonise ulungelelwaniso olubalulekileyo phakathi kwezilingo kunye nembono ezimeleyo ye-stimuli eqhubekayo.Olu lungelelaniso lukhulu kwiiyunithi ezibonisa intshukumo yehlabathi kwiipateni ezinezalathiso ezininzi zasekhaya.Ke, sigqiba kwelokuba indawo yexesha eliphakathi iqulethe imiqondiso esetyenziselwa ukwahlula imiboniso entsonkothileyo kwizinto eziyinxalenye kunye nemigangatho.
Umbono awuxhomekanga kuphela kucalucalulo oluchanekileyo lweempawu zomfanekiso osisiseko ezifana nokuqhelaniswa nomphetho kunye nesantya, kodwa kubaluleke ngakumbi ekudityanisweni okuchanekileyo kwezi mpawu ukubala iimpawu zokusingqongileyo ezifana nokumila kwento kunye netrajectory1.Nangona kunjalo, iingxaki zivela xa imifanekiso ye-retinal ixhasa amaqela amaninzi afanelekileyo ngokulinganayo 2, 3, 4 (Fig. 1a).Ngokomzekelo, xa iiseti ezimbini zemiqondiso yesantya zisondele kakhulu, oku kunokutolikwa ngokufanelekileyo njengento enye ehambayo okanye izinto ezininzi ezihambayo (Umfanekiso 1b).Oku kubonisa ubume bokwahlulahlula, okt asiyonto isisigxina yomfanekiso, kodwa yinkqubo yokutolika.Ngaphandle kokubaluleka kwayo okucacileyo kwimbono eqhelekileyo, ukuqonda kwethu isiseko se-neural sokwahlulahlula ngokwengqiqo kuhlala kungagqibekanga kokona kulungileyo.
Umzobo wekhathuni wengxaki yokwahlulwa kokuqonda.Imbono yomkhi-mkhanyo ngobunzulu kwityhubhu ye-Necker (ekhohlo) itshintshana phakathi kweengcaciso ezimbini ezinokubakho (ekunene).Oku kungenxa yokuba akukho miqondiso emfanekisweni evumela ingqondo ukuba imisele ngokukodwa umzobo we-3D we-3D (ebonelelwa ngumqondiso we-monocular occlusion signal ekunene).b Xa imiqondiso eshukumayo emininzi ibonakaliswe kufutshane nendawo, inkqubo yombono kufuneka imisele ukuba iisampulu zengingqi zivela kwinto enye okanye ngaphezulu.Ukungacaci kakuhle kweempawu zentshukumo zasekhaya, oko kukuthi, ulandelelwano lwezindululo zento inokuvelisa isindululo sasekhaya esifanayo, okukhokelela kutoliko oluphindaphindekayo ngokulinganayo lwegalelo elibonakalayo, oko kukuthi amabala eVektha apha anokuvela kwintshukumo ehambelanayo yomphezulu omnye okanye intshukumo ecacileyo yemigangatho ethe kratya.c (ekhohlo) Umzekelo wovuselelo lwethu lwegridi eyenziweyo.Iigrayiti ezijikelezayo ezijikelezayo zihamba ngokuthe ngqo kwicala lazo ("izalathiso zecandelo" - iintolo ezimhlophe) zidibanisana ukuze zenze ipateni yokugaya.I-lattice inokubonwa njengento enye, eqhelekileyo, edityanisiweyo yentshukumo yezalathiso (iintolo ezibomvu) okanye intshukumo ecacileyo yezalathiso ezihlanganisiweyo.Umbono we-lattice uphazamiseka ngokongezwa kweempawu ze-random texture.(Embindini) Indawo ephawulwe ngomthubi iyandiswa kwaye iboniswe njengoluhlu lwezakhelo zemiqondiso ehambelanayo neselubala, ngokulandelelanayo.Ukuhamba kwechaphaza kwimeko nganye imelwe ngeentolo eziluhlaza kunye nobomvu.(Ekunene) Igrafu yendawo (x, y) yendawo yokukhetha ngokuchasene nenani lezakhelo.Kwimeko ehambelanayo, zonke ii-texture zikhukuliseka kwicala elinye.Kwimeko yokungafihli, ukuthungwa kuhamba kwicala lecandelo.d Umzobo wekhathuni womsebenzi wethu wokwahlulahlula intshukumo.Iinkawu zaqalisa ulingo ngalunye ngokulungisa ichaphaza elincinane.Emva kolibaziseko olufutshane, uhlobo oluthile lwepateni yokugrayitha (ukuhambelana/ukungafihli) kunye nobukhulu bomqondiso wokuthungwa (umzekelo, umahluko) wavela kwindawo yeMT RF.Ngexesha lovavanyo ngalunye, igrayiti inokukhukuliseka kwelinye lamacala amabini anokwenzeka epateni.Emva kokurhoxiswa kovuselelo, iithagethi zokukhetha zavela ngasentla nangaphantsi kweMT RF.Iinkawu kufuneka zibonise imbono yazo yegridi kwi-saccades kwithagethi yokukhetha efanelekileyo.
Ukusetyenzwa kweentshukumo ezibonakalayo kubonakaliswe kakuhle kwaye ngaloo ndlela kubonelela ngemodeli egqwesileyo yokufunda iisekethe ze-neural zokwahlulahlula okubonwayo.Izifundo ezininzi zokubala ziye zaqaphela ukusetyenziswa kweemodeli zokulungiswa kwezinyathelo ezimbini apho uqikelelo lokuqala lwesisombululo esiphezulu lulandelwa ukuhlanganiswa okukhethiweyo kweesampuli zendawo ukususa ingxolo kunye nokubuyisela into ye-velocity7,8.Kubalulekile ukuqaphela ukuba iinkqubo zombono kufuneka zithathele ingqalelo ukunciphisa le ensemble kuphela kwezo sampuli zasekhaya ezivela kwizinto eziqhelekileyo.Izifundo zengqondo zichaze izinto zomzimba ezichaphazela indlela imiqondiso yentshukumo yendawo eyahlulwe ngayo, kodwa imilo yeetrajethi ze-anatomical kunye neekhowudi ze-neural zihlala ziyimibuzo evulekileyo.Iingxelo ezininzi zibonisa ukuba iiseli ezikhethiweyo zokukhetha kwi-temporal (MT) ye-primate cortex ngabaviwa be-neural substrates.
Okubalulekileyo, kolu vavanyo lwangaphambili, utshintsho kwimisebenzi ye-neural ehambelana nokutshintsha komzimba kwi-stimuli ebonakalayo.Nangona kunjalo, njengoko kukhankanyiwe ngasentla, ukwahlulahlula kuyinkqubo yokuqonda.Ngoko ke, ukufundwa kwe-neural substrate yayo kufuna ukudibanisa utshintsho kumsebenzi we-neural kunye neenguqu kwimbono ye-stimuli esisigxina.Ke ngoko, siye saqeqesha iinkawu ezimbini ukuba zinike ingxelo yokuba ingaba ipateni yokugayiwa kwe-bistable eyenziwe ngokukhukuliseka okungaphezulu koxande yayingumphezulu omnye okanye imiphezulu emibini ezimeleyo.Ukufunda ubudlelwane phakathi komsebenzi we-neural kunye nezigwebo zezahlulo, sibhale umsebenzi omnye kwi-MT xa iinkawu zenza lo msebenzi.
Sifumene ulungelelwaniso olubalulekileyo phakathi kokufunda umsebenzi weMT kunye nokuqonda.Olu lungelelaniso belukhona nokuba isivuseleli siqulathe iimpawu zokwahlulahlula okuthe kratya.Ukongezelela, amandla esi siphumo sinxulumene novakalelo kwiimpawu zezahlulo, kunye nesalathiso sepateni.Le yokugqibela ilinganisa iqondo apho iyunithi isasaza kwihlabathi kunentshukumo yasekhaya kwiipateni ezinzima.Nangona ukukhethwa kolwalathiso lwefashoni sele kuqondwa njengento ecacileyo ye-MT, kwaye iiseli ezikhethiweyo zefashoni zibonisa ukulungelelaniswa kwe-stimuli eyinkimbinkimbi ehambelana nombono womntu wezo zivuseleli, ngokolwazi lwethu, obu bubungqina bokuqala bokulungelelaniswa phakathi kweepateni.isalathiso kunye nokwahlulahlula ngokwengqiqo.
Siqeqeshe iinkawu ezimbini ukubonisa imbono yazo yegrid evuselelayo (iintshukumo ezihambelanayo okanye ezicacileyo).Abakhi-mkhanyo abangabantu badla ngokubona ezi zivuseleli njengeentshukumo ezihambelanayo okanye ezicacileyo eziphantse zibe ngamaza afanayo.Ukuze unikeze impendulo echanekileyo kolu vavanyo kwaye usete isiseko somvuzo osebenzayo, senze izibonakaliso zecandelo ngokubhala i-raster yecandelo elenza i-lattice (Umfanekiso 1c, d).Ngaphantsi kweemeko ezihambelanayo, zonke izinto eziphathekayo zihamba ecaleni kwendlela yomzekelo (Umfanekiso 1c, "uhambelana").Kwimeko ecacileyo, i-texture ihamba nge-perpendicular ukuya kwicala le-grating apho ifakwe khona (umzobo 1c, "i-transparent").Silawula ubunzima bomsebenzi ngokutshintsha umahluko wale leyibhile yokuthungwa.Kwiimvavanyo ezichongiweyo, iinkawu zavuzwa ngenxa yeempendulo ezihambelana nokuthungwa, kwaye imivuzo yanikezelwa ngokungaqhelekanga (i-50/50 odds) kwizilingo eziqulethe iipatheni ngaphandle kweempawu ze-texture (zero texture difference condition).
Idatha yokuziphatha evela kwiimvavanyo ezimbini zommeli ziboniswa kuMfanekiso 2a, kwaye iimpendulo zicwangciswe njengenxalenye yezigwebo zokubambisana ngokuchasene nokuchasana kweempawu zokuthungwa (ukuchasana kokungafihli kucingelwa ukuba kubi ngenkcazo) kwiipateni ezihamba phezulu okanye phantsi, ngokulandelanayo. Ngokubanzi, imbono yeenkawu zokuhambelana / ukungafihli kwachatshazelwa ngokuthembekileyo kuzo zombini uphawu (olucacileyo, oluhambelanayo) kunye namandla (uchasi) lwe-texture cue (ANOVA; inkawu N: isalathiso - F = 0.58, p = 0.45, uphawu - F = 1248, p <10−1 = 1, 2, mon-1, 2, 2, 2, 2, 2, 2, 2, 2, 3, 2, 2, 3, 10 = 10 = 1, 2, 2, 2, 2. S: isalathiso - F = 0.41, p = 0.52, uphawu - F = 2876.7, p <10-10, umahluko - F = 36.5, p <10−10). Ngokubanzi, imbono yeenkawu zokuhambelana / ukungafihli kwachatshazelwa ngokuthembekileyo kuzo zombini uphawu (olucacileyo, oluhambelanayo) kunye namandla (uchasi) we-texture cue (ANOVA; inkawu N: isalathiso - F = 0.58, p = 0.45, uphawu - F = 1248, p <10−1 = 1, 2, 2, 2, 2, 2, p <10 - 3, 2, 2, 2, mon - 1, 2; isitshixo S: ulwalathiso - F = 0.41, p = 0.52, uphawu - F = 2876.7, p <10-10, umahluko - F = 36.5, p <10-10). В целом на восприятие обезьянами когерентности/прозрачности достоверно влияли как знак (прозрачность, когерентность), так исток (кон) обезьяна N: направление — F = 0,58, p = 0,45, знак — F = 1248, p < 10−10, контраст – F = 22,63, p < 10; −10 обезьнак - 10 = F, p, p = 4, 2, p, p = ризнак - F = 2876,7, p <10−10, контраст - F = 36,5, р <10-10). Ngokuqhelekileyo, imbono yokubambisana / ukungafihli kweenkawu kwakuchatshazelwa kakhulu zombini isibonakaliso (ukungafihli, ukuhambelana) kunye namandla (ukuchasana) kwesicatshulwa sombhalo (ANOVA; inkawu N: ulwalathiso - F = 0.58, p = 0.45, uphawu - F = 1248, p <10-= 3 <0, i-10, i-monkey , i-10, i-1, i-1, i-monkey, i-1, i-1, i-1, i-1, i-1, i-monkey, i-1, i-1, i-1, i-monkey, i-1, i-1, i-1. F = 0.41, p = 0.52, uphawu - F = 2876.7, p <10 -10, umahluko - F = 36.5, p <10-10).总体而言，猴子对连贯性/透明度的感知受到纹理提示(ANOVA）的符号)靠影响；猴子N: 方向- F = 0.58，p = 0.45, 符号- F = 1248, p <10−10, 对比度– F = 22.63, p <10;–10 = F = 10;–10: S = 52, 符号– F = 2876.7, p <10−10, 对比度– F = 36.5,p <10-10）.总体而言，猴子对连贯性/透明度的感知受到纹理提示(ANOVA）的符号)靠影响；猴子N: 方向- F = 0.58,p = 0.45, 符号- F = 1248, p <10−10, 对比度– F = 22.63, p <10;−0010;−10 p.Ngokuqhelekileyo, imbono yokubambisana kweenkawu / ukungafihli kwachatshazelwa kakhulu ngumqondiso (ukukhanya, ukuhambelana) kunye nokuqina (ukuchasana) kweempawu zokuthungwa (ANOVA);обезьяна N: ориентация – F = 0,58, p = 0,45, знак – F = 1248, p <10−10, Контрастность — F = 22,63, p <10; inkawu N: i-orientation - F = 0.58, p = 0.45, uphawu - F = 1248, p <10-10, Ukuchasana - F = 22.63, p <10; −10 Обезьяна S: Ориентация — F = 0,41, p = 0,52, Знак — F = 2876,7, p < 10−10, Контрастность — F = 36,5, p < 10-10). −10 I-Monkey S: Ukuqhelaniswa - F = 0.41, p = 0.52, Uphawu - F = 2876.7, p <10-10, Ukuchasana - F = 36.5, p <10-10).Imisebenzi ye-Gaussian cumulative yafakwa kwidatha yeseshoni nganye ukuze ibonise iimpawu ze-psychophysical zeenkawu.Kwikhiwane.I-2b ibonisa ukusasazwa kwesivumelwano kule mifuziselo kuzo zonke iiseshoni kuzo zombini iinkawu.Ngokubanzi, iinkawu zawugqiba umsebenzi ngokuchanekileyo nangokungaguquguqukiyo, kwaye sakhaba ngaphantsi kwe-13% yeeseshoni zeenkawu ezimbini ngenxa yokungangeni kakuhle kwimodeli yeGaussian eqokelelweyo.
imizekelo yokuziphatha kweenkawu kwiiseshoni ezimele (n ​​≥ izilingo ze-20 kwimeko yokuvuselela).Kwiiphaneli zasekhohlo (ekunene), idatha evela kwiseshoni yenkawu ye-N(S) icwangciswe njengamanqaku okhetho ahambelanayo (ulungelelwaniso) ngokuchasene nophawu olwahlukileyo lwemiqondiso yendlela (abscissa).Apha kucingelwa ukuba izinto ezicacileyo (ezihambelanayo) zinexabiso elibi (elilungileyo).Iimpendulo zakhiwe ngokwahlukileyo ngokwesikhokelo sokuhamba kwepateni (phezulu (90 °) okanye phantsi (270 °)) kuvavanyo.Kuzo zombini izilwanyana, ukusebenza, nokuba impendulo yahlulahlulwe ngo-50/50 umahluko (PSE - iintolo eziqinileyo) okanye ubungakanani bokuchasana okubhaliweyo okufunekayo ukuxhasa umgangatho othile wokusebenza (umda - iintolo ezivulekileyo), kule migaqo ye-drift.b Ifakelwe i-histogram yee-R2 zamaxabiso omsebenzi oqokelelweyo we-Gaussian.Idatha yeMonkey S(N) iboniswa ekhohlo (ekunene).c (Phezulu) I-PSE elinganisiweyo yegridi itshintshelwe ezantsi (ulungelelwaniso) xa kuthelekiswa ne-PSE yatshintshela phezulu igridi (i-abscissa) ecwangcisiweyo, enemiphetho emele unikezelo lwe-PSE kwimeko nganye kunye neentolo ezibonisa intsingiselo yemeko nganye.Idatha yazo zonke iiseshoni ze-N (S) zenkawu zinikwe kwikholamu yasekhohlo (ekunene).(Ezantsi) Ingqungquthela efanayo neyedatha ye-PSE, kodwa yeempawu ezifakiweyo.Kwakungekho nantlukwano ibalulekileyo kwimida ye-PSE okanye iintsingiselo zefashoni (jonga isicatshulwa).d I-PSE kunye ne-slope (i-ordination) zicwangciswe ngokuxhomekeke kwi-raster ye-normalized orientation ye-angular separation component ("integral grating angle" - abscissa).Izangqa ezivulekileyo ziyindlela, umgca oqinileyo ngowona mzekelo ufanelekileyo wokubuyisela umva, kwaye umgca onamachaphaza yi-95% yexesha lokuzithemba kwimodeli yokubuyisela.Kukho ulungelelwaniso olubalulekileyo phakathi kwe-PSE kunye ne-engile yokudityaniswa okuqhelekileyo, kodwa hayi ukuthambeka kunye ne-angle yokudibanisa eqhelekileyo, ecebisa ukuba umsebenzi we-psychometric uyatshintsha njengoko i-engile isahlula iileyile zecandelo, kodwa hayi ukulola okanye ukucaba.(I-Monkey N, n = iiseshoni ze-32; i-Monkey S, n = iiseshoni ze-43).Kuzo zonke iipaneli, iibar zempazamo zibonisa imposiso eqhelekileyo yentsingiselo.Haha.Ukuyondelelana, iPSE inqaku elilinganayo elilinganayo, eliqhelekileyo.ukubekwa emgangathweni.
Njengoko kuphawuliwe ngasentla, zombini umahluko weempawu zokuthungwa kunye nesalathiso sokuhamba kwepateni zahluka kwizilingo, kunye ne-stimuli ekhukuliseka phezulu okanye ezantsi kwisilingo esinikiweyo.Oku kwenziwa ukunciphisa i-psychophysical11 kunye ne-neuronal28 adaptive effects.Ipatheni yokuqhelaniswa necala (inqaku elilinganayo lokulinganisa okanye i-PSE) (i-Wilcoxon yenqanaba lovavanyo lwesambuku; inkawu N: z = 0.25, p = 0.8; inkawu S: z = 0.86, p = 0.39) okanye i-threshold efakiweyo (i-sum of Wilcoxon irenki: 1, monkey = 4, monkey = 4, monkey; z = 0.49, p = 0.62) (Umfanekiso 2c).Ukongezelela, kwakungekho mahluko abalulekileyo phakathi kweenkawu kwiqondo lokungafani kokuthungwa okufunekayo ukuze kugcinwe amanqanaba omgangatho wokusebenza (N monkey = 24.5% ± 3.9%, i-S monkey = 18.9% ± 1.9%; i-Wilcoxon isamba senqanaba, z = 1.01, p = 0.31).
Kwiseshoni nganye, sitshintshe i-angle ye-interlattice eyahlula ukuqhelaniswa kweeleti zecandelo.Uphononongo lwePsychophysical lubonise ukuba abantu banokubona iseli ye-10 njengedityanisiweyo xa le engile incinci.Ukuba iinkawu zazixela ngokuthembekileyo imbono yazo yokuhambelana / ukungafihli, ngoko ngokusekwe kwezi ziphumo, umntu unokulindela i-PSE, umahluko wokuthungwa ohambelana nokwahlulwa okufanayo phakathi kokuhambelana kunye nokukhetha ukungafihli, ukwandisa phezu kokunxibelelana.i-engile yelathisi. Oku kwakunjalo ngokwenene (umzobo we-2d; ukuwa ngapha kweendlela zepateni, i-Kruskal-Wallis; inkawu N: χ2 = 23.06, p <10-3; inkawu S: χ2 = 22.22, p <10-3; ulungelelwaniso phakathi kwe-normalized inter-grating −3, i-angle ye-7, i-P, i-P, i-P, i-6, i-P, i-P. 9; inkawu S: r = 0.76, p <10−13). Oku kwakunjalo ngokwenene (Umfanekiso we-2d; ukudilika kwiindlela zepateni, i-Kruskal-Wallis; inkawu N: χ2 = 23.06, p <10-3; inkawu S: χ2 = 22.22, p <10-3; ukulungelelaniswa phakathi kwe-normalized inter- 6-0 i-angle = i-P-0 −3; i-angle ye-Normon-1 = i-P (i-7-0, i-P, i-P, i-P, i-P) 9; inkawu S: r = 0.76, p <10−13). Это действительно имело место (рис. 2d; коллапс поперек направления паттерна, Крускал-Уоллис; обезьяна N: χ2 = 23,06, p <2,2,2,3; p. < 10–3 корреляция между нормализованными угол решетки и PSE – обезьяна N: r = 0,67, p < 10-9, обезьяна S: r = 0,76-1, p. Oku kwenzeka ngokwenene (umzobo we-2d; ukuwa ngaphesheya kolwalathiso lwepateni, i-Kruskal-Wallis; inkawu N: χ2 = 23.06, p <10-3; inkawu S: χ2 = 22.22, p <10-3; ukulungelelaniswa phakathi kwe-angle ye-lattice eqhelekileyo kunye ne-PSE: 1, i-monkey N. r = 0,76, p <10-13).情况确实如此（图2d；跨模式方向折叠，Kruskal-Wallis；子子N:χ2 = 23.06，p <10-3猴子S:χ20; χ2 = 3 p.光栅角和PSE – 猴子N:r = 0.67,p <10-9；猴子S:r = 0.76,p <10-13）.情况 确实 如此 （图 图 2D ； 方向 折叠 , kruskal-wallis ; n : :2 = 23.06 , p <10-3 ;栅角 和 pse-猴子 猴子 猴子 猴子 猴子 猴子N：r = 0.67，p <10-9；猴子S:r = 0.76，p <10-13）. Это действительно имело место (рис. 2d; кратность по оси моды, Крускал-Уоллис; обезьяна N: χ2 = 23,06, p <10-3; обез,3,2,2,2,3; лизованный межрешеточный угол). Oku kwakunjalo ngokwenene (Umfanekiso 2d; phinda ulandele i-axis yemowudi, i-Kruskal-Wallis; inkawu N: χ2 = 23.06, p <10-3; inkawu S: χ2 = 22.22, p <10-3; ikona ye-interlattice eqhelekileyo). PSE-обезьяна N: r = 0,67, p < 10–9, обезьяна S: r = 0,76, p < 10–13). PSE monkey N: r = 0.67, p <10-9, inkawu S: r = 0.76, p <10-13).Ngokwahlukileyo, ukutshintsha i-angle ye-interlattice kwakungenayo impembelelo ebalulekileyo kwi-slope yomsebenzi we-psychometric (Umfanekiso we-2d; i-cross-modal orientation fold, i-Kruskal-Wallis; i-monkey N: χ2 = 8.09, p = 0.23; inkawu S χ2 = 3.18 = i-angle ye-3.18, i-angle ephakathi kunye ne-7, i-angle ephakathi, i-7, i-p. N: r = -0.4, p = 0.2, inkawu S: r = 0.03, p = 0.76).Ke, ngokuhambelana nedatha ye-psychophysical yomntu, isiphumo esiphakathi sokutshintsha i-angle phakathi kwe-gratings kukutshintsha kwamanqaku okufuduka, kwaye hayi ukwanda okanye ukuncipha kobuntununtunu kwimiqondiso yokwahlulahlula.
Okokugqibela, imbuyekezo yabelwa ngokungacwangciswanga kunye nokuba nokwenzeka kwe-0.5 kulingo kunye ne-zero texture umahluko.Ukuba zonke iinkawu bezazi le randomness eyodwa kwaye bakwazi ukwahlula phakathi zero ukuthungwa umahluko kunye cue stimuli, babenokuphuhlisa amaqhinga ezahlukeneyo iintlobo ezimbini zolingo.Imibono emibini ibonisa ngamandla ukuba oku akunjalo.Okokuqala, ukutshintsha i-angle ye-grating kuneempembelelo ezifanayo ngokomgangatho kwi-cue kunye ne-zero texture amanqaku okungafaniyo (Umfanekiso we-2d kunye ne-Supplementary Fig. 1).Okwesibini, kuzo zombini iinkawu, ukhetho lovavanyo lwe-bistable alunakwenzeka ukuba luphinde lube yinto ephindaphindiweyo (edlulileyo) ukhetho lomvuzo (uvavanyo lwe-binomial, i-N monkeys: 0.52, z = 0.74, p = 0.22; S iinkawu: 0.51, r = 0.9, p = 0.18).
Ukuqukumbela, ukuziphatha kweenkawu kumsebenzi wethu wokwahlula kwakuphantsi kolawulo olulungileyo lokuvuselela.Ukuxhomekeka kwezigwebo zokuqonda kumqondiso kunye nobukhulu beempawu zokuthungwa, kunye notshintsho kwi-PSE nge-angle ye-grating, ibonisa ukuba iinkawu zichaze imbono yazo ye-motor coherence / transparency.Ekugqibeleni, iimpendulo zeenkawu kwizilingo zokuchasana kwe-zero texture azizange zichaphazeleke yimbali yomvuzo wezilingo zangaphambili kwaye zachatshazelwa kakhulu lutshintsho lwe-angular inter-raster.Oku kubonisa ukuba iinkawu ukuqhubeka ingxelo imbono yabo subjective uqwalaselo umphezulu ileti phantsi le meko ibalulekileyo.
Njengoko kukhankanyiwe ngasentla, inguqu yenguqu yokuthungwa ukusuka kwi-negative ukuya kwi-positive ilingana nenguqu yengqondo ye-stimuli ukusuka ekukhanyeni ukuya kwi-coherent.Ngokuqhelekileyo, kwiseli enikeziweyo, impendulo ye-MT ivame ukunyuka okanye iyancipha njengoko ukungafani kwe-texture kuguquka ukusuka kwi-negative ukuya kwi-positive, kwaye isalathiso salo mphumela ngokuqhelekileyo sixhomekeke kwindlela yokuhamba kwepateni / inxalenye.Ngokomzekelo, i-curves yokuqondisa i-curve yeeseli ezimbini ze-MT ezimele ziboniswe kwi-Figure 3 kunye neempendulo zala maseli kwi-gratings equlethe izibonakaliso eziphantsi okanye eziphezulu ezihambelanayo okanye ezibonakalayo ezibonakalayo.Sizame ngandlela thile ukulinganisa ngcono ezi mpendulo zegridi, ezinokunxulumana nokusebenza kwengqondo kweenkawu zethu.
Iploti yePolar yegophe lokuqondisa igophe lommeli wenkawu MT cell S ekuphenduleni kuluhlu olulodwa lwe-sinusoidal.I-angle ibonisa isalathiso sokuhamba kwe-grating, ubukhulu bubonisa ukukhutshwa, kunye nolwalathiso lweseli olukhethiweyo lugqithise malunga ne-90 ° (phezulu) kunye necala lelinye lamacandelo kwicala lepatheni ye-grating.b I-histogram yeveki ye-stimulus-time (PSTH) yegridi yokuphendula, itshintshelwe kwicala letemplate ngama-90 ° (eboniswe ngokucwangcisekileyo ekhohlo) kwiseli eboniswe kwi-.Iimpendulo zihlelwe ngohlobo lwengcebiso ye-texture (i-cohesive / transparent - panel ephakathi / ekunene ngokulandelelanayo) kunye ne-Michelson umahluko (i-PSTH ingcebiso yombala).Iinzame ezichanekileyo kuphela ziboniswa kuhlobo ngalunye lweempawu zokuthungwa okuphantsi kunye nokuchasana okuphezulu.Iiseli ziphendule bhetele kwiipateni zelathisi ezikhukulisekayo ezibonisa ukuthungwa okubonakalayo, kwaye impendulo kwezi patheni iye yanda ngokwanda kokuthungwa.c, d zizinto ezifanayo njengaku-a kunye no-b, kodwa kwiiseli zeMT ngaphandle kwenkawu S, ukuqhelaniswa kwazo ezikhethwayo kuphantse ukugqithe kwegridi ehlayo.Iyunithi ikhetha i-gratings ezantsi ekhukulisekayo eneempawu zokuthungwa ezihambelanayo, kwaye impendulo kule pateni iyanda ngokungafaniyo kokuthungwa.Kuzo zonke iipaneli, indawo enomthunzi imele impazamo eqhelekileyo yentsingiselo.Izithethi.Izikhonkwane, imizuzwana.Isibini.
Ukuphonononga ubudlelwane phakathi kokulungelelaniswa komphezulu we-lattice (ehambelanayo okanye ecacileyo) njengoko kubonakaliswe yimiqondiso yethu ye-texture kunye nomsebenzi we-MT, saqala ukubuyisela ulungelelwaniso phakathi kweeseli zentshukumo ehambelanayo (i-slope enhle) okanye intshukumo ebonakalayo (i-slope engathandekiyo) ngokuhlehla.inikwe ukuhlelwa kweeseli ngokwereyithi yempendulo yomqondiso xa kuthelekiswa nothelekiso (ngokwahlukileyo kwimowudi nganye icala).Imizekelo yale jiko lokulungisa i-lattice ukusuka kwiseli yomzekelo ofanayo kuMfanekiso 3 ibonisiwe kuMfanekiso 4a.Emva kokuhlelwa, sisebenzise uhlalutyo lokusebenza kommkeli (ROC) ukulinganisa uvakalelo lweseli nganye ekumodareyithweni kweempawu zokuthungwa (bona Iindlela).Imisebenzi ye-neurometric efunyenwe ngolu hlobo inokuthelekiswa ngokuthe ngqo kunye neempawu ze-psychophysical zeenkawu kwiseshoni enye ukwenzela ukuthelekisa ngokuthe ngqo uvakalelo lwe-psychophysical lwe-neurons kwi-lattice textures.Senze uhlalutyo lweempawu ezimbini kuzo zonke iiyunithi kwisampulu, sibala iimpawu ezihlukeneyo ze-neurometric kwisalathiso ngasinye sepateni (kwakhona, phezulu okanye phantsi).Kubalulekile ukuqaphela ukuba, kolu hlalutyo, sibandakanya kuphela iimvavanyo apho (i) i-stimuli iqulethe i-texture cue kwaye (ii) iinkawu ziphendule ngokuhambelana nelo nqaku (oko kukuthi, izilingo "ezichanekileyo").
Amazinga omlilo acetywayo ngokuchasene nokuchasana kweempawu ze-texture, ngokulandelanayo, kwi-gratings eguqukayo phezulu (ekhohlo) okanye ezantsi (ekunene), umgca oqinileyo ubonisa ukuhlenga komgca ofanelekileyo, kunye nedatha ephezulu (ezantsi) kumqolo ithathwa kwezo ziboniswe kuMfanekiso.Irayisi.3a Iseli, b (Umfanekiso 3c, d).Iimpawu ze-slope zokuhlehla zisetyenziselwe ukwabela iindlela zokuthungwa ezikhethiweyo (ezihambelanayo / ezicacileyo) kwiseli nganye / i-lattice orientation orientation indibaniselwano (n ≥ izilingo ze-20 kwimeko yokuvuselela).Iibar zemposiso zibonisa ukutenxa okusemgangathweni kwentsingiselo.Imisebenzi ye-neurometric yeeyunithi eziboniswe kwi-ba zichazwe kunye nemisebenzi ye-psychometric eqokelelwe ngexesha leseshoni enye.Ngoku, kuphawu ngalunye, siceba ingcebiso yesixhobo esikhethiweyo (ulungelelwaniso) (jonga okubhaliweyo) njengepesenti yomahluko wophawu (abscissa).Umahluko wobume utshintshiwe ukuze iincam zesixhobo ezikhethiweyo zibe zilungile kwaye iingcebiso ezingenanto zingalunganga.Idatha evela kwiigridi ezihamba phezulu (ezantsi) ziboniswa kwiipaneli zekhohlo (ekunene), kwimiqolo ephezulu (ephantsi) - idatha evela kwiiseli eziboniswe kwi-Fig. 3a, b (Fig. 3c, d).Umlinganiselo we-neurometric kunye ne-psychometric threshold (N / P) iboniswe kwiqela ngalinye.Izithethi.Izikhonkwane, imizuzwana.umzuzwana, ulawulo.isikhokelo, iphondo elikhethwayo, psi.Psychometry, Neurology.
I-Lattice tuning curves kunye nemisebenzi ye-neurometric yeeseli ezimbini ze-MT ezimele kunye nemisebenzi yazo ye-psychometric ehambelana nayo, edityaniswe kunye nezi mpendulo, ziboniswe kwiiphaneli eziphezulu kunye nezantsi zoMzobo 4a,b, ngokulandelanayo.Ezi seli zibonisa ukonyuka okurhabaxa okanye ukwehla njengoko uthsuphe wokuthungwa lusuka elubala luye ludibane.Ukongeza, isalathiso kunye namandla alo bhondi kuxhomekeke kwicala le-lattice motion.Ekugqibeleni, imisebenzi ye-neurometric ibalwe kwiimpendulo zala maseli asondele kuphela (kodwa ayikahambelani nayo) iimpawu ze-psychophysical ye-unidirectional grid movement.Zombini imisebenzi ye-neurometric kunye ne-psychometric yashwankathelwa kunye ne-thresholds, okt ihambelana malunga ne-84% yomahluko okhethwe ngokuchanekileyo (ohambelana nentsingiselo + 1 sd yomsebenzi ohlanganisiweyo we-Gaussian).Kuyo yonke isampuli, umlinganiselo we-N/P, umlinganiselo we-neurometric threshold ukuya kwi-psychometric one, i-avareji ye-12.4 ± 1.2 kwi-monkey N kunye ne-15.9 ± 1.8 kwi-monkey S, kunye ne-lattice ukuba ihambe ubuncinane kwicala elinye, kuphela Kwi- ~ 16% (18).%) iiyunithi ukusuka kwinkawu N (inkawu S) (Fig. 5a).Ukususela kumzekelo weseli oboniswe kumzobo.Njengoko kubonwe kwiMifanekiso 3 kunye ne-4, uvakalelo lwe-neurons lunokuchaphazeleka ngobudlelwane phakathi kokukhetha okukhethiweyo kweseli kunye nolwalathiso lwentshukumo ye-lattice esetyenziswe kwiimvavanyo.Ngokukodwa, i-orientation adjustment curves kwi-Fig. 3a, c ibonisa ubudlelwane phakathi kwe-neuron orientation setting ye-sinusoidal array enye kunye novakalelo lwayo kwintshukumo ecacileyo / ehambelanayo kwi-textured array. Oku bekunjalo kuzo zombini iinkawu (ANOVA; isalamane esikhethwayo sezalathiso sifakwe kwisisombululo se-10 °; inkawu N: F = 2.12, p <0.01; inkawu S: F = 2.01, p <0.01). Oku bekunjalo kuzo zombini iinkawu (ANOVA; isalamane esikhethwayo sezalathiso sifakwe kwisisombululo se-10 °; inkawu N: F = 2.12, p <0.01; inkawu S: F = 2.01, p <0.01). Это имело место для обеих обезьян (ANOVA; относительные предпочтительные направления объединены в группы с разрешениен, 10°, p 1; обезьяна S: F = 2,01, p <0,01). Oku bekunjalo kuzo zombini iinkawu (ANOVA; izalathiso ezikhethwayo zesalamane ezidityaniswe kwisisombululo se-10°; inkawu N: F=2.12, p<0.01; inkawu S: F=2.01, p<0.01) .两只猴子都是這种情况（方差分析；以10° 分辨率合并的相对首选方向；猴子N:F = 2.10=S = 2.10,p = 2.10,p = 10; ）.两 只 猴子 都 是 這 种 （方差 分析 以 10 ° 分辨率 合并 的 相对 方向 : : f = 10 p ; f = 2 p. <0.01.........） Это имело место для обеих обезьян (ANOVA; относительная предпочтительная ориентация объединена при разрешении 10°; обез:1,0 = 10°; S: F = 2,01, p <0,01). Oku bekunjalo kuzo zombini iinkawu (ANOVA; isalamane esikhethwayo sokuqhelaniswa nesombululo esiyi-10°; inkawu N: F=2.12, p<0.01; inkawu S: F=2.01, p<0.01).Ukunika iqondo elikhulu lokuguquguquka kubuntununtunu be-neuron (Fig. 5a), ukuze ube nombono wokuxhomekeka kwi-neuron ubuntununtunu kwiindlela ezikhethwayo ezihambelanayo, siqale saqhelanisa iseli nganye yokuqhelaniswa ekhethwayo ukuya kweyona “yeyona” yokuqhelanisa intshukumo yepateni yegridi (okt icala).apho i-grating yenza i-angle encinci phakathi kwe-cell orientation ekhethiweyo kunye nokuqhelaniswa kwepateni yokugaya).Sifumene ukuba imilinganiselo ehambelanayo ye-neurons (i-threshold ye-lattice "eyona" imbi kakhulu yokuqhelaniswa ne-lattice orientation "yeyona nto ingcono" ye-lattice orientation) yahluka ngolu lungelelwaniso luqhelekileyo olukhethiweyo, kunye neencopho kulo mlinganiselo we-threshold owenzeka malunga neepateni okanye ukuqhelaniswa kwecandelo (Umfanekiso 5b).)). Esi siphumo asikwazanga ukucaciswa ngocalucalulo ekusasazweni kwezikhokelo ezikhethiweyo kwiiyunithi kwisampulu nganye ukuya kwelinye lepatheni yeplaid okanye imiyalelo yecandelo (umzobo 5c; uvavanyo lukaRayleigh; inkawu N: z = 8.33, p <10-3, isetyhula intsingiselo = 190.13 deg ± 0.9 , 8 deg ± 0; 9. I-5) kwaye yayihambelana kwii-angles ze-plaid inter-grating (I-Fig. 2 eyongezelelweyo). Esi siphumo asikwazanga ukucaciswa ngocalucalulo ekusasazweni kwezikhokelo ezikhethiweyo kwiiyunithi kwisampulu nganye ukuya kwelinye lepatheni yeplaid okanye izikhombisi zecandelo (Umfanekiso 5c; uvavanyo lweRayleigh; inkawu N: z = 8.33, p <10-3, isetyhula intsingiselo = 190.13 demon 80 = ± 9; 45) kwaye yayingaguquguquki kwi-plaid inter-grating angles (iFig. 2 eyongezelelweyo). Этот эффект нельзя было объяснить смещением распределения предпочтительных направлений в единицах в каждой выборкей выборке й или направлений компонентов (рис. 5в; критерий Рэлея; обезьяна N: z = 8,33, p < 10–3). Esi siphumo asikwazanga ukuchazwa ngokutshintshwa kokusasazwa kweendlela ezikhethiweyo kwiiyunithi kwisampuli nganye ukuya kwelinye leendlela zokutshekisha okanye izikhokelo zecandelo (umzobo 5c; uvavanyo lweRayleigh; inkawu N: z = 8.33, p <10-3)., intsingiselo yesetyhula = 190.13 degrees ± 9.83 degrees;inkawu S: z = 0.79, p = 0.45) kwaye yayifana kuzo zonke iikona zeplaid grid (Figure Supplementary 2).7猴子N:z = 8.33，p <10-3 ，圆形平均值= 190.13 度上± 9.83 度；猴子S:z = 0.79，p = 0.45图2）.7图 图 瑞利 测试 ；猴子 n : z = 8.33 , p <10-3 , iintyatyambo ze- 平均值 圆形圆形 圆形 圆形 圆形z Этот эффект не может быть объяснен тем, что распределение предпочтительных ориентаций в клетках в каждом образце смеще ки, либо в сторону одной из ориентаций компонентов (рис. 5в; критерий Рэлея; обезьяна N: z = 8,33, p < 10–3). Esi siphumo asikwazi ukuchazwa yinto yokuba ukuhanjiswa kokukhetha okukhethiweyo kwiiseli kwisampuli nganye kuthambekele ekubhekiseleleni kwisakhiwo se-lattice okanye ngokubhekiselele kwi-orientation yecandelo (umzobo 5c; uvavanyo lukaRayleigh; inkawu N: z = 8.33, p <10-3)., umyinge wesetyhula) = 190.13 degrees ± 9.83 degrees;inkawu S: z = 0.79, p = 0.45) kwaye babelingana kwii-engile ze-lattice phakathi kweegridi (i-Supplementary Fig. 2).Ke, ubuntununtunu bee-neurons kwiigridi ezenziweyo buxhomekeke, ubuncinci ngokuyinxenye, kwiipropathi ezisisiseko ze-MT tuning.
Iphaneli ekhohlo ibonisa ukuhanjiswa kwe-N / P (i-neuron / psychophysiological threshold);iseli nganye ibonelela ngamanqaku amabini edatha, enye kwicala ngalinye apho ipateni ihamba khona.Iphaneli yasekunene icwangcisa imiqobo ye-psychophysical (ilungelelaniso) ngokuchasene ne-neuronal thresholds (abscissa) yazo zonke iiyunithi kwisampulu.Idatha kumqolo ophezulu (osezantsi) zivela kwinkawu N (S).b Umlinganiselo oqhelekileyo we-threshold ucwangciswe ngokuchasene nobukhulu bomahluko phakathi kweyona ndawo ilungileyo yokuqhelaniswa nelathisi kunye nokuqhelaniswa neseli ekhethwayo.Isalathiso "esigqwesileyo" sichazwa njengesalathiso sesakhiwo se-grating (kulinganiswa nge-sinusoidal grating eyodwa) esondeleyo kwindlela yeseli ekhethiweyo.Idatha yaqala ukubotshelelwa ngokuqhelaniswa okukhethiweyo okuqhelekileyo (imigqomo eyi-10 °), emva koko imilinganiselo ye-threshold yaye yenziwa yesiqhelo ukuya kwixabiso eliphezulu kwaye i-avareji ngaphakathi komgqomo ngamnye.Iiseli ezinolungelelwaniso olukhethwayo ukuba nkulu kancinane okanye zincinci kuneyokuqhelaniswa nenxalenye yelethisi inomahluko omkhulu kubuntununtunu kuqhelaniso lwepateni yelethiyisi.c Ihistogram epinki yosasazo olukhethwayo lokuqhelaniswa nazo zonke iiyunithi zeMT ezirekhodwe kwinkawu nganye.
Ekugqibeleni, impendulo ye-MT ihlaziywa yi-direction of grating movement kunye neenkcukacha zeempawu zethu zokwahlula (i-texture).Ukuthelekiswa kwe-neuronal kunye ne-psychophysical sensitivity kubonise ukuba, ngokubanzi, iiyunithi ze-MT zazingaphantsi kakhulu kwiimpawu ze-texture ezichaseneyo kuneenkawu.Nangona kunjalo, ubuntununtunu be-neuron butshintshile ngokuxhomekeke kumahluko phakathi kokukhetha okukhethiweyo kweyunithi kunye nesalathiso sokuhamba kwegridi.Ezona seli zinovakalelo zithande ukuba nokukhethwa kokuqhelaniswayo okuphantse kwagquma ipateni yelathisi okanye enye yeendlela zokumiselwa, kunye neseti encinci yeesampulu zethu bezinovakalelo okanye ezinovakalelo kunombono weenkawu umahluko umahluko.Ukuqonda ukuba ingaba imiqondiso evela kula mayunithi abuthathaka ngakumbi anxulunyaniswa ngokusondeleyo nombono kwiinkawu, sivavanye unxibelelwano phakathi kokuqonda kunye neempendulo ze-neuronal.
Isinyathelo esibalulekileyo ekusekeni uxhulumaniso phakathi komsebenzi we-neural kunye nokuziphatha kukuseka ulungelelwaniso phakathi kwe-neurons kunye neempendulo zokuziphatha kwi-stimuli eqhubekayo.Ukuze udibanise iimpendulo ze-neural kwizigwebo zokwahlulahlula, kubalulekile ukudala i-stimulus ethi, nangona ifana, ibonwa ngokuhlukileyo kwizilingo ezahlukeneyo.Kuphononongo lwangoku, oku kuboniswa ngokucacileyo ngunothi wokuchasa umahluko wegrayiti.Nangona sigxininisa ukuba, ngokusekelwe kwimisebenzi ye-psychometric yezilwanyana, iigratings ezinobuncinci (ngaphantsi kwe- ~ 20%) ukuchasana kombhalo ngokuqhelekileyo kubonwa njengento ehambelanayo okanye ecacileyo.
Ukulinganisa ubungakanani beempendulo zeMT ezihambelana neengxelo zokuqonda, senze uhlalutyo olunokwenzeka (CP) lwedatha yethu yegridi (jonga i-3).Ngamafutshane, i-CP i-non-parametric, imilinganiselo engekho-standard ebonisa ubudlelwane phakathi kweempendulo ze-spike kunye nezigwebo zokuqonda30.Ukukhawulela uhlalutyo kwiimvavanyo kusetyenziswa iigridi kunye ne-zero yokuchasana kombhalo kunye neeseshoni apho iinkawu zenze ukhetho oluhlanu ubuncinane kuhlobo ngalunye lwezilingo, sibale i-SR ngokwahlukileyo kwisalathiso ngasinye sokuhamba kwegridi. Kuzo zonke iinkawu, siye sabona ixabiso eliqhelekileyo le-CP elikhulu kakhulu kunokuba sinokulindela ngethuba (umzobo 6a, d; i-monkey N: ithetha i-CP: 0.54, 95% CI: (0.53, 0.56), ukuvavanya kwe-t-macala amabini ngokuchasene ne-null ye-CP = 0.5, t = 6.5 monkey: 9; I-5% CI: (0.54, 0.57), i-t-side t-test, t = 9.4, p <10-13). Kuzo zonke iinkawu, siye sabona ixabiso eliqhelekileyo le-CP elikhulu kakhulu kunokuba sinokulindela ngethuba (umzobo 6a, d; i-monkey N: ithetha i-CP: 0.54, 95% CI: (0.53, 0.56), ukuvavanya kwe-t-macala amabini ngokuchasene ne-null ye-CP = 0.5, t = 6.5 monkey: 9; I-5% CI: (0.54, 0.57), i-t-side t-test, t = 9.4, p <10-13).Kwiimfene, siye sabona i-CP ephakathi kakhulu kunokuba ilindeleke ngokungalindelekanga (Umfanekiso 6a, d; inkawu N: ithetha i-CP: 0.54, 95% CI: (0.53, 0.56), i-t-tailed t-test vs. null values).CP = 0,5, t = 6,7, p <10-9; CP = 0.5, t = 6.7, p <10-9; обезьяна S: среднее CP: 0,55, 95% ДИ: (0,54, 0,57), двусторонний t-критерий, t = 9,4, p < 10–13) . inkawu S: ithetha i-CP: 0.55, 95% CI: (0.54, 0.57), uvavanyo lwe-t-tailed ezimbini, t = 9.4, p <10-13).在猴子中，我們观察到平均CP 值显着大于我們偶然预期的值（图6a，d；猴子N:平均CP:0.54,95% CI:3,000.双边t 检验CP = 0.5, t = 6.7, p <10−9; 猴子S: 平均CP: 0.55, 95% CI: (0.54, 0.57), 双边t <10−9; 检验 −1 = 10, 4 检验在 猴子 中 , 我們 观察 平均 平均 值 显着 大于 我們 偶然 的值.56） , 空值 检验 CP = 0.5, t = 6.7, p <10−9;猴子S: 平均CP: 0.55, 95% CI: (0.54, 0.57), 双边t检验, t=9.4, p <10−13) У обезьян мы наблюдали средние значения CP, значительно превышающие то, что мы могли бы ожидать случайно (рис. 6a, CP: 9, 5:5; ДИ: (0,53, 0,56), двусторонний t- тест CP против нуля = 0,5, t = 6,7, p < 10-9, обезьяна S: средний CP: 0,55, 90, 5,55%, 55% критерий, t = 9,4, p < 10- 13) . Kwiimfene, siqaphele amaxabiso e-CP athetha kakuhle ngaphezu koko sinokukulindela ngethuba (umzobo 6a, d; inkawu N: ithetha i-CP: 0.54, 95% CI: (0.53, 0.56), uvavanyo lwe-t-tailed t-t-2 vs. S. I-CI: (0.54, 0.57), i-t-tailed t-criterion, t = 9.4, p <10-13).Ke, ii-neuron zeMT zikholisa ukuvutha ngamandla ngakumbi naxa kungekho nakuphi na ukwahlulahlula okuthe kratya kweempawu, xa isilwanyana esibona ngaso intshukumo yelathisi ihambelana nezinto ezikhethwa yiseli.
a Ukhetho lokusasazwa kwamathuba kwiigridi ngaphandle kwemiqondiso yokuthungwa kweesampulu ezirekhodiweyo kwi-monkey N. Iseli nganye inokufaka isandla ukuya kumanqaku amabini edatha (enye kwicala ngalinye lentshukumo yegridi).Ixabiso le-CP eliqhelekileyo ngaphezu kwe-random (iintolo ezimhlophe) libonisa ukuba ngokubanzi kukho ubudlelwane obubalulekileyo phakathi komsebenzi we-MT kunye nombono.b Ukuphonononga impembelelo yalo naluphi na ukhetho olunokuthi lubekho, sibale i-CP ngokwahlukileyo kuyo nayiphi na i-stimulus apho iinkawu zenze impazamo enye ubuncinane.Amathuba okukhetha acwangciswe njengomsebenzi wokhetho lomlinganiselo (pref/null) kuzo zonke izivuseleli (ekhohlo) kunye nexabiso elipheleleyo lokumahluko kwamanqaku okuthungwa (ekunene, idatha esuka kwiiseli ezili-120).Umgca oqinileyo kunye nendawo enomthunzi kwindawo yolawulo ekhohlo imele intsingiselo ± isiqingatha somndilili ohambayo wamanqaku angama-20.Ukhetho olunokwenzeka olubalwe kwi-stimuli kunye nomlinganiselo wokhetho olungalungelelananga, olufana neegridi ezinophawu oluphezulu olwahlukileyo, zahluke kakhulu kwaye zadityaniswa malunga nokunokwenzeka.Indawo enomthunzi ongwevu kwiphaneli elungileyo igxininisa ukuchasana kweempawu ezibandakanyiweyo ekubalweni kwamathuba aphezulu okukhetha.c Amathuba okhetho olukhulu (olungelelanisiweyo) lucetywe ngokuchasene nomda we-neuron (abscissa).Amathuba okhetho aye anxulunyaniswa kakubi kakhulu nomda.I-convention df iyafana ne-ac kodwa isebenza kwi-157 data eyodwa evela kwi-monkey S ngaphandle kokuba kuchazwe ngenye indlela.g Eyona ndlela iphezulu yokhetho (ulungelelwaniso) icwangciswe ngokuchasene necala elithandwayo eliqhelekileyo (abscissa) kwinkawu nganye kwezi zimbini.Iseli nganye yeMT yanikela ngamanqaku amabini edatha (enye kwicala ngalinye lesakhiwo se-lattice).h Iziza zebhokisi ezinkulu zokhetho olunokwenzeka kwi-engile nganye ye-inter-raster.Umgca oqinileyo uphawula i-median, imiphetho engezantsi kunye nephezulu yebhokisi imele i-25th kunye ne-75th percentiles ngokulandelanayo, i-whiskers yandiswe ukuya kuma-1.5 amaxesha oluhlu lwe-interquartile, kunye nabangaphandle ngaphaya kwalo mda bayaphawulwa.Idatha kwiiphaneli zasekhohlo (ekunene) zisuka kwi-120 (157) yeeseli zeenkawu ze-N(S) zomntu ngamnye.i Amathuba aphezulu okukhetha (ulungelelwaniso) acwangciswe ngokuchasene nexesha lokuqala kwe-stimulus (abscissa).I-CP enkulu yabalwa kwii-rectangles ezityibilikayo (ububanzi be-100 ms, inyathelo le-10 ms) kulo lonke uvavanyo kwaye emva koko i-avareji kwiiyunithi.
Olunye uphando lwangaphambili luye lwaxela ukuba i-CP ixhomekeke kwinani elihambelanayo lezilingo kwi-basal rate distribution, oku kuthetha ukuba lo mlinganiso awuthembekanga kangako kwi-stimuli ebangela ukubahluko omkhulu kumlinganiselo wokhetho ngalunye.Ukuvavanya le mpembelelo kwidatha yethu, sibale i-CP ngokwahlukileyo kuzo zonke izivuseleli, kungakhathaliseki ukuba yintoni umehluko wokuthungwa kweempawu, kwaye iinkawu zenze ubuncinane uvavanyo olulodwa lobuxoki.I-CP icwangciswe malunga nomlinganiselo wokukhetha (pref / null) kwisilwanyana ngasinye kwi-Figure 6b kunye ne-e (iphaneli yasekhohlo), ngokulandelanayo.Ukujonga i-avareji ezihambayo, kucacile ukuba i-CP ihlala ingaphezulu kwe-probability phezu koluhlu olubanzi lwee-odds zokukhetha, iyancipha kuphela xa amathuba okuwa (ukwanda) ngaphantsi (ngaphezulu) 0.2 (0.8).Ngokusekelwe kwiimpawu ze-psychometric zezilwanyana, sinokulindela ukuba i-coefficients yokukhetha yolu bukhulu isebenze kuphela kwi-stimuli kunye ne-high-contrast texture cues (ehambelanayo okanye ecacileyo) (jonga imizekelo yeempawu zengqondo kwi-Fig. 2a, b).Ukufumanisa ukuba ngaba le nto yayinjalo kwaye nokuba i-PC ebalulekileyo iqhubekile nokuba i-stimuli eneempawu ezicacileyo zokwahlula, siye savavanya umphumo wexabiso elipheleleyo lombhalo we-PC (Fig. 6b, e-right).Njengoko bekulindelekile, i-CP yayiphezulu kakhulu kunokuba kunokwenzeka ukuba i-stimuli equle ukuya kwimodareyitha (~ 20% umahluko okanye ngaphantsi) i-segmentation cues.
Kwi-orientation, isantya, kunye nemisebenzi engafanelekanga yokuqaphela, i-MT CP ithande ukuba phezulu kwii-neurons ezithintekayo, mhlawumbi ngenxa yokuba ezi neurons zithwala izibonakaliso ezinolwazi30,32,33,34.Ngokuhambelana nezi ziphumo siye sabona ulungelelwaniso oluthobekileyo kodwa olubalulekileyo phakathi kwe-CP enkulu, ebalwa ukusuka kumazinga okudubula afunyenwe ngu-z kuwo wonke umahluko wecue wokuthungwa oboniswe kwiphaneli yenene yeFig.I-6b, i-e, kunye ne-neuronal threshold (Umfanekiso 6c, f; i-geometric ithetha ukuhla; inkawu N: r = -0.12, p = 0.07 monkey S: r = -0.18, p <10-3). I-6b, i-e, kunye ne-neuronal threshold (Umfanekiso 6c, f; i-geometric ithetha ukuhla; inkawu N: r = -0.12, p = 0.07 monkey S: r = -0.18, p <10-3).Ngokuhambelana nezi ziphumo, siye sabona ukulungelelaniswa okuthobekileyo kodwa okubalulekileyo phakathi kwe-CP enkulu ebalwa ukusuka kwi-frequency excitation z-score ukusuka kwi-textura yesignali ye-texture egxininiswe kwiphaneli echanekileyo ye-Fig. 6b, e, kunye ne-neuronal threshold (Fig. 6c, f; geometric).ijometri ithetha ukuhlehla;обезьяна N: r = -0,12, p = 0,07 обезьяна S: r = -0,18, p < 10-3). inkawu N: r = -0.12, p = 0,07 inkawu S: r = -0.18, p <10-3).与這些发现一致，我們观對到大CP 之间存在适度但显着的相关性，這是根据图6b, e 和神经元阈候，和神经元阈倚回归；猴子N:r = -0.12，p = 0.07 iifom zeS:r = -0.18,p <10-3）。於 這些 发现 一致 ， 我們到 大 大 之间 存在 适度 但 显着 的 相关性這 是根据 图 6b , e . . . , f ; 回归 ； 猴子 n : r = -0.12 , p = 0.07 猴子S:r = -0.18,p <10-3）.Ngokuhambelana nezi ziphumo, siye sabona ukulungelelaniswa okuthobekileyo kodwa okubalulekileyo phakathi kwee-CV ezinkulu njengoko kuboniswe kwi-Fig.Обезьяна S: г = -0,18, р < 10-3). I-Monkey S: r = -0.18, p <10-3).Ngoko ke, iimpawu ezivela kwiiyunithi ezifundisayo zivame ukubonisa i-covariance enkulu kunye nezigwebo ze-subjective segmentation kwiinkawu, okubalulekileyo kungakhathaliseki ukuba nayiphi na isicatshulwa sombhalo esongeziweyo kwi-bias yokuqonda.
Ngenxa yokuba ngaphambili sasisungule ubudlelwane phakathi kovakalelo kwiimpawu zegrid texture kunye nokukhethwa kwe-neuronal orientation, sasizibuza ukuba kukho ubudlelwane obufanayo phakathi kwe-CP kunye nokuqhelaniswa okukhethiweyo (Umfanekiso we-6g).Olu nxulumano lwalubaluleke kancinane kuphela kwinkawu S (ANOVA; inkawu N: 1.03, p=0.46; inkawu S: F=1.73, p=0.04).Asibonanga mahluko kwi-CP yee-angles ze-lattice phakathi kwee-lattices nakwesiphi na isilwanyana (umzobo 6h; ANOVA; inkawu N: F = 1.8, p = 0.11; inkawu S: F = 0.32, p = 0. 9).
Ekugqibeleni, umsebenzi wangaphambili ubonise ukuba i-CP iyatshintsha kulo lonke uvavanyo.Olunye uphando luye lwabika ukwanda okubukhali okulandelwa yimpembelelo yokhetho egudileyo, i-30 ngelixa abanye baye babika ukunyuka okuzinzileyo kwisignali yokukhetha ngexesha lovavanyo31.Kwinkawu nganye, sibale i-CP yeyunithi nganye kwiimvavanyo kunye ne-zero textural difference (ngokulandelelanayo, ngokwepateni yokuqhelaniswa) kwiiseli ze-100 ms zinyathela zonke i-20 ms ukusuka kwi-pre-stimulus ukuqala ukuposa kuthetha ukutshatyalaliswa kwangaphambili kwe-stimulus.I-avareji ye-CP dynamics yeenkawu ezimbini iboniswe kwi-Fig. 6i.Kuzo zombini iimeko, i-CP yahlala kwinqanaba elingaqhelekanga okanye isondele kakhulu kude kube malunga ne-500 ms emva kokuqala kwe-stimulus, emva koko i-CP yanda ngokukhawuleza.
Ukongeza ekutshintsheni ubuntununtunu, i-CP ibonakaliswe ukuba ichatshazelwa ziimpawu ezithile zeempawu zokulungisa iiseli.Umzekelo, u-Uka kunye ne-DeAngelis34 bafumanisa ukuba i-CP kumsebenzi we-binocular mismatch recognition ixhomekeke kwi-symmetry ye-binocular mismatch tuning curve yesixhobo.Kule meko, umbuzo onxulumeneyo ngowokuba ingaba iiseli ze-Pattern direction selective (PDS) zinovakalelo ngakumbi kuneeseli zeCDS.Iiseli ze-PDS zifakela i-orientation jikelele yeepatheni eziqulethe iindawo ezininzi zokuqhelaniswa kwendawo, ngelixa iiseli ze-CDS ziphendula ukunyakaza kwamacandelo epateni yolwalathiso (Umfanekiso 7a).
Umelo olucwangcisiweyo lwemowudi yolungelelwaniso lwecandelo kunye ne-hypothetical grating (ekhohlo) kunye ne-grating orientation tuning curves (ekunene) (jonga iiMathiriyeli kunye neeNdlela).Ngamafutshane, ukuba iseli idibanisa kuwo wonke amacandelo egridi umqondiso wokuhamba kwepateni, umntu unokulindela i-curves efanayo yokulungisa igridi kunye negridi ye-stimuli (ikholamu yokugqibela, igophe eliqinileyo).Ngokuchaseneyo, ukuba iseli ayizihlanganisi izikhokelo zamacandelo kwintshukumo yepateni yomqondiso, umntu unokulindela ijiko lokulungisa kabini elinencopho kwicala ngalinye lentshukumo yegrayiti eguqulela icandelo elinye kwicala elikhethiweyo leseli (umhlathi wokugqibela, igophe elidayiweyo)..b (ngasekhohlo) iijika zokulungelelanisa ukuqhelaniswa kwe-sinusoidal array kwiiseli eziboniswe kwiMifanekiso 1 kunye ne-2. 3 kunye ne-4 (umqolo ophezulu - iiseli kwiMifanekiso 3a, b kunye ne-4a, b (phezulu); iphaneli ephantsi - iiseli zeFig. 3c, d kunye ne-4a, b (ezantsi)).(Embindini) Ipateni kunye noqikelelo lwecandelo olubalwe kwiiprofayili zokulungisa iiletiyisi.(Ekunene) Ukulungelelanisa igridi yezi seli.Iiseli zephaneli ephezulu (esezantsi) zihlelwa njenge template (component) cells.Qaphela ukuba akukho mbalelwano phakathi kokuhlelwa kwamacandelo epateni kunye nokukhethwa kwentshukumo yeseli ehambelanayo / ecacileyo (jonga iimpendulo ze-texture lattice kwezi seli kwi-Fig. 4a).c I-Coefficient ye-partial corelation ye-z-score mode (ulungelelwaniso) ecetywe ngokuchasene ne-partial coefficient coefficient ye-z-score component (abscissa) yazo zonke iiseli ezirekhodwe kwi-N (ekhohlo) kunye ne-S (ekunene) iinkawu.Imigca eshinyeneyo ibonisa indlela yokubaluleka esetyenziselwa ukwahlula iiseli.d Iploti yokhetho oluphezulu olunokwenzeka (ulungelelwaniso) ngokuchasene nesalathiso semowudi (Zp – Zc) (abscissa).Idatha kwiiphaneli zasekhohlo (ekunene) zibhekisa kwinkawu N(S).Izangqa ezimnyama zibonisa idatha kwiiyunithi eziqikelelweyo.Kuzo zombini izilwanyana, bekukho ulungelelwaniso olubalulekileyo phakathi kokukhethwa okuphezulu okunokwenzeka kunye nesalathiso sepateni, ecebisa ulungelelwaniso olukhulu lweeseli ezinepateni yomqondiso wokuqhelaniswa ne-stimuli kunye nokuqhelaniswa kwamacandelo amaninzi.
Ngoko ke, kwisethi yovavanyo eyahlukileyo, silinganise iimpendulo kwiigridi ze-sinusoidal kunye neegridi zokwahlula i-neurons kwiisampuli zethu njenge-PDS okanye i-CDS (jonga Iindlela).I-Lattice tuning curves, i-template component predictions eyakhelwe kule datha yokulungisa, kunye ne-lattice tuning curves kwiiseli eziboniswe kwiMizobo 1 kunye ne-3. I-Figure 3 kunye ne-4 kunye ne-Supplementary Figure 3 iboniswe kuMfanekiso 7b.Ukuhanjiswa kwepateni kunye nokukhethwa kwecandelo, kunye nokuqhelaniswa kweseli ekhethiweyo kudidi ngalunye, kuboniswa kwinkawu nganye kwi-Fig. 7c kunye nefig eyongezelelweyo.4 ngokulandelelanayo.
Ukuvavanya ukuxhomekeka kwe-CP ekulungisweni kwamacandelo epatheni, saqala ukubala isalathiso sepateni 35 (PI), amaxabiso amakhulu (amancinci) abonisa i-PDS enkulu (CDS) ukuziphatha okufanayo.Ukunikezelwa kolu mboniso ungasentla: (i) uvakalelo lwe-neuronal luyahluka kunye nomahluko phakathi kwe-cell orientation ekhethiweyo kunye nesalathiso sokunyakaza kwe-stimulus, kwaye (ii) kukho ukulungelelaniswa okubalulekileyo phakathi kovakalelo lwe-neuronal kunye nokwenzeka kokukhetha kwisampuli yethu, sifumene ubudlelwane phakathi kwe-PI kunye ne-CP iyonke ifundwe ngeyona ndlela "engcono" yolwalathiso lokuhamba kweseli nganye (jonga ngasentla). Sifumene ukuba i-CP ihambelana kakhulu ne-PI (Umfanekiso we-7d; i-geometric ithetha ukuhla; i-CP monkey enkulu N: r = 0.23, p <0.01; bi-stable CP monkey N r = 0.21, p = 0.013; grand CP monkey S: r = ps 3; 0.29, p <10−3), ebonisa ukuba iiseli ezihlelwe njenge-PDS zibonise umsebenzi omkhulu onxulumene nokukhetha kune-CDS kunye neeseli ezingabhalwanga. Sifumene ukuba i-CP ihambelana kakhulu ne-PI (Umfanekiso we-7d; i-geometric ithetha ukuhla; i-CP monkey enkulu N: r = 0.23, p <0.01; bi-stable CP monkey N r = 0.21, p = 0.013; grand CP monkey S: r = ps 3; 0.29, p <10−3), ebonisa ukuba iiseli ezihlelwe njenge-PDS zibonise umsebenzi omkhulu onxulumene nokukhetha kune-CDS kunye neeseli ezingabhalwanga. Мы обнаружили, что CP значительно коррелирует с PI (рис. 7d; регрессия среднего геометрического; большая обезьяна CP N: 3, r0; 0,25 зьяна CP N r = 0,21, p = 0,013; большая обезьяна CP S: r = 0,30, p < 10-4; бистабильный CP <обезьяны S: r = 0,29, ут клетки, классифицированные как PDS, проявляли большую активность, связанную с выбором, чем CDS и неклассифицированные клетки. Sifumene ukuba i-CP ihambelana kakhulu ne-PI (Umfanekiso we-7d; i-geometric ithetha ukuhla; inkawu enkulu CP N: r = 0.23, p <0.01; i-bistable monkey CP N r = 0.21, p = 0.013; inkawu enkulu CP S: r = CP monkey 0: 20; 9, p <10-3), ebonisa ukuba iiseli ezichazwe njengePDS zibonise umsebenzi omningi, ohambelana nokukhetha kuneCDS kunye neeseli ezingabhalwanga.我們发现CP 与PI 显着相关（图7d；几何平均回归；大CP 猴N:r = 0.23,p <0.01；双稳态CP = 猴0: CP = 30.02 CP = 猴00: CP = 10.02r. r = 0.30, p < 10-4；双稳态CP 猴S:r = 0.29,p < 10-3），表明分类為PDS 的细胞比CDS 和未分类的细诚诚该大衛的细胞比CDS活性. CP 与PI 显着相关（图7d；几何平均回归；大CP猴N:r = 0.23，p <0.01；双稳态CP 猴N r = 0.20,r = 0.20; Мы обнаружили, что CP был значительно связан с PI (рис. 7d; регрессия среднего геометрического; большая обезьяна CP N:3, r0, p = 0,25 зьяна CP N r = 0,21, p = 0,013; большая обезьяна CP S: r = 0,013) 0,30, p <10-4; Sifumene ukuba i-CP yayinxulumene kakhulu ne-PI (Umfanekiso 7d; i-geometric mean regression; inkawu enkulu CP N: r = 0.23, p <0.01; i-bistable monkey CP N r = 0.21, p = 0.013; inkawu enkulu CP S: r = 0.010-4; бистабильный CP обезьяны S: r = 0,29, p < 10-3), что указывает на то, что клетки, классифицированные как PDS, прояотинцикс ть, чем клетки, классифицированные как CDS и неклассифицированные. monkey S bistable CP: r = 0.29, p <10-3), ebonisa ukuba iiseli ezihlelwa njenge-PDS zibonise umsebenzi omkhulu wokukhetha kuneeseli ezichazwe njenge-CDS kwaye zingachazwanga.Ngenxa yokuba zombini i-PI kunye ne-neuron sensitivity ehambelana ne-CP, senze uhlalutyo oluninzi lwe-regression (kunye ne-PI kunye ne-neuron sensitivity njengezinto ezizimeleyo kunye ne-CP enkulu njenge-variable variable) ukulawula ulungelelwaniso phakathi kwamanyathelo amabini akhokelela ekubeni kwenzeke umphumo.. Zombini i-coefficients yokulungelelaniswa kwenxalenye yayibalulekile (inkawu N: i-threshold vs. CP: r = -0.13, p = 0.04, PI vs. CP: r = 0.23, p <0.01; inkawu S: i-threshold vs. CP: r = -0.0.10, CP = 0.10, p. −3), icebisa ukuba i-CP inyuke ngovakalelo kwaye ngendlela ezimeleyo inyuka nge-PI. Zombini i-coefficients yokulungelelaniswa kwenxalenye yayibalulekile (inkawu N: i-threshold vs. CP: r = -0.13, p = 0.04, PI vs. CP: r = 0.23, p <0.01; inkawu S: i-threshold vs. CP: r = -0.0.10, CP = 0.10, p. −3), icebisa ukuba i-CP inyuke ngovakalelo kwaye ngendlela ezimeleyo inyuka nge-PI. Inkcazelo ngokuthe gabalala были значимыми (обезьяна N: порог против CP: r = -0,13, p = 0,04, PI против CP: 3, p = 0, p = 0, прок; ротив CP: r = -0,16, p = 0,03, PI vs CP: 0,29, p < 10-3), предполагая, что CP увеличивается с чувствительностью и независимзомым обетым обегий против обезе. Zombini i-coefficients yokulungelelaniswa kwenxalenye yayibalulekile (inkawu N: i-threshold vs. CP: r=-0.13, p=0.04, PI vs. CP: r=0.23, p <0.01; inkawu S: i-threshold vs. CP: r = -0.16, p = 0.03 , CP-3, i-CP = 0.03, iphakamisa ngovakalelo kunye nokwandisa ngokuzimeleyo ngePI.两个偏相关系数均显着（猴子N:阈值与CP:r = -0.13,p = 0.04,PI 与CP:r = 0.23，p <0.01 = 3: 阈 CP10 = S = 3.23 ,PI vs CP:0.29,p <10-3），表明CP 随灵敏度增加而增加，并且以独立方式随PI 增加。两个偏相关系数均显着（猴子N:阈值与CP:r = -0.13,p = 0.04，PI = 0.03，PI vs CP:0.29，p <10-3明， Inkcazelo ye-ораа коэффчьиецннннннннццнн I-R = поти против cp: R. -0,16, P <10-3), что пто пчозвазвает На пто, ччо CP ууелителе и увеличась с с с с независим обзомомо.comм. Zombini i-coefficients yokulungelelaniswa kwenxalenye yayibalulekile (inkawu N: i-threshold vs. CP: r=-0.13, p=0.04, PI vs. CP: r=0.23, p <0.01; inkawu S: i-threshold vs. CP: r = -0.16, p = 0.03 CPdic: CP-3, p = 0.03, CPdic 0.03, p. ukwanda ngovakalelo kunye nokunyuka nge-PI ngendlela ezimeleyo.
Sirekhode umsebenzi omnye kwindawo ye-MT, kwaye iinkawu zabika imbono yazo yeepateni ezinokubonakala zihamba ngokuhambelanayo okanye ezicacileyo.Uvakalelo lwe-neurons kwi-segmentation cues ezongeziweyo kwiimbono ezicalu-caluleyo ziyahluka ngokubanzi kwaye zimiselwa, ubuncinci ngokuyinxenye, ngobudlelwane phakathi kweyunithi yoqhelaniso olukhethwayo kunye nesalathiso sentshukumo yovuselelo.Kubo bonke abantu, uvakalelo lwe-neuronal lwalusezantsi kakhulu kunovakalelo lwe-psychophysical, nangona ezona yunithi zibuthathaka zihambelana okanye zigqithise ubuntununtunu bokuziphatha kwimiqondiso yokwahlulahlula.Ukongezelela, kukho i-covariance ebalulekileyo phakathi kokuphindaphindiweyo kokudubula kunye nokuqonda, ebonisa ukuba ukubonakaliswa kweMT kudlala indima ekuhlukaniseni.Iiseli ezikhethayo ziluphucule uvakalelo lwazo kumahluko kwimiqondiso yolwahlulo lweletiyisi kwaye zithande ukubonakalisa intshukumo yehlabathi kwizivuseleli ezinemikhomba-ndlela emininzi yendawo, ebonisa olona lungelelwaniso lwengqiqo luphezulu.Apha siqwalasela ezinye iingxaki ezinokubakho phambi kokuthelekisa ezi ziphumo nomsebenzi wangaphambili.
Ingxaki enkulu ngophando olusebenzisa i-bistable stimuli kwiimodeli zezilwanyana kukuba iimpendulo zokuziphatha azinakusekelwe kumlinganiselo womdla.Umzekelo, iinkawu zethu zinokuxela imbono yazo yendlela yokuthungwa ngokuzimeleyo kwimbono yazo yokudibana kwelathisi.Imiba emibini yedatha ibonisa ukuba oku akunjalo.Okokuqala, ngokuhambelana neengxelo zangaphambili, ukutshintsha i-engile yokuqhelaniswa nesalamana yokwahlula kuluhlu lwamacandelo ngokucwangcisiweyo kutshintshe ukubakho kokuqonda okuhambelanayo.Okwesibini, ngokomyinge, umphumo uyafana kwiipateni eziqulethe okanye ezingenazo iimpawu zokuthungwa.Xa zidibene, ezi ngqwalasela zibonisa ukuba iimpendulo zeenkawu zihlala zibonisa imbono yazo yoqhagamshelo / elubala.
Enye ingxaki enokubakho kukuba asikaziphuculanga iiparamitha zentshukumo yegrayiti yemeko ethile.Kwimisebenzi emininzi yangaphambili ethelekisa i-neuronal kunye ne-psychophysical sensitivity, i-stimuli ikhethwe ngabanye kwiyunithi nganye ebhalisiweyo [31, 32, 34, 36, 37, 38, 39, 40, 41, 42, 43, 44, 45].Apha sisebenzise izikhokelo ezimbini ezifanayo zokuhamba kwepateni yelathisi kungakhathaliseki ukulungelelaniswa kokuqhelaniswa neseli nganye.Olu luyilo luye lwasivumela ukuba sifunde indlela ubuntununtunu obutshintshile ngayo ngokugqithelana phakathi kokunyakaza kwe-lattice kunye ne-orientation ekhethiweyo, nangona kunjalo, ayizange ibonelele ngesiseko sokuqala sokugqiba ukuba iiseli zikhetha iileyile ezihambelanayo okanye ezicacileyo.Ke ngoko, sixhomekeke kwiikhrayitheriya zobuchule, sisebenzisa impendulo yeseli nganye kumnatha owenziweyo, ukwabela okuthandwayo kunye neelebhile zero kudidi ngalunye lwentshukumo yemesh.Nangona kungenakwenzeka, oku kunokugqwesa ngokucwangcisiweyo iziphumo zohlalutyo lwethu lobuntununtunu kunye nokufunyanwa komqondiso we-CP, okunokuthi kuqikelelwe ngokugqithisileyo nawuphi na umlinganiselo.Nangona kunjalo, imiba emininzi yohlalutyo kunye nedatha exutyushwa ngezantsi ibonisa ukuba oku akunjalo.
Okokuqala, ukwabela amagama akhethiweyo (okungekho nto) kwi-stimuli eyenza umsebenzi omningi (ongaphantsi) awuchaphazelekanga ukwahlulahlula kwezi mpendulo.Kunoko, iqinisekisa kuphela ukuba imisebenzi ye-neurometric kunye ne-psychometric inophawu olufanayo, ngoko inokuthelekiswa ngokuthe ngqo.Okwesibini, iimpendulo ezisetyenziselwa ukubala i-CP (izilingo "ezingalunganga" ze-textured gratings kunye nazo zonke izilingo ze-gratings ngaphandle kokungafani kokuthungwa) azizange zibandakanywe kuhlalutyo lokubuyisela olunqume ukuba iseli nganye "ikhetha" edibeneyo okanye imidlalo ebonakalayo.Oku kuqinisekisa ukuba iziphumo zokhetho azikhethi cala kubizo olukhethiweyo/olungasebenziyo, okukhokelela ekukhetheni okubonakalayo okunokwenzeka.
Izifundo zeNewsom kunye noogxa bakhe [36, 39, 46, 47] babe ngabokuqala ukumisela indima ye-MT ekuqikelweni okuqikelelwayo kolwalathiso lwentshukumo.Iingxelo ezilandelayo ziqokelele idatha malunga nokuthatha inxaxheba kwe-MT kwi-deep34,44,48,49,50,51 kunye ne-speed32,52, i-fine orientation33 kunye nombono wesakhiwo se-3D ukusuka kwintshukumo31,53,54 (3D amahlathi azinzileyo).ukulawula).Sandisa ezi ziphumo ngeendlela ezimbini ezibalulekileyo.Okokuqala, sinika ubungqina bokuba iimpendulo zeMT zifaka isandla kwisahlulo sokuqonda kweempawu ze-visuomotor.Okwesibini, siqaphele ubudlelwane phakathi kwe-MT yokukhetha imo yokukhetha kunye nesi siginali yokukhetha.
Ngokwengqiqo, iziphumo zangoku zifana kakhulu nomsebenzi we-3-D SFM, njengoko zombini ziyingcamango enzima ye-bistable ebandakanya ukunyakaza kunye nokucwangcisa ubunzulu.U-Dodd et al.31 ufumene ithuba elikhulu lokukhetha (0.67) kumsebenzi wenkawu echaza ukujikeleza okujikelezayo kwe-bistable 3D SFM cylinder.Sifumene isiphumo esincinci sokukhetha se-bistable grid stimuli (malunga ne-0.55 kuzo zombini iinkawu).Ekubeni uvavanyo lwe-CP luxhomekeke kwi-coefficient yokukhetha, kunzima ukutolika i-CP efunyenwe phantsi kweemeko ezahlukeneyo kwimisebenzi eyahlukeneyo.Nangona kunjalo, ubukhulu besiphumo sokukhetha esisibonileyo sasifana ne-zero kunye ne-texture ephantsi yokuchasa i-gratings, kwaye naxa sidibanisa i-low / no-text-maxtivity stimuli ukwandisa amandla.Ngoko ke, lo mahluko kwi-CP akunakwenzeka ukuba ube ngenxa yokungafani kumazinga okhetho phakathi kwedatha.
Utshintsho oluncinci kwireyithi yokudubula yeMT ehamba kunye nembono kule meko yokugqibela ibonakala idida xa kuthelekiswa neemeko ezibukhali nezingokomgangatho ezahlukeneyo zokucinga ezibangelwa yi-3-D SFM yokuvuselela kunye nolwakhiwo lwegridi ye-bistable.Enye into enokwenzeka kukuba sijonge phantsi isiphumo sokukhetha ngokubala izinga lokudubula kulo lonke ixesha lokuvuselela.Ngokwahlukileyo kwimeko ye-31 3-D SFM, apho ulwahlulo kumsebenzi we-MT luphuhliswe malunga ne-250 ms kwizilingo kwaye lwanda ngokukhawuleza kulo lonke izilingo, uhlalutyo lwethu lwamandla exeshana leempawu zokukhetha (jonga i-500 ms emva kokuvuselela ukuqala kuzo zombini iinkawu. Ukongeza, emva kokunyuka okubukhali ngeli xesha lengxelo, sibona i-Rubin ye-fluc ngexesha lovavanyo lwe-Hubin5 ngexesha le-Hubin5. Umbono we-bistable rectangular arrays kaninzi utshintsha ngexesha lolingo olude.Nangona izivuseleli zethu zanikwa imizuzwana eyi-1.5 kuphela, imbono yeenkawu zethu zinokuhluka ukusuka ekudibaneni ukuya elubala ngexesha lovavanyo (iimpendulo zabo zibonise umbono wabo wokugqibela ekukhetheni i-cue.) Ngoko ke, uguqulelo lwexesha lokuphendula lomsebenzi wethu, okanye isicwangciso apho iinkawu zinokuthi zibe nombono olindelekileyo ukuba isiphumo sabo siqhubeke. funda ngokwahlukileyo kule misebenzi mibini.Nangona kudala kucingwa ukuba imiqondiso ye-CPU iphuma kwi-sensory decoding kunye nengxolo ehambelanayo, i-56 Gu kunye noogxa57 bafumanisa ukuba kwiimodeli zokubala, izicwangciso ezahlukeneyo zokudibanisa, kunokuba kumanqanaba okuguquguquka okuhambelanayo, kunokucacisa ngcono i-CPU kwi-dorsal media-superior temporal neurons.Utshintsho lweshiti lokuqaphela umsebenzi wokuqaphela (MSTd).Isiphumo esincinci sokukhetha esisibonileyo kwi-MT mhlawumbi sibonisa ukudityaniswa okubanzi kwee-neurons ezininzi ezisezantsi zolwazi ukwenza iimbono zokuhambelana okanye ukungafihli.Kwimeko nayiphi na imeko apho iimpawu zentshukumo zendawo kufuneka zihlanganiswe zibe yinto enye okanye ezimbini (i-bistable gratings) okanye iindawo ezihlukeneyo zezinto eziqhelekileyo (i-3-D SFM), ubungqina obuzimeleyo bokuba iimpendulo ze-MT zidibene kakhulu nezigwebo zokuqonda, kwakukho iimpendulo ezinamandla zeMT.kucetywa ukuba udlale indima ekwahlulweni kwemifanekiso entsonkothileyo kwimiboniso yezinto ezininzi usebenzisa ulwazi lwentshukumo olubonakalayo.
Njengoko kukhankanyiwe ngasentla, sibe ngabokuqala ukuchaza unxibelelwano phakathi komsebenzi weselula we-MT kunye nokuqonda.Njengoko kuqulunqwe kwimodeli yokuqala yamanqanaba amabini nguMovshon kunye noogxa, iyunithi yemowudi yinqanaba lemveliso yeMT.Nangona kunjalo, umsebenzi wamva nje ubonise ukuba imo kunye neeseli zecandelo zimelela iziphelo ezahlukeneyo zokuqhubeka kunye nokuba ukungafani kweparametric kwisakhiwo sendawo yokwamkela kujongene ne-spectrum yokulungiswa kwamacandelo emowudi.Ngoko ke, sifumene ulungelelwaniso olubalulekileyo phakathi kwe-CP kunye ne-PI, efana nobudlelwane phakathi kwe-binocular mismatch adjustment symmetry kunye ne-CP kumsebenzi wokuqaphela ubunzulu okanye ulungelelwaniso lokumisela ulungelelwaniso kumsebenzi wocalucalulo olufanelekileyo.Ubudlelwane phakathi kwamaxwebhu kunye ne-CP 33.U-Wang kunye ne-Movshon62 bahlalutye inani elikhulu leeseli kunye ne-MT yokukhetha ukhetho kwaye bafumanisa ukuba, ngokomyinge, isalathisi semowudi sasinxulumene neendawo ezininzi zokulungisa, ebonisa ukuba ukhetho lwemowudi lukhona kwezinye iintlobo ezininzi zeempawu ezinokufundwa kwi-MT yabemi..Ngoko ke, kwizifundo zexesha elizayo zobudlelwane phakathi komsebenzi we-MT kunye nombono we-subjective, kuya kubaluleka ukugqiba ukuba isalathisi sepateni sihambelana ngokufana nomnye umsebenzi kunye neempawu zokukhetha i-stimulus, okanye ukuba olu budlelwane buchanekile kwimeko yokwahlulwa kokuqonda.
Ngokufanayo, i-Nienborg kunye ne-Cumming 42 yafumanisa ukuba nangona iiseli ezikufutshane nezikude zikhethele ukungahambelani kwebinocular kwi-V2 zazinovakalelo ngokulinganayo kumsebenzi wocalucalulo olunzulu, kuphela ngabantu beeseli abasondele-bakhethayo ababonisa i-CP ebalulekileyo.Nangona kunjalo, ukuqeqesha iinkawu kwakhona ukuba zikhethe ubunzima bokuhluka okukude kubangele ii-CPs ezibalulekileyo kwiikheji ezithandwayo.Olunye uphando luye lwaxela ukuba imbali yoqeqesho ixhomekeke kwi-perceptual correlations34,40,63 okanye ubudlelwane be-causal phakathi komsebenzi we-MT kunye nocalucalulo oluhlukeneyo48.Ubudlelwane esibubonileyo phakathi kwe-CP kunye nokukhethwa kolwalathiso lwerejimeni kusenokwenzeka ukuba kubonisa iqhinga elithile elisetyenziselwe iinkawu ukusombulula ingxaki yethu, kwaye hayi indima ethile yeempawu zokukhetha imo kumbono wokubonwa kwemoto.Kumsebenzi wexesha elizayo, kuya kubaluleka ukugqiba ukuba imbali yokufunda inempembelelo ebalulekileyo ekunqumeni ukuba zeziphi iimpawu zeMT ezilinganiswe ngokufanelekileyo kunye nokuguquguqukayo ukwenza izigwebo zokwahlula.
I-Stoner kunye noogxa14,23 babe ngabokuqala ukunikezela ingxelo yokuba ukuguqula ukukhanya kwemimandla yegridi edibeneyo ngokuqikelelwayo kuchaphazele ukuhambelana kunye nokungafihli kweengxelo zomkhangeli womntu kunye nohlengahlengiso lwesikhokelo kwi-macaque MT neurons.Ababhali bafumanise ukuba xa ukuqaqamba kwemimandla egqagqeneyo kuhambelana nokucaca, abakhi-mkhanyo babika imbono ecacileyo ngakumbi, ngelixa i-MT neurons ibonakalisa ukunyakaza kwamacandelo e-raster.Ngokwahlukileyo, xa ukukhanya okugqithekileyo kunye nokugqithiswa okubonakalayo kungahambelani ngokwasemzimbeni, umbonisi ubona intshukumo ehambelanayo, kwaye i-MT neurons ibonakalisa intshukumo yehlabathi yepateni.Ngaloo ndlela, ezi zifundo zibonisa ukuba utshintsho lomzimba kwi-stimuli ebonakalayo echaphazela ngokuthembekileyo iingxelo zezahlulo nazo zenza utshintsho oluqikelelweyo kwi-MT arousal.Umsebenzi wakutshanje kule ndawo uhlolisise ukuba yeyiphi imiqondiso yeMT elandelelanisa imbonakalo yombono we-stimuli enzima18,24,64.Ngokomzekelo, i-subset ye-MT neurons ibonakaliswe ukuba ibonise ukulungiswa kwe-bimodal kwimephu ye-random motion map (RDK) kunye nezalathiso ezimbini ezingaphantsi kokwahlukana kune-RDK engabonakaliyo.Ububanzi bokulungelelaniswa kweselula 19, 25.Ababukeli bahlala bebona ipateni yokuqala njengentshukumo ecacileyo, nangona i-MT neurons ezininzi zibonisa ukulungelelaniswa okungafaniyo ekuphenduleni ezi zivuseleli, kwaye i-avareji elula yazo zonke iiseli ze-MT zinika impendulo yabantu abangabodwa.Ke, iseti engaphantsi yeeseli ezibonisa i-bimodal tuning inokwenza i-neural substrate yalo mbono.Okubangela umdla kukuba, kwii-marmosets, eli nani labemi lihambelana neeseli zePDS xa zivavanywa kusetyenziswa igridi eqhelekileyo kunye negridi yokuvuselela.
Iziphumo zethu zihamba inyathelo elinye ngaphezu kwezi zingentla, ezibaluleke kakhulu ekusekeni indima ye-MT ekuhlukaniseni ingqiqo.Enyanisweni, ulwahlulo yinto eyenzeka ngokuzimeleyo.Uninzi lwemiboniso ebonakalayo ye-polystable ibonisa amandla enkqubo yokujonga ukucwangcisa nokutolika i-stimuli eqhubekayo ngeendlela ezingaphezu kweyodwa.Ngexesha elifanayo ukuqokelela iimpendulo ze-neural kunye neengxelo zokuqonda kwisifundo sethu kwasivumela ukuba sihlolisise i-covariance phakathi kwezinga lokudubula kweMT kunye nokutolikwa kwengqondo yokuvuselela rhoqo.Emva kokubonisa olu budlelwane, siyavuma ukuba ulwalathiso lwe-causality aluzange lusekwe, oko kukuthi, iimvavanyo ezongezelelweyo ziyafuneka ukuze kuqinisekiswe ukuba isibonakaliso sokwahlulwa kokuqonda okubonwa ngathi, njengoko abanye bephikisana [65, 66, 67], ngokuzenzekelayo.Inkqubo iphinda imele izibonakaliso ezihlayo ezibuyela kwi-cortex ye-sensory kwiindawo eziphakamileyo 68, 69, 70 (umzobo 8).Iingxelo zenxalenye enkulu yeeseli ezikhethiweyo zepateni kwi-MSTd71, enye yeethagethi eziphambili ze-cortical ye-MT, zibonisa ukuba ukwandisa olu mvavanyo ukubandakanya ukurekhoda kwangaxeshanye kwe-MT kunye ne-MSTd kuya kuba linyathelo lokuqala elilungileyo ekuqondeni ngakumbi iindlela ze-neural zokuqonda.ukwahlulahlula.
Imodeli yamanqanaba amabini ecandelo kunye nemowudi yokukhetha ukhetho kunye nefuthe elinokubakho lempendulo ephezulu-phantsi malunga nomsebenzi onxulumene nokukhetha ekuguquleleni komatshini.Apha, i-mode direction selectivity (PDS - "P") kwisinyathelo se-MT senziwa (i) isampula enkulu yedatha ekhethiweyo yokufakwayo ehambelanayo nesantya semowudi ethile, kunye (ii) nokunyanzeliswa kokulungiswa okunamandla.I-directionally selective (CDS) icandelo leMT ("C") inqanaba linoluhlu oluncinci lwesampulu kwindlela yokufaka kwaye ayinakho ukunyanzeliswa kwe-tuning.Uthintelo olungaguqukiyo lunika ulawulo kubo bobabini abantu.Iintolo ezinemibala zibonisa ukuqhelaniswa nesixhobo esikhethwayo.Ukucacisa, kuphela iseti esezantsi yoqhagamshelo lwe-V1-MT kunye nemowudi yecandelo elinye kunye nebhokisi yokukhetha uqhelaniso eboniswayo.Kumxholo wokutolika iziphumo zethu ze-feed-forward (FF), ukusetwa kwegalelo elibanzi kunye ne-inhibition eqinile yokuguqula (ekugqanyiswe kubomvu) kwiiseli ze-PDS zenze umahluko omkhulu kumsebenzi ekuphenduleni kwiipatheni ezininzi zokuhamba.Kwingxaki yethu yokwahlulahlula, eli qela liqhuba amakhonkco ezigqibo kwaye ligqwetha imbono.Ngokwahlukileyo, kwimeko yempendulo (i-FB), izigqibo zokuqonda ziveliswa kwiisekethe eziphakamileyo ngedatha ye-sensory kunye ne-cognitive biases, kunye nefuthe elikhulu le-FB esezantsi kwiiseli ze-PDS (imigca engqingqwa) ivelisa izibonakaliso zokukhetha.b Umelo olucwangcisiweyo lwezinye iimodeli zeCDS kunye nezixhobo zePDS.Apha iimpawu ze-PDS kwi-MT ziveliswa kungekuphela nje ngegalelo elithe ngqo le-V1, kodwa nangegalelo elingathanga ngqo le-V1-V2-MT indlela.Iindlela ezingathanga ngqo zemodeli zilungelelaniswa ukunika ukhetho kwimida yokuthungwa (iindawo ezinqamlezileyo zegridi).Imodyuli ye-CDS yomaleko weMT yenza isixa esilinganisiweyo segalelo elithe ngqo kunye nelingathanga ngqo kwaye lithumela imveliso kwimodyuli yePDS.I-PDS ilawulwa ngokuthintela ukukhuphisana.Kwakhona, kuphela ezo ziqhagamshelo ziboniswa ukuba ziyimfuneko ukuzoba isiseko soyilo lwemodeli.Apha, indlela ye-FF eyahlukileyo kunaleyo icetywayo kwi-a ingakhokelela ekuhlukeni okukhulu kwimpendulo ye-lattice yeselula kwi-PDS, kwakhona ekhokelela ekukhetheni kwiipatheni zesigqibo.Ngenye indlela, i-CP enkulu kwiiseli ze-PDS isenokuba sisiphumo sokungakhethi amandla okanye ukusebenza kakuhle kwe-FB encamathiselwe kwiiseli zePDS.Ubungqina buxhasa imodeli ye-MT PDS enemigangatho emibini nantathu kunye ne-CP FF kunye notoliko lwe-FB.
I-macaques yabantu abadala ababini (i-macaca mulatta), enye indoda kunye neyesifazana (i-7 kunye ne-5 iminyaka ngokulandelanayo), ubunzima be-4.5 ukuya kwi-9.0 kg, isetyenziswe njengezinto zokufunda.Ngaphambi kwazo zonke iimvavanyo zotyando oluyinyumba, izilwanyana zafakelwa kwigumbi lokurekhoda elenziwe ngokwezifiso ze-electrode ezithe nkqo ezisondela kwindawo ye-MT, indawo yokumisa yentsimbi engenasici (i-Crist Instruments, i-Hagerstown, MD), kunye nesikhundla samehlo kunye ne-coil yokukhangela i-scleral.(Cooner Wire, San Diego, California).Zonke iiprotocol zihambelana nemimiselo yeSebe lezoLimo lase-United States (i-USDA) kunye ne-National Institutes of Health (NIH) Izikhokelo zoLondolozo loLuntu kunye nokuSetyenziswa kweZilwanyana zeLabhorethri kwaye zivunyiwe yiYunivesithi yaseChicago Institutional Animal Care and Use Committee (IAUKC).
Zonke izivuseleli ezibonwayo zaboniswa kwindawo engqukuva ngokuchasene nemvelaphi emnyama okanye engwevu.Ngexesha lokurekhoda, isikhundla kunye nobubanzi balo mngxuma kwahlengahlengiswa ngokuhambelana nentsimi ye-classical receptive ye-neurons kwi-tip electrode.Sisebenzise iindidi ezimbini ezibanzi ze-stimuli ebonakalayo: i-psychometric stimuli kunye ne-tuning stimuli.
I-psychometric stimulus yi-grating pattern (i-20 cd / m2, i-50% umehluko, i-50% ye-duty cycle, i-5 degrees / i-sec) eyenziwe ngokugqithisa i-gratings ezimbini ezixande zikhukuliseka kwi-directional perpendicular direction (Fig. 1b).Kuye kwaboniswa ngaphambili ukuba ababukeli bomntu babona le patheni yegridi njenge-stimuli ye-bistable, ngamanye amaxesha njengepatheni enye ehamba ngendlela efanayo (intshukumo ehambelanayo) kwaye ngamanye amaxesha njengeendawo ezimbini ezihlukeneyo ezihamba ngeendlela ezahlukeneyo (ukunyakaza okucacileyo).Amacandelo epateni yelathisi, ejoliswe symmetrically - i-engile phakathi kweeletisi ukusuka 95 ° ukuya 130 ° (etsalwa kwisethi: 95 °, 100 °, 105 °, 115 °, 120 °, 125 °, 130 ° °, kuyo yonke iseshoni Isolation1 angle - 5 ° sigcina i-engile ye-Isolation1, malunga ne-neurons ye-psychod ayibandakanyi apha. ° okanye 270 ° (pattern orientation).Kwiseshoni nganye, ikona enye kuphela ye-interlattice lattice yasetyenziswa;ngexesha leseshoni nganye, ukuqhelaniswa kwepateni yovavanyo ngalunye kukhethwe ngokungenamkhethe kwizinto ezimbini ezinokwenzeka.
Ukuze singagxininisi imbono yegridi kunye nokubonelela ngesiseko sobungqina bomvuzo wesenzo, sazisa i-random point textures kwibar yokukhanya inyathelo lama-72 kwigridi nganye yecandelo.Oku kuphunyezwa ngokunyuka okanye ukunciphisa (ngexabiso elimiselweyo) ukuqaqamba kweseti ekhethiweyo ekhethiweyo yeepikseli (umzobo 1c).Isalathiso sentshukumo yokuthungwa inika umqondiso oqinileyo oguqula imbono yombukeli kwintshukumo ehambelanayo okanye ecacileyo (Umfanekiso 1c).Ngaphantsi kweemeko ezihambelanayo, zonke izinto eziphathekayo, kungakhathaliseki ukuba yeyiphi icandelo le-texture lattice covers, ziguqulelwe kwicala lepatheni (umzobo 1c, ohambelanayo).Kwimeko ecacileyo, ukuthungwa kuhamba nge-perpendicular ukuya kwicala le-grating eligubungelayo (umzobo 1c, obala) (i-Movie eyongezelelweyo 1).Ukulawula ubunzima bomsebenzi, kwiiseshoni ezininzi ukuchasana kweMichelson (Lmax-Lmin / Lmax + Lmin) kolu phawu lokuthungwa luhluke kwisethi (-80, -40, -20, -10, -5, 0, 5)., 10, 20, 40, 80).Ukuchasana kuchazwa njengokukhanya okuhambelanayo kwe-raster (ngoko ixabiso elichaseneyo le-80% liya kubangela ukuthungwa kwe-36 okanye i-6 cd / m2).Kwiiseshoni ezi-6 kwi-monkey N kunye neeseshoni ezi-5 kwi-monkey S, sasebenzisa uluhlu oluncinci lokungafani okubhaliweyo (-30, -20, -15, -10, -5, 0, 5, 10, 15, 20, 30), apho iimpawu ze-psychophysical zilandela ipateni efanayo njengoluhlu olupheleleyo, kodwa ngaphandle kokuhluthwa.
I-Tuning stimuli i-sinusoidal grids (i-contrast 50%, i-1 cycle / degree, i-5 degrees / sec) ihamba kwenye ye-16 yeendlela ezilinganayo, okanye iigridi ze-sinusoidal ezihamba kula macala (ezibandakanya ezimbini ezichasene ne-135 ° ii-angles ezibekwe phezulu kwi-sinusoidal gratings phezu komnye nomnye).kwicala elinye lepateni.