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Kuona kwepamusoro-resolution kunoda yakanaka retinal sampling uye kubatanidzwa kuvakazve zvivakwa zvechinhu.Zvakakosha kucherechedza kuti kana uchisanganisa masampuli emunharaunda kubva kune zvinhu zvakasiyana-siyana, kururamisa kunorasika.Naizvozvo, kupatsanura, kuunganidzwa kwenzvimbo dzechifananidzo kuti zvigadziriswe zvakasiyana, zvakakosha pakuona.Mubasa rakapfuura, bistable lattice zvimiro, izvo zvinogona kutariswa seimwe kana kupfuura inofamba nzvimbo, yakashandiswa kudzidza maitiro aya.Pano, isu tinoshuma hukama pakati pechiitwa uye segmentation kutonga munzvimbo dzepakati dzenzira yekuona yeprimate.Kunyanya, takaona kuti kusarudza kutarisisa kwemukati menguva neurons kwainzwa kune mameseji cues anoshandiswa kukanganisa maonero emabistable gratings uye yakaratidza kuwirirana kwakakosha pakati pemiedzo uye kuzviisa pasi pekuona kwekuramba kuchikurudzira.Uku kuwirirana kwakakura mumayuniti anoratidza mafambiro epasi rose mumapateni ane akawanda enzvimbo.Nekudaro, isu tinogumisa kuti iyo yepakati nguva domain ine masaini anoshandiswa kupatsanura zvakaomarara kuita zvinhu zvinoumba uye nzvimbo.
Vision inotsamira kwete chete parusarura chairwo rwekutanga mufananidzo hunhu senge kumucheto uye kumhanya, asi zvakanyanya kukosha pakubatanidzwa kwakaringana kwehunhu uhwu kuverenga zvimiro zvezvakatipoteredza zvakaita sechimiro chechinhu uye trajectory1.Nekudaro, matambudziko anomuka kana retinal mifananidzo inotsigira akati wandei akaenzana anonzwisisika emhando mapoka 2, 3, 4 (Fig. 1a).Semuenzaniso, kana mapoka maviri ezviratidzo zvekumhanya ari pedyo zvakanyanya, izvi zvinogona kududzirwa sechinhu chimwe chinofamba kana zvinhu zvakawanda zvinofamba (Fig. 1b).Izvi zvinoratidzira chimiro chekuzvimiririra chechikamu, kureva kuti haisi pfuma yakatarwa yemufananidzo, asi nzira yekududzira.Pasinei nekukosha kwayo kuri pachena kwekuona kwakajairika, kunzwisisa kwedu kweiyo neural hwaro hwekuona segmentation inoramba isina kukwana zvachose.
Mufananidzo wekatuni wedambudziko rekuona segmentation.Maonero emucherechedzi ekudzika muNecker cube (kuruboshwe) anochinjanisa pakati petsananguro mbiri dzinobvira (kurudyi).Izvi zvinodaro nekuti hapana masaini mumufananidzo anobvumira uropi kuti husarudzike iyo nhamba ye3D kutaridzika (inopihwa ne monocular occlusion chiratidzo kurudyi).b Kana masaini akawanda ekufamba achiiswa munzvimbo iri padhuze, sisitimu yekuona inofanirwa kuona kuti samples dzenzvimbo dziri kubva kune chimwe kana zvimwe zvinhu.Kusanzwisiswa kurimo kwezviratidzo zvemafambiro enzvimbo, kureva kutevedzana kwekufamba kwechinhu kunogona kuburitsa kufamba kwakafanana kwenzvimbo, zvichikonzera kududzira kwakafanana kwakaenzana kwemaonero ekuisa, kureva minda yevekita pano inogona kusimuka kubva mukufamba kwakabatana kwenzvimbo imwe chete kana kufamba kwakajeka kwenzvimbo dzakapiringana.c (kuruboshwe) Muenzaniso wekukurudzira kwegridi yedu.Rectangular gratings drifting perpendicular to their direction ("component directions" - white miseve) inopindirana kuti iite grating pateni.Iyo latisi inogona kunzwisiswa seimwe, yakajairwa, yakabatana mafambiro enzira (tsvuku miseve) kana kufamba kwakajeka kwemukomboni nzira.Maonero e lattice anokanganiswa nekuwedzera kwe random point texture cues.(Pakati) Nzvimbo yakamisikidzwa muyero inowedzerwa uye inoratidzwa senhevedzano yemafuremu ezviratidziro zvinopindirana uye zvakajeka, zvichiteerana.Kufamba kwedoti mune yega yega inomiririrwa negirinhi uye tsvuku miseve.(Kurudyi) Girafu yenzvimbo (x, y) yenzvimbo yekusarudza maringe nenhamba yemafuremu.Muchiitiko chakabatana, zvese zvimiro zvinokwevera munzira imwechete.Panyaya yekujeka, kuumbwa kunofamba nenzira yechikamu.d Mufananidzo wekatuni webasa redu rechikamu chekufamba.Tsoko dzakatanga kuedza kwega kwega nekugadzirisa kadonhwe kadiki.Mushure mekunonoka kupfupi, imwe mhando ye grating pateni (kubatana / kujeka) uye maumbirwo echiratidzo saizi (semuenzaniso musiyano) yakaonekwa panzvimbo yeMT RF.Panguva yekuedzwa kwega kwega, giraiti rinogona kukukurwa mune imwe yenzira mbiri dzinogoneka dzepateni.Mushure mekusimudzira kubviswa, zvibodzwa zvekusarudza zvakaonekwa pamusoro uye pazasi MT RF.Tsoko dzinofanirwa kuratidza maonerwo adzo egidhi mumasakedhi kune yakakodzera kusarudzwa chinangwa.
Kugadziriswa kwekufamba kwekuona kwakanyatso kuratidzwa uye nekudaro kunopa yakanakisa modhi yekudzidza neural maseketi ekuona segmentation.Ongororo dzinoverengeka dzemakomputa dzakacherekedza kushandiswa kwematanho maviri-matanho ekugadzirisa mafambiro umo yakakwirira-resolution yekutanga fungidziro inoteverwa nekusarudza kubatanidzwa kwemasampuli emunharaunda kubvisa ruzha uye kudzorera chinhu velocity7,8.Izvo zvakakosha kuti uzive kuti masisitimu ekuona anofanirwa kungwarira kudzikamisa iyi ensemble kune iwo chete emunharaunda samples kubva kune zvakajairika zvinhu.Zvidzidzo zve Psychophysical zvakatsanangura zvinhu zvemuviri zvinopesvedzera kuti masiginecha emunharaunda akakamurwa sei, asi chimiro cheanatomical trajectories uye neural codes inoramba iri mibvunzo yakavhurika.Mishumo yakawanda inoratidza kuti kutarisisa-selective masero munzvimbo yenguva (MT) yeprimate cortex vanokwikwidza neural substrates.
Zvakakosha, mune izvi zviedzo zvakapfuura, shanduko mukuita kweneural yakabatana nekuchinja kwemuviri mukukurudzira kwekuona.Zvisinei, sezvataurwa pamusoro apa, kuparadzanisa inzira yekunzwisisa.Naizvozvo, chidzidzo cheiyo neural substrate chinoda kubatanidza shanduko muchiitiko cheneural nekuchinja mumaonero ezvakagadziriswa zvinokurudzira.Naizvozvo, takadzidzisa tsoko mbiri kuti dzitaure kana iyo inofungirwa bhistable grating pateni yakaumbwa nepamusoro-soro inodonhedza rectangular girati yaive nzvimbo imwechete kana mbiri dzakazvimirira.Kuti tidzidze hukama pakati pekuita neural uye segmentation kutonga, takanyora chiitiko chimwe chete muMT apo tsoko dzakaita basa iri.
Takawana kuwirirana kwakakosha pakati pekudzidza kweMT basa uye maonero.Uku kuwirirana kwaivepo kunyangwe iyo yekukurudzira yaive neapachena segmentation cues.Mukuwedzera, kusimba kweichi chiitiko kune chekuita nekunzwa kune zvikwangwani zvezvikamu, pamwe neiyo index index.Iyo yekupedzisira inotara dhigirii iyo unit inopenya pasi rose kwete kufamba kwenzvimbo mumapateni akaomarara.Kunyange zvazvo kusarudzwa kwehutungamiri hwemafashoni kwagara kuchizivikanwa sechinhu chinotsanangura cheMT, uye masero anosarudza fashoni anoratidza kugadzirisa kune zvakaoma zvinokurudzira zvinopindirana nemafungiro evanhu kune izvo zvinokurudzira, kusvika pakuziva kwedu, uhwu ndihwo humbowo hwekutanga hwekubatana pakati pemaitiro.index uye segmentation yemafungiro.
Takadzidzisa tsoko mbiri kuratidza maonero adzo ekukukurwa kwegridi zvinokurudzira (kufamba kwakabatana kana kujeka).Vacherechedzi vevanhu vanowanzoona izvi zvinokurudzira sekufamba kwakabatana kana kwakajeka kweanenge frequency yakafanana.Kuti tipe mhinduro chaiyo mumuedzo uyu uye kuisa hwaro hwemubairo wekushanda, takagadzira zvikwangwani zvezvikamu nekutumira mameseji raster yechikamu chinoumba lattice (Fig. 1c, d).Pasi pemamiriro ezvinhu akabatana, zvinyorwa zvose zvinofamba nenzira yemuenzaniso (Fig. 1c, "coherent").Mumamiriro ezvinhu akajeka, chimiro chinofambisa perpendicular kune kutungamira kwegrating iyo yakanyanyisa (Fig. 1c, "transparent").Isu tinodzora kuoma kwebasa nekushandura mutsauko weiyi mavara label.Mumiyedzo yakacherechedzwa, tsoko dzakapihwa mubairo wemhinduro dzinoenderana nemavara emavara, uye mibairo yaipihwa zvisina tsarukano (50/50 odds) mumiedzo ine mapatani pasina mameseji cues (zero texture musiyano mamiriro).
Maitiro ekuzvibata kubva mumiedzo miviri yevamiriri inoratidzwa muMufananidzo 2a, uye mhinduro dzakarongwa sechikamu chemitongo yekubatana zvichienderana nekusiyana kwezviratidzo zvemavara (kujeka kusiyanisa kunofungidzirwa kuve kwakashata netsanangudzo) kune mapatani anochinja kumusoro kana pasi, zvichiteerana. Pakazere, maonero etsoko ekubatana/kubudira pachena akabatwa nekuvimbika nechiratidzo (pachena, chakabatana) uye simba (kusiyana) kwechimiro chemavara (ANOVA; tsoko N: kutungamira - F = 0.58, p = 0.45, chiratidzo - F = 1248, p <10−0 −1 = 1, mon-1, mutsauko; S: kutungamira - F = 0.41, p = 0.52, chiratidzo - F = 2876.7, p <10-10, kusiyana - F = 36.5, p <10-10). Pakazere, maonero etsoko ekubatana/kujeka akakanganiswa nechiratidzo (pachena, chakabatana) uye simba (kusiyana) kwechimiro chemavara (ANOVA; tsoko N: kutungamira - F = 0.58, p = 0.45, chiratidzo - F = 1248, p <10-10 = 0 = 1, mon −1 = 1, 2 −1 = 1, 2 −1 = 1, 2, p <10 − 3, −1 = kiyi S: kutungamira - F = 0.41, p = 0.52, chiratidzo - F = 2876.7, p <10-10, kusiyana - F = 36.5, p <10-10). В целом на восприятие обезьянами когерентности/прозрачности достоверно влияли как знак (прозрачность, когерентность), так и сила (кон); обезьяна N: направление — F = 0,58, p = 0,45, знак — F = 1248, p < 10−10, контраст – F = 22,63, p < 10; −10 обезья - 5, p = 2, 63, p <10; ризнак – F = 2876,7, p <10−10, контраст – F = 36,5, р <10-10). Kazhinji, maonero ekubatana / kujekesa netsoko zvakakanganiswa zvakanyanya nechiratidzo (kujeka, kuwirirana) uye simba (kusiyana) kwechimiro chemavara (ANOVA; monkey N: kutungamira - F = 0.58, p = 0.45, chiratidzo - F = 1248, p <10- = 3 <10, 3, 10 , monkey - 10 : 10, 1 monkey - 10: 1 monkey; F = 0.41, p = 0.52, chiratidzo - F = 2876.7, p <10 -10, kusiyana - 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； 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: 幑 = S = 22.63, p <10;–10: S = F = 10;–10: S = F = S = 10;−1: 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;−0 010 <10;−0010.Kazhinji, maonero ekubatana kwemonkey / kujekesa kwakakanganiswa zvakanyanya nechiratidzo (kujeka, kuwirirana) uye kusimba (kusiyana) kwemavara ezvinyorwa (ANOVA);обезьяна N: ориентация – F = 0,58, p = 0,45, знак – F = 1248, p < 10−10, Контрастность — F = 22,63, p <10; monkey N: kutaridzika - F = 0.58, p = 0.45, chiratidzo - F = 1248, p <10-10, Kusiyana - 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 Tsoko S: Kutarisana - F = 0.41, p = 0.52, Chiratidzo - F = 2876.7, p <10-10, Kusiyana - F = 36.5, p <10-10).Gaussian cumulative mabasa akaiswa kune data kubva pachikamu chega chega kuratidza psychophysical maitiro etsoko.Pamusoro pemuonde.2b inoratidza kugoverwa kwechibvumirano chemhando idzi pazvikamu zvese zvetsoko dzese.Pakazara, tsoko dzakapedza basa nemazvo uye zvakangofanana, uye isu takaramba isingasviki 13% yemasesheni etsoko mbiri nekuda kwekutadza kukwana kune yakawedzera Gaussian modhi.
a Behavioral mienzaniso yetsoko mumimiriri yezvikamu (n ≥ 20 miedzo pamamiriro ekusimudzira).Kuruboshwe (kurudyi) mapaneru, data kubva kune imwe N (S) tsoko sesheni inorongwa seyakabatana sarudzo zvibodzwa (ordinate) maringe nechiratidzo chakasiyana chemavara masaini (abscissa).Pano panofungidzirwa kuti mavara akajeka (akabatana) ane maitiro asina kunaka (akanaka).Mhinduro dzakavakwa zvakasiyana maererano negwara rekufamba kwepateni (kumusoro (90 °) kana pasi (270 °)) muyedzo.Pamhuka dzose dziri mbiri, kuita, ingave mhinduro yakakamurwa ne50/50 mutsauko (PSE - miseve yakasimba) kana kuwanda kwekusiyana kwemavara kunodiwa kutsigira imwe nhanho yekuita (chikumbaridzo - miseve yakavhurika), iri mune idzi nzira dzekukukurwa.b Yakakwana histogram yeR2 kukosha kweGaussian cumulative basa.Tsoko S(N) data inoratidzwa kuruboshwe (kurudyi).c (Pamusoro) PSE yakayerwa kuti grid yakachinjika pasi (yakarongeka) kana ichienzaniswa nePSE yakasimudza grid (abscissa) yakarongwa, ine micheto inomiririra kugoverwa kwePSE kune yega yega mamiriro uye miseve inoratidza zvinoreva mamiriro ese.Data yeese N(S) tsoko masesheni anopiwa kuruboshwe (kurudyi) column.(Pazasi) Same gungano se data rePSE, asi kune akaiswa maficha zvikumbaridzo.Pakanga pasina misiyano yakakosha muPSE zvikumbaridzo kana fashoni mafambiro (ona chinyorwa).d PSE uye mutserendende (ordinate) zvakarongwa zvichienderana neyakajairwa raster kutaridzika kweangular separation chikamu ("integral grating angle" - abscissa).Denderedzwa rakavhurika ndiyo nzira, mutsara wakasimba ndiyo wakanakisa wakakodzera regression modhi, uye mutsara une doti ndiyo 95% yekuvimbika kupindirana kweiyo regression modhi.Pane kuwirirana kwakakosha pakati pePSE uye yakajairwa yekubatanidza kona, asi kwete kutsetseka uye yakajairwa yekubatanidza kona, zvichiratidza kuti psychometric basa rinoshanduka sezvo kona inopatsanura mareti echikamu, asi kwete kurodza kana kubata kumeso.( Tsoko N, n = 32 zvikamu; Tsoko S, n = 43 zvikamu).Mumapanera ese, mabhawa ekukanganisa anomiririra kukanganisa kwakajairwa kwezvinoreva.Haha.Kubatana, PSE subjective kuenzana mamakisi, zvakajairwa.standardization.
Sezvakataurwa pamusoro, zvese mutsauko wezvinyorwa zvemavara uye mafambiro ekufamba kwepateni zvakasiyana pamiyedzo, paine zvinokurudzira zvichikwira kumusoro kana pasi muyedzo yakapihwa.Izvi zvinoitirwa kuderedza psychophysical11 uye neuronal28 adaptive mhedzisiro.Patani inotarisana neyakarerekera (subjective equality point kana PSE) (Wilcoxon rank sum test; tsoko N: z = 0.25, p = 0.8; tsoko S: z = 0.86, p = 0.39) kana chikumbaridzo chakaiswa (chiverengero cheWilcoxon chiyero: 1, monkey = 1, monkey = 4, monkey; z = 0.49, p = 0.62) (Mufananidzo 2c).Pamusoro pezvo, pakanga pasina misiyano yakakosha pakati petsoko muchiyero chekusiyana kwemavara anodiwa kuchengetedza maitiro echikumbaridzo (N tsoko = 24.5% ± 3.9%, S tsoko = 18.9% ± 1.9%; Wilcoxon rank sum , z = 1.01, p = 0.31).
Muchikamu chega chega, isu takashandura iyo interlattice angle inoparadzanisa matarisiro echikamu chelattices.Zvidzidzo zvePsychophysical zvakaratidza kuti vanhu vanogona kuona sero 10 seyakabatana kana kona iyi idiki.Kana tsoko dzaitaura zvakavimbika maonero adzo ekubatana/kubudirana pachena, zvichibva pane zvakawanikwa izvi, munhu aitarisira PSE, mutsauko wemavara unoenderana nekupatsanurwa kweyunifomu pakati pekubatana uye kujeka kwesarudzo, kuwedzera pakudyidzana.lattice angle. Izvi ndizvo chaizvo zvakanga zvakaita (Fig. 2d; kukoromoka nepakati penzira, Kruskal-Wallis; tsoko N: χ2 = 23.06, p <10−3; tsoko S: χ2 = 22.22, p <10-3; kuwirirana pakati pe-normalized inter-grating −3, pSE = 6, pSE = 0, pSE = 1, 0 . 9; tsoko S: r = 0.76, p <10−13). Izvi ndizvo chaizvo zvakanga zvakaita (Fig. 2d; kukoromoka nemativi epatani, Kruskal-Wallis; tsoko N: χ2 = 23.06, p <10−3; tsoko S: χ2 = 22.22, p <10-3; kuwirirana pakati pe-normalized inter- 0 −3 angle---------------------------------------------------3 ne-monkey--------------------------------------------------------------- monkey ne-monkey angle-7-------------------------------------------3 9; tsoko S: r = 0.76, p <10−13). Это действительно имело место (рис. 2d; коллапс поперек направления паттерна, Крускал-Уоллис; обезьяна N: χ2 = 23,06, pya 2,2 = 23,06, 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. Izvi zvakaitika zvechokwadi (Fig. 2d; kuputsika mhiri kwegwaro remuenzaniso, Kruskal-Wallis; monkey N: χ2 = 23.06, p <10-3; monkey S: χ2 = 22.22, p <10-3; kuwirirana pakati pe normalized lattice angle uye PSE: 6: 0, monkey N, 9, pSE = monkey. r = 0.76, p <10-13).情况确实如此（图2d；跨模式方向折叠，Kruskal-Wallis;子N:χ2 = 23.06，p <10-3猴子S:χ2-3; 2 = 2;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 ; <10-3 ; <10-3 ; <10-3 ; <10-3 ; <10-3 ; <10-3 ; <10-3 ; 阿 : 2 - : 12 = 2 - :栅角 和 pse-猴子 猴子 猴子 猴子 猴子 猴子N：r = 0.67，p <10-9; 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, 3; лизованный межрешеточный угол). Izvi ndizvo zvaive zvechokwadi (Fig. 2d; peta pamwe chete neaxis mode, Kruskal-Wallis; monkey N: χ2 = 23.06, p <10-3; monkey S: χ2 = 22.22, p <10-3; normalized interlattice kona). PSE-обезьяна N: r = 0,67, p < 10–9, обезьяна S: r = 0,76, p < 10–13). PSE tsoko N: r = 0.67, p <10–9, tsoko S: r = 0.76, p <10–13).Kusiyana neizvi, kushandura kona ye interlattice kwakanga kusina chinhu chakakosha pamateru e psychometric function (Fig. 2d; cross-modal orientation fold, Kruskal-Wallis; monkey N: χ2 = 8.09, p = 0.23; monkey S χ2 = 3.18 = 3.18 , p, 7, p. N: r = -0.4, p = 0.2, tsoko S: r = 0.03, p = 0.76).Nekudaro, zvinoenderana neiyo psychophysical data yemunhu, avhareji mhedzisiro yekushandura kona pakati pemagirodhi ndeyekuchinja kwenzvimbo dzekutamisa, uye kwete kuwedzera kana kuderera kwekunzwa kune zvikamu zvezviratidzo.
Chekupedzisira, mibairo inopihwa zvisina tsarukano ine mukana we 0.5 mumiedzo ine zero mavara akasiyana.Dai tsoko dzese dzaiziva nezve kusarudzika uku uye dzichikwanisa kusiyanisa pakati zero mavara akasiyana uye cue stimuli, vaigona kugadzira nzira dzakasiyana dzemhando mbiri dzemiedzo.Ongororo mbiri dzinoratidza zvakasimba kuti izvi handizvo.Chekutanga, kushandura kona yegrating yaive nehunhu hwakafanana mhedzisiro pane cue uye zero manyorerwo akasiyana zvibodzwa (Fig. 2d uye Supplementary Fig. 1).Chechipiri, kune nyani mbiri, bistable kutongwa kusarudzwa hazvibviri kuva kudzokorora kwekupedzisira (yapfuura) kusarudzwa kwemubairo (binomial test, N monkeys: 0.52, z = 0.74, p = 0.22; S tsoko: 0.51, r = 0.9, p = 0.18).
Mukupedzisa, maitiro etsoko mubasa redu rezvikamu aive pasi pekutonga kwakanaka kwekukurudzira.Kutsamira kwekutonga kwekunzwisisa pachiratidzo uye saizi yezvimiro zvemavara, pamwe nekuchinja muPSE ine grating angle, zvinoratidza kuti tsoko dzakashuma maonero adzo epamhepo ekubatana / kujeka.Pakupedzisira, mhinduro dzetsoko mu zero texture misiyano miyedzo haina kukanganiswa nemubairo wenhoroondo yemiedzo yapfuura uye yakakanganiswa zvakanyanya ne-inter-raster angular shanduko.Izvi zvinoratidza kuti tsoko dzinoramba dzichitaura maonero adzo ega ega pasi pemamiriro ezvinhu akakosha aya.
Sezvambotaurwa pamusoro apa, kushanduka kwemavara akasiyana kubva kune yakaipa kusvika kune yakanaka yakaenzana nemafungiro ekuchinja kwekusimudzira kubva pakujeka kuenda kune kubatana.Kazhinji, kune imwe sero, mhinduro yeMT inowanzowedzera kana kuderera sezvo kusiyana kwemavara kunoshanduka kubva kune yakaipa kusvika kune yakanaka, uye kutungamira kwechiitiko ichi kunowanzoenderana negwara rekufamba kwemuenzaniso / chikamu.Semuyenzaniso, kwakananga tuning macurves e2 anomiririra MT maseru anoratidzwa muFigure 3 pamwe nemhinduro dzemasero aya kune magirating ane yakaderera kana yakakwirira musiyano inobatana kana yakajeka masaini masaini.Isu takaedza neimwe nzira kuverengera zvirinani mhinduro dzegidhi, izvo zvinogona kunge zvine hukama nekuita kwe psychophysiological yetsoko dzedu.
Polar plot ye directional tuning curve yeanomiririra tsoko MT cell S mukupindura kune imwechete sinusoidal array.Kona inotaridza mafambiro ekufamba kwegireti, ukuru hunoratidza kubuda, uye yakasarudzika sero kutungamira kunopindirana neanosvika 90 ° (kumusoro) negwara rechimwe chezvikamu mugwara regwati patani.b Weekly stimulus-time histogram (PSTH) yegedhi remhinduro, inochinjirwa mugwara retemplate ne 90° (inoratidzwa schematically kuruboshwe) sero rinoratidzwa mu a.Mhinduro dzakarongwa nemhando yemhando yemhando (yakabatana / yakajeka - yepakati / yekurudyi pani zvichiteerana) uye Michelson musiyano (PSTH color hint).Kuedza kwakarurama chete kunoratidzwa kune imwe neimwe yemhando yepasi-yakasiyana-siyana uye yakakwirira-kusiyanisa masaini masaini.Masero aipindura zvirinani kune anokwira kumusoro elattice mapatani ane ruvara rwakajeka, uye mhinduro kune aya mapatani yakawedzera nekuwedzera kwemavara akasiyana.c, d ndiwo makongiresheni akafanana nea uye b, asi kune MT maseru kunze kwetsoko S, maitiro avanoda anotoda kudarika iwo egidhi inodonha ichidzika.Iyo unit inofarira kudzika kudonhedza magireti ane akabatana manyorerwo cues, uye mhinduro kune aya mapatani inowedzera nekuwedzera kwemavara akasiyana.Mumapanera ese, nzvimbo ine mumvuri inomiririra kukanganisa kwakajairwa kwezvinoreva.Spokes.Spikes, masekondi.chepiri.
Kuti tiongorore hukama huri pakati pelattice surface configuration (yakabatana kana yakajeka) sezvinoratidzwa nemaitiro edu ezvinyorwa uye MT basa, isu takatanga kudzoreredza kuwirirana pakati pemasero ekufambisa kwakabatana (positive slope) kana kufamba kwakajeka (negative slope) nekudzokera shure.yakapihwa kupatsanura maseru nechiyero chekupindura kwechiratidzo zvichienzaniswa nekusiyanisa (zvakasiyana kune yega yega modhi nzira).Mienzaniso yeiyi lattice tuning curves kubva kune imwecheteyo muenzaniso sero muFigure 3 inoratidzwa muMufananidzo 4a.Mushure mekupatsanurwa, takashandisa receiver performance analysis (ROC) kuyera kunzwisiswa kwesero rega rega pakugadzirisa masaini masaini (ona Nzira).Iyo mashandiro eurometric anowanikwa nenzira iyi anogona kuenzaniswa zvakananga nehunhu hwe psychophysical hwetsoko muchikamu chimwe chete kuitira kuenzanisa zvakananga psychophysical senitivity ye neurons kune lattice textures.Isu takaita maviri masaini ekuongorora ekuongorora kune ese mayuniti mumuenzaniso, tichiverenga akaparadzana neurometric maficha kune yega yega nzira yepati (zvakare, kumusoro kana pasi).Zvakakosha kuziva kuti, pakuongorora uku, takangobatanidza miedzo umo (i) zvinokurudzira zvaive nechimiro chemavara uye (ii) tsoko dzakapindura maererano nechocho (kureva, "chaiyo" miedzo).
Mitengo yemoto inorongerwa kusiyanisa chiratidzo chemavara, zvichiteerana, kune gratings inosimuka kumusoro (kuruboshwe) kana pasi (kurudyi), mutsara wakasimba unomiririra yakanakisa kuenderana mutsara regression, uye data iri kumusoro (pasi) mutsara inotorwa kubva kune inoratidzwa muFig.Mupunga.3a Sero, b (Mufananidzo 3c, d).Regression materu maficha akashandiswa kugovera akasarudzika manyorerwo cues (akabatana / akajeka) kune yega yega sero / lattice orientation musanganiswa (n ≥ 20 miedzo pamamiriro ekusimudzira).Error bars inomiririra kutsauswa kwechirevo.Mabasa etsinga dzemayuniti anoratidzwa muba anotsanangurwa pamwe chete nemabasa epfungwa anounganidzwa panguva imwe chete.Ikozvino, kune chimwe nechimwe chinhu, tinoronga sarudzo yakasarudzika yetip (ona zvinyorwa) sechikamu chechikamu chechiratidzo chekusiyanisa (abscissa).Musiyano wemavara wakashandurwa kuitira kuti matipi ezvishandiso anodiwa ave akanaka uye asina chinhu matipi ekushandisa haana kunaka.Iyo data kubva kumusoro (pasi) inodonha grids inoratidzwa kuruboshwe (kurudyi) mapaneru, mumitsetse yepamusoro (pasi) - data kubva kumasero anoratidzwa muFig. 3a, b (Fig. 3c, d).Izvo zviyero zveneurometric uye psychometric threshold (N/P) inoratidzwa pane yega yega.Spokes.Spikes, masekondi.sec, dhairekitori.gwara, dunhu rinodiwa, psi.Psychometry, Neurology.
Lattice tuning curves uye neurometric mabasa e maviri anomiririra MT masero uye anoenderana nawo psychometric mabasa, akaunganidzwa pamwechete nemhinduro idzi, anoratidzwa kumusoro uye pasi mapaneru eMufananidzo 4a,b, zvichiteerana.Maseru aya anoratidza kuwedzera kusinganzwisisike kana kudzikira sezvo ratidziro yemavara inoenda kubva pakujeka kuenda kune inobatana.Pamusoro pezvo, kutungamira uye kusimba kwechisungo ichi zvinoenderana negwara rekufamba kwelatisi.Pakupedzisira, mashandiro eurometric akaverengerwa kubva kumhinduro dzemasero aya akaswedera chete (asi zvakadaro haana kuenderana) maitiro e psychophysical eiyo unidirectional grid movement.Ose ari maviri neurometric uye psychometric mabasa akapfupikiswa nezvikumbaridzo, kureva zvinoenderana ne84% yemusiyano wakasarudzwa nemazvo (zvichienderana nerevo + 1 sd yeyakagadzirirwa cumulative Gaussian basa).Pakati pemuenzaniso wose, chiyero cheN/P, chiyero cheurometric pachikumbaridzo kune psychometric one, yakaenzana ne12.4 ± 1.2 mutsoko N uye 15.9 ± 1.8 mutsoko S, uye kuti lattice ifambe nenzira imwe chete, chete Pa ~ 16% (18).%) zvikamu kubva kutsoko N (tsoko S) (Fig. 5a).Kubva pane cell muenzaniso inoratidzwa mumufananidzo.Sezvinoonekwa muFigure 3 uye 4, kunzwisiswa kwemaneuroni kunogona kukanganiswa nehukama pakati penzvimbo yakasarudzika yesero uye kutungamira kwekufamba kwelattice kunoshandiswa mukuyedza.Kunyanya, iyo orientation adjustment curves muFig. 3a, c inoratidza hukama huri pakati pe neuron orientation setting ye single sinusoidal array uye kunzwisiswa kwayo kune pachena / kuwirirana kwekufamba mune yedu textured array. Izvi ndizvo zvakanga zvakaita tsoko dzose (ANOVA; hama dzaida nzira dzakasungwa ne10° resolution; tsoko N: F = 2.12, p <0.01; tsoko S: F = 2.01, p <0.01). Izvi ndizvo zvakanga zvakaita tsoko dzose (ANOVA; hama dzaida nzira dzakasungwa ne10° resolution; tsoko N: F = 2.12, p <0.01; tsoko S: F = 2.01, p <0.01). Это имело место для обеих обезьян (ANOVA; относительные предпочтительные направления объединены в группы с разрешениение, 10,10; 1; обезьяна S: F = 2,01, p <0,01). Izvi ndizvo zvakanga zvakaita tsoko dzose dziri mbiri (ANOVA; hama dzaida nzira dzakaiswa mu10° resolution; tsoko N: F=2.12, p<0.01; tsoko 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 = 1 p = 2. ; f = 1 p = p. <0.01........） Это имело место для обеих обезьян (ANOVA; относительная предпочтительная ориентация объединена при разрешении 10°; S: F = 2,01, p <0,01). Izvi ndizvo zvanga zvakaita tsoko dzose dziri mbiri (ANOVA; hukama hwakasarudzika hwakaunganidzwa pa10° resolution; tsoko N: F=2.12, p<0.01; tsoko S: F=2.01, p<0.01).Tichifunga hukuru hukuru hwekusiyana kwekunzwa kweneuron (Fig. 5a), kuitira kuti tione kutsamira kwekunzwa kweuron pane hukama hwakasarudzika, takatanga tajaira kutarisisa sero rega rega kune "yakanakisa" kutarisisa kwekufamba kwegridhi pateni (kureva kutungamira).umo gireti rinoumba kona diki pakati pesero rinoda kutariswa uye kutariswa kwepateni yekugezera).Takaona kuti zvikumbaridzo zvemaneuroni (chikumbaridzo che "yakaipisisa" kutarisisa lattice / chikumbaridzo che "yakanakisa" lattice kutarisisa) akasiyana neaya akajairwa maitiro akasarudzika, aine mapeaks echikumbaridzo reshiyo anoitika akatenderedza mapatani kana chikamu chechikamu (Mufananidzo 5b).)). Mhedzisiro iyi haigone kutsanangurwa nekusarura mukugovaniswa kwemaitiro anodiwa muzvikamu mune imwe neimwe sampu yakananga kune imwe yeplaid pateni kana zvikamu zvinongedzo (Fig. 5c; Rayleigh test; monkey N: z = 8.33, p <10-3, denderedzwa zvinoreva = 190.13 deg 30 ± 9; 9 = 190.13 deg 3 deg ± 7, 8 deg 3 deg mon ± 0 ± 9. 5) uye yaienderana mhiri plaid inter-grating angles (Supplementary Fig. 2). Mhedzisiro iyi haigone kutsanangurwa nekusarura mukugoverwa kwemaitiro anodiwa muzvikamu mune imwe neimwe sampu yakananga kune imwe yeplaid patani kana zvikamu zvinongedzo (Fig. 5c; Rayleigh test; monkey N: z = 8.33, p <10-3, denderedzwa zvinoreva = 190.13 deg 30 ± 9; 45) uye yaienderana mhiri plaid inter-grating angles (Supplementary Fig. 2). Этот эффект было объяснить смещением распределения предпочтительных направлений в единицах в каждой выборкей выборке здороженник й или направлений компонентов (рис. 5в; критерий Рэлея; обезьяна N: z = 8,33, p < 10–3). Mhedzisiro iyi haigone kutsanangurwa nekuchinja kwekugoverwa kwemaitiro akasarudzika mumayuniti mune imwe neimwe sampu yakananga kune imwe yecheckered mirayiridzo kana zvikamu zvechikamu (Fig. 5c; Rayleigh test; monkey N: z = 8.33, p <10-3)., denderedzwa kureva = 190.13 madhigirii ± 9.83 madhigirii;tsoko S: z = 0.79, p = 0.45) uye yakanga yakafanana kumakona ose egridi yeplaid (Supplementary Figure 2).7猴子N:z = 8.33，p <10-3 ，圆形平均值= 190.13 度上± 9.83 度；子S:z = 0.79，p = 0.45图2）.這 种 效应 不 能 通过 每 樣本 中 单元 中 优选 方向 分布 偏向 偏向 图案 元 戻 光 偏向 图案 元 元 成 戻 成 布图 图 瑞利 测试 ； 猴子 n : z = 8.33 , p <10-3 , 平均值 平均值 圆形 圆形 圆墢 圆彆 彆彆 彆彆 彆彆 彆彆 彆彆 彆彆 彆彆 彆彆 彆 彆彆 彆 彆 彆彆 彆 彆 彆 彆 彆 彆 彆圆形 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). Mhedzisiro iyi haigone kutsanangurwa nenyaya yekuti kugoverwa kwemaitiro akasarudzika mumasero mune imwe neimwe sampu inorerekera kune yakanangana neretiti chimiro kana kune chimwe chechikamu chechikamu (Fig. 5c; Rayleigh's test; monkey N: z = 8.33, p <10-3)., avhareji yedenderedzwa) = 190.13 madhigirii ± 9.83 madhigirii;tsoko S: z = 0.79, p = 0.45) uye vakanga vakaenzana mumakona eresiti pakati pegridi (Supplementary Fig. 2).Nekudaro, kunzwisiswa kwemaneuroni kune akagadzirwa grids kunoenderana, zvishoma muchikamu, pane zvakakosha zveMT tuning.
Chikamu chekuruboshwe chinoratidza kugoverwa kweN / P ratios (neuron / psychophysiological threshold);sero rimwe nerimwe rinopa maviri data mapoinzi, imwe kune yega yega nzira iyo pateni inofamba.Iyo yekurudyi pani inoronga psychophysical zvikumbaridzo (kuronga) kupesana neuronal zvikumbaridzo (abscissa) kune ese mayuniti mumuenzaniso.Iwo data mumutsara wepamusoro (pazasi) anobva kutsoko N (S).b Normalized threshold ratios inorongwa maererano nehukuru hwemutsauko pakati peiyo optimal lattice orientation uye inosarudzwa sero orientation.Iyo "yakanakisa" kutungamira inotsanangurwa sekutungamira kweiyo grating chimiro (yakayerwa neine imwe sinusoidal grating) iri padyo neinodiwa sero kutungamira.Dhata yakatanga kuvharirwa neyakajairwa yakasarudzika yakasarudzika (10 ° mabhini), ipapo zviyero zvechikumbaridzo zvakajairwa kune iyo yakanyanya kukosha uye yakaenzana mukati mebhini rega rega.Masero ane chinongedzo chaanoda akakura zvishoma kana kuti madiki pane kutariswa kwezvikamu zvelatisi aive nemusiyano mukuru mukunzwisiswa kune kutaridzika kwepateni yelatisi.c Pink histogram yeanofarira yekutarisa kugovera kwese MT zvikamu zvakarekodhwa mutsoko yega yega.
Pakupedzisira, mhinduro yeMT inogadziriswa nehutungamiri hwekufamba kwegrating uye zvinyorwa zvezvikamu zvedu zvezvikamu (zvinyorwa).Kuenzanisa neuronal uye psychophysical senitivity kwakaratidza kuti, kazhinji, MT mauniti aive asinganyanyi kutarisisa kusiyanisa masaini masaini kupfuura tsoko.Nekudaro, kunzwisiswa kweiyo neuron kwakachinja zvichienderana nemusiyano uripo pakati pekuda kutaridzika kweyuniti uye kutungamira kwegridi kufamba.Masero akanyanya kutarisisa aiwanzova neatarisiro aifarira anenge akavhara retisi patani kana imwe yenzvimbo, uye chikamu chidiki chemasamples edu chaive chakapfava kana kunzwa zvakanyanya kupfuura maonero etsoko dzekusiyana.Kuti tione kana masaini kubva kune aya akanyanya kujeka mayuniti ainyanya kubatanidzwa nekuona mutsoko, takaongorora kuwirirana pakati pemaonero uye neuronal mhinduro.
Nhanho yakakosha pakugadzira hukama pakati pezviitwa neural uye maitiro ndeyekumisa kuwirirana pakati peeuroni uye maitiro emhinduro kune zvinogara zvichikurudzira.Kuti ubatanidze mhinduro dzeneural kune segmentation kutonga, zvakakosha kugadzira kukurudzira iyo, kunyangwe yakafanana, inoonekwa zvakasiyana mumiedzo yakasiyana.Muchidzidzo chazvino, izvi zvinomiririrwa zvakajeka ne zero texture musiyano grating.Kunyangwe isu tichisimbisa kuti, zvichibva pamabasa e psychometric emhuka, magirating ane mashoma (asingasviki ~ 20%) musiyano wemavara anowanzoonekwa seanowirirana kana akajeka.
Kuti tienzanise kuti mhinduro dzeMT dzinopindirana sei nemishumo yekunzwisisa, takaita sarudzo yekusarudza (CP) kuongororwa kwedata redu regidhi (ona 3).Muchidimbu, CP isiri-parametric, isiri-yakajairwa chiyero chinotaridza hukama pakati pemhinduro dzespike uye maonero ekutonga30.Kuganhurira kuongororwa kumiyedzo tichishandisa magridi ane zero mavara akasiyana uye zvikamu umo tsoko dzakaita kanenge kashanu sarudzo dzerudzi rwega rwemiedzo iyi, isu takaverenga SR zvakasiyana kune yega yega nzira yekufamba kwegridi. Pakati petsoko, takaona kukosha kweCP kwakakura zvakanyanya kupfuura zvatingatarisira netsaona (Fig. 6a, d; tsoko N: zvinoreva CP: 0.54, 95% CI: (0.53, 0.56), t-test yemativi maviri kurwisa null yeCP = 0.5, t = 6.0 mon : 9, p. 5% CI: (0.54, 0.57), mativi maviri-t-test, t = 9.4, p <10-13). Pakati petsoko, takaona kukosha kweCP kwakakura zvakanyanya kupfuura zvatingatarisira netsaona (Fig. 6a, d; tsoko N: zvinoreva CP: 0.54, 95% CI: (0.53, 0.56), t-test yemativi maviri kurwisa null yeCP = 0.5, t = 6.0 mon : 9, p. 5% CI: (0.54, 0.57), mativi maviri-t-test, t = 9.4, p <10-13).Mutsoko, takaona CP inorehwa yakakwira zvakanyanya kupfuura yaitarisirwa (Fig. 6a, d; tsoko N: zvinoreva CP: 0.54, 95% CI: (0.53, 0.56), maviri-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) . tsoko S: zvinoreva CP: 0.55, 95% CI: (0.54, 0.57), maviri-tailed t-test, 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:3,54,95% 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; 猴子S: 平均CP: 0.55, 95% CI: (0.54, 0.57), 双边t <10−9;在 猴子 中 ， 我們 观察 平均 平均 值 显着 大于 我們 偶然 的 值 图 图 6a , d: : ; 5 : 5 , 5 , 5 .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, d: 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% критерий, t = 9,4, p <10- 13) . Mutsoko, takaona zvinoreva kukosha kweCP zviri pamusoro pezvatingatarisira netsaona (Fig. 6a, d; tsoko N: zvinoreva CP: 0.54, 95% CI: (0.53, 0.56), maviri-tailed t-test CP vs. zero = 0.5, t = CP mon10: 5, % 5. CI: (0.54, 0.57), maviri-tailed t- criterion, t = 9.4, p <10-13).Saka, MT neurons anowanzopisa zvakanyanya kunyangwe pasina chero akajeka segmentation cues, kana maonerwo emhuka ekufamba kwelattice achienderana nezvido zvesero.
a Selection probability distribution for grids without texture signs for samples recorded from monkey N. Sero rega rega rinogona kubatsira kusvika pamapoinzi maviri edata (rimwe kune rimwe divi rekufamba kwegridi).Chirevo cheCP kukosha pamusoro pechisina (chichena miseve) chinoratidza kuti pane hukama hwakakosha pakati pekuita kweMT uye maonero.b Kuti tiongorore kukanganisa kwechero sarudzo yekusarura, isu takaverenga CP zvakasiyana kune chero kukurudzira uko tsoko dzakaita kamwechete kukanganisa.Iwo mukana wekusarudza wakarongerwa sebasa reyero yekusarudza (pref/null) yezvese zvinokurudzira (kuruboshwe) uye mhedziso kukosha kwemavara maki musiyano (kurudyi, data kubva ku120 masero ega ega).Mutsara wakasimba uye nzvimbo ine shaded mupaneru yekuruboshwe inomiririra zvinoreva ± sem ye20-point inofamba pakati.Sarudzo ingangoita yakaverengerwa kurudziro ine nhanho dzesarudzo dzisina kuenzana, senge magidhi ane misiyano yechiratidzo chepamusoro, aisiyana zvakanyanya uye akaunganidzwa zvakatenderedza zvingangoitika.Iyo grey-shaded nzvimbo mupaneru yekurudyi inosimbisa kusiyana kwezvinhu zvakabatanidzwa mukuverenga kwepamusoro kusarudzwa kwepamusoro.c Mukana wesarudzo huru (ordination) inorongerwa pachikumbaridzo cheneuron (abscissa).Mukana wekusarudzwa waive wakabatana zvakanyanya nehuremu.Gungano df rakafanana neac asi rinoshanda ku157 data rimwechete kubva kutsoko S kunze kwekunge zvaonekwa.g Mukana wepamusoro wekusarudza (kuronga) wakarongerwa kutungamira kwakasarudzika (abscissa) kune imwe neimwe yetsoko mbiri.Sero rega rega reMT rakapa mapoinzi maviri edata (imwe kune imwe neimwe nzira yechimiro chelattice).h Mahombe mabhokisi ezvirongwa zvekusarudza mukana kune yega yega inter-raster angle.Mutsara wakasimba unoisa mitsetse yepakati, yepasi nekumusoro mipendero yebhokisi inomiririra 25th uye 75th percentiles zvichiteerana, ndebvu dzinowedzerwa kusvika 1.5 times the interquartile range, uye kunze kweiyi muganhu zvinocherechedzwa.Iyo data iri kuruboshwe (kurudyi) mapaneru anobva ku120 (157) yega N(S) maseru etsoko.i Mukana wepamusoro wekusarudza (kuronga) wakarongerwa nguva yekutanga kwekukurudzira (abscissa).Huru CP yakaverengerwa mumarectangles anotsvedza (hupamhi 100 ms, nhanho 10 ms) muyedzo yese uyezve avhareji pamusoro pemayuniti.
Zvimwe zvidzidzo zvekare zvakashuma kuti CP inoenderana nehuwandu hwemiyedzo muyedzo yebasal kugovera, zvichireva kuti chiyero ichi hachina kuvimbika kune zvinokurudzira zvinokonzeresa misiyano mikuru muchikamu chesarudzo yega yega.Kuti tiedze mhedzisiro iyi mune yedu data, isu takaverenga iyo CP zvakasiyana kune zvese zvinokurudzira, zvisinei nechiratidzo chekusiyanisa, uye tsoko dzakaita kamwechete nhema kuyedza.Iyo CP inorongerwa kutarisana nechiyero chekusarudza (pref / null) yemhuka yega yega muMufananidzo 6b uye e (kuruboshwe panhi), zvichiteerana.Tichitarisa paavhareji inofamba, zviri pachena kuti CP inoramba iri pamusoro pezvingabvira pamusoro pemhando dzakasiyana dzesarudzo, ichidzikira chete kana kusawirirana kuchidonha (kuwedzera) pazasi (pamusoro) 0.2 (0.8).Kubva pane psychometric maitiro emhuka, isu taizotarisira kusarudzwa coefficients ehukuru uhu kuti hushandiswe chete kune zvinokurudzira nepamusoro-kusiyana kwemavara mavara (akabatana kana akajeka) (ona mienzaniso yezvimiro zvepfungwa muFig. 2a, b).Kuti tione kana izvi zvaive zvakadaro uye kana PC yakakosha yakaramba ichikurudzira nezviratidzo zvezvikamu zvakajeka, takaongorora mhedzisiro yeabsolute textural mutsauko kukosha paPC (Fig. 6b, e-kurudyi).Sezvaitarisirwa, CP yaive yakakwira zvakanyanya kupfuura mukana wekukurudzira kunosvika pakati nepakati (~ 20% kusiyanisa kana kudzika) segmentation cues.
Mukutarisa, kumhanya, uye kusapindirana kwekuziva mabasa, MT CP inowanzove yepamusoro-soro mumaneuron anonyanya kuoma, zvichida nekuti idzi neuroni dzinotakura zviratidzo zvinodzidzisa30,32,33,34.Zvichienderana nezvakawanikwa izvi takaona hukama hune mwero asi hwakakosha pakati pehukuru CP, yakaverengerwa kubva ku-z-yakapimwa mareti ekupfura mukati meiyo mameseji cue misiyano yakasimbiswa mune yekurudyi pani yeFig.6b, e, uye neuronal threshold (Fig. 6c, f; geometric mean regression; monkey N: r = -0.12, p = 0.07 monkey S: r = -0.18, p <10-3). 6b, e, uye neuronal threshold (Fig. 6c, f; geometric mean regression; monkey N: r = -0.12, p = 0.07 monkey S: r = -0.18, p <10-3).Zvichienderana nezvakawanikwa izvi, takaona kuwirirana kwakadzikama asi kwakakosha pakati peCP yakakura yakaverengerwa kubva pamhepo yekukurudzira z-score kubva kune yemavara maratidziro akasiyana anoratidzwa muchikamu chekurudyi cheFig. 6b, e, uye neuronal threshold (Fig. 6c, f; geometric).geometric zvinoreva kuderera;обезьяна N: r = -0,12, p = 0,07 обезьяна S: r = -0,18, p < 10-3). tsoko N: r = -0.12, p = 0.07 tsoko 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 和神经元阈倚回归；猴子N：r = -0.12，p = 0.07 猴子S：r = -0.18，p <10-3）。与 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 6b , e . , f ; 回归 ； 猴子 n : r = -0.12 , p = 0.07 猴子S：r = -0.18，p <10-3）.Zvichienderana nezvakawanikwa izvi, takaona kuwirirana kwakadzikama asi kwakakosha pakati peCVs huru sezvinoratidzwa muFig. 6b, e uye neuron thresholds (Mufananidzo 6c, f; geometric zvinoreva kuregeredza; monkey N: r = -0.12, p = 0.07).Обезьяна S: г = -0,18, р < 10-3). Tsoko S: r = -0.18, p <10-3).Naizvozvo, zvinyorwa kubva kune zvakanyanya ruzivo mayuniti zvairatidza hukuru covariance ne subjective segmentation mitongo mutsoko, izvo zvakakosha zvisinei nechero mavara ezvinyorwa akawedzerwa kune yekufunga.
Tichifunga kuti isu takambogadzira hukama pakati pekunzwa kune grid masaini masaini uye aifarira neuronal orientation, takashamisika kana paine hukama hwakafanana pakati peCP uye yaidiwa kutarisisa (Fig. 6g).Kubatana uku kwaingokosha zvishoma mutsoko S (ANOVA; tsoko N: 1.03, p=0.46; tsoko S: F=1.73, p=0.04).Hatina kucherechedza mutsauko muCP yemakona e lattice pakati pemachira mune chero mhuka (Fig. 6h; ANOVA; monkey N: F = 1.8, p = 0.11; monkey S: F = 0.32, p = 0. 9).
Pakupedzisira, basa rekare rakaratidza kuti CP inoshanduka munguva yose yekuedza.Zvimwe zvidzidzo zvakashuma kuwedzera kwakanyanya kunoteverwa nesarudzo yakanyatsonaka, 30 apo vamwe vakashuma kuwedzera kwakasimba kwechiratidzo chekusarudza pamusoro penguva yekuedzwa31.Patsoko yega yega, takaverenga CP yeyuniti yega yega mumiyedzo ine zero textural musiyano (zvichitevedzana, zvinoenderana nemaitiro ekumisikidza) mumaseru 100 ms achikwira yega 20 ms kubva pre-stimulus kutanga kutumira zvinoreva pre-stimulus offset.Avhareji yeCP dynamics yetsoko mbiri inoratidzwa muFig. 6i.Muzviitiko zvese izvi, CP yakaramba iri padanho risingarondedzereki kana padyo padyo kusvika panosvika 500 ms mushure mekutanga kwekukurudzira, mushure meizvozvo CP yakawedzera zvakanyanya.
Pamusoro pekuchinja kunzwisiswa, CP yakaratidza kukanganiswawo nehumwe hunhu hwesero tuning hunhu.Semuyenzaniso, Uka naDeAngelis34 vakawana kuti CP mubinocular mismatch recognition basa rinoenderana nesymmetry yebiinocular mismatch tuning curve yemuchina.Muchiitiko ichi, mubvunzo une hukama ndewekuti masero epattern direction selective (PDS) anotarisisa here pane masero anosarudza (CDS).Masero ePDS encode kutaridzika kwemaitiro ane maitiro akawanda emunharaunda, nepo masero eCDS anopindura kufamba kwemaitiro ekutungamirira maitiro (Fig. 7a).
a Schematic inomiririra yemodhi yechikamu tuning kukurudzira uye yekufungidzira grating (kuruboshwe) uye grating yekumisikidza tuning macurves (kurudyi) (ona Zvishandiso uye Nzira).Muchidimbu, kana sero rakabatana nepakati pegidhi zvikamu kuratidza mafambiro epateni, munhu angatarisira iwo matuning macurves ega ega grid uye grid stimuli (yekupedzisira koramu, yakasimba curve).Zvikasadaro, kana sero rikasabatanidza mafambiro ezvikamu mukufamba kwechiratidzo chechiratidzo, munhu angatarisira bipartite tuning curve ine peak munzira yega yega yekugezesa inofamba inoshandura chikamu chimwe kuenda kudivi rinodiwa nesero (koramu yekupedzisira, dashed curve)..b (kuruboshwe) mavhavha ekugadzirisa kutaridzika kwe sinusoidal array yemasero anoratidzwa muMifananidzo 1 uye 2. 3 uye 4 (mutsara wepamusoro - masero muFig. 3a, b uye 4a, b (kumusoro); pasi pasi - masero eFig. 3c, d uye 4a, b (pasi)).(Pakati) Patani uye fungidziro yechikamu yakaverengerwa kubva kune lattice tuning profiles.(Kurudyi) Kugadzirisa grid yemaseru aya.Iwo maseru ekumusoro (pazasi) mapaneru anoiswa se template (chikamu) maseru.Cherechedza kuti hapana-kune-mumwe kunyorerana pakati pezvikamu zvezvikamu zvepateni uye zvido zvekubatana / kujeka kwesero kufamba (ona mhinduro dzelatiti yekugadzirwa kwemasero aya mumufananidzo 4a).c Coefficient of partial correlation of the z-score mode (ordinate) akarongedzerwa nechikamu chekubatana kwechikamu chez-score component (abscissa) yemasero ose akanyorwa muN (kuruboshwe) uye S (kurudyi) tsoko.Mitsetse mikobvu inoratidza kukosha kwemaitiro anoshandiswa kuronga maseru.d Chirongwa chepamusoro sarudzo mukana (kuronga) maringe nemodhi index (Zp - Zc) (abscissa).Data iri kuruboshwe (kurudyi) mapaneru anoreva tsoko N(S).Denderedzwa dema rinoratidza data mumayuniti akaenzana.Mumhuka dzose dziri mbiri, pakanga paine kuwirirana kwakakosha pakati pekugona kusarudzwa kwepamusoro uye pateni index, zvichiratidza kuwirirana kukuru kwemaseru ane chiratidzo cheratidziro yekumisikidza mune inosimudzira ine yakawanda chikamu chekutarisa.
Naizvozvo, mune imwe bvunzo seti, takayera mhinduro kune sinusoidal grids uye grids kurongedza neuroni mumasampuli edu sePDS kana CDS (ona Nzira).Lattice tuning curves, template component predictions yakavakwa kubva pane iyi tuning data, uye lattice tuning curves yemasero anoratidzwa muFigure 1 ne3. Figure 3 uye 4 uye Supplementary Figure 3 inoratidzwa muFigure 7b.Iko kugoverwa kwepateni uye kusarudzwa kwechikamu, pamwe chete neinodiwa sero kutaridzika muchikamu chimwe nechimwe, inoratidzwa kune imwe neimwe tsoko muFig. 7c uye yekuwedzera fig.4 zvichiteerana.
Kuongorora kutsamira kweCP pakugadziriswa kwezvikamu zvepateni, takatanga kuverenga iyo index index 35 (PI), iyo yakakura (idiki) hunhu hunoratidza yakakura PDS (CDS) maitiro akafanana.Tichipa kuratidzwa kwepamusoro kuti: (i) neuronal senitivity inosiyana nemusiyano pakati peyakasarudzwa sero orientation uye kutungamira kwekukurudzira kufamba, uye (ii) pane kuwirirana kwakakosha pakati pekunzwa neuronal uye mukana wekusarudza mumuenzaniso wedu, takawana hukama pakati pePI uye iyo yakazara CP yakadzidzwa kune "yakanakisa" kutungamira kwekufamba kwesero rega rega (ona pamusoro). Takaona kuti CP yakanga yakabatana zvikuru nePI (Fig. 7d; geometric mean regression; grand CP tsoko N: r = 0.23, p <0.01; bi-stable CP monkey N r = 0.21, p = 0.013; grand CP tsoko S: r = 4 , CP monkey S: r = 4 , 1 − CP 0.29, p <10−3), zvichiratidza kuti masero akarongedzerwa sePDS airatidza basa rakakura rine chekuita nesarudzo kupfuura maCDS nemaseru asina kunyorwa. Takaona kuti CP yakanga yakabatana zvikuru nePI (Fig. 7d; geometric mean regression; grand CP tsoko N: r = 0.23, p <0.01; bi-stable CP monkey N r = 0.21, p = 0.013; grand CP tsoko S: r = 4 , CP monkey S: r = 4 , 1 − CP 0.29, p <10−3), zvichiratidza kuti masero akarongedzerwa sePDS airatidza basa rakakura rine chekuita nesarudzo kupfuura maCDS nemaseru asina kunyorwa. Мы обнаружили, что CP значительно коррелирует с PI (рис. 7d; регрессия среднего геометрического; большая обезьбяна CP N:3, r0, p = 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 = 0,10, ут 0,29, ут клетки, классифицированные как PDS, проявляли большую активность, связанную с выбором, чем CDS и неклассифицированные клетки. Takaona kuti CP yaive yakabatana zvakanyanya nePI (Mufananidzo 7d; geometric zvinoreva kuregeredza; tsoko huru CP N: r = 0.23, p <0.01; bistable tsoko CP N r = 0.21, p = 0.013; tsoko hombe CP S: r = 0.10 S: r = 0.10 s. 9.我們发现CP 与PI 显着相关（图7d; 大CP 猴N:r = 0.23，p <0.01；双稳态CP = 猴0 CP = 3 0: CP = 3 0. 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 和未分类的细类的细类的细类的细类 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 ; Мы обнаружили, что CP был значительно связан с PI (рис. 7d; регрессия среднего геометрического; большая обезьбяна CP N:3, r0, p = 0,21; зьяна CP N r = 0,21, p = 0,013; большая обезьяна CP S: r = 0,013) 0,30, p <10-4; Takaona kuti CP yakabatanidzwa zvakanyanya nePI (Mufananidzo 7d; geometric zvinoreva kuregeredza; tsoko huru CP N: r = 0.23, p <0.01; bistable tsoko CP N r = 0.21, p = 0.013; tsoko huru 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, прояолунцевлеклиши ть, чем клетки, классифицированные как CDS и неклассифицированные. monkey S bistable CP: r = 0.29, p <10-3), zvichiratidza kuti masero akarongwa sePDS airatidza basa guru rekusarudza kupfuura masero anotsanangurwa seCDS uye asina kunyorwa.Nekuti zvese PI uye neuron senitivity yakabatana neCP, takaita akawanda regression ongororo (nePI uye neuron senitivity sezvizvimiririra zvinosiyana uye huru CP seinotsamira shanduko) kutonga kunze kuwirirana pakati pezviyero zviviri zvinotungamira kune mukana wekuita.. Ose ari maviri chidimbu chekubatanidza coefficient aive akakosha (tsoko N: chikumbaridzo ichipikisana neCP: r = -0.13, p = 0.04, PI vs. CP: r = 0.23, p <0.01; tsoko S: chikumbaridzo vs. CP: r = −0.0 PI. −3), zvichiratidza kuti CP inowedzera nekunzwa uye nenzira yakazvimirira inowedzera nePI. Ose ari maviri chidimbu chekubatanidza coefficient aive akakosha (tsoko N: chikumbaridzo ichipikisana neCP: r = -0.13, p = 0.04, PI vs. CP: r = 0.23, p <0.01; tsoko S: chikumbaridzo vs. CP: r = −0.0 PI. −3), zvichiratidza kuti CP inowedzera nekunzwa uye nenzira yakazvimirira inowedzera nePI. Оба частных коэффициента корреляции были значимыми (обезьяна N: порог против CP: r = -0,13, p = 0,04, PI против CP: 3, p = 0, p = 0, p = 0, про ротив CP: r = -0,16, p = 0,03, PI vs CP: 0,29, p < 10-3), предполагая, что CP увеличивается с чувствительностью и независимзечи обется в PI. Ose ari maviri chidimbu coefficients akakosha (tsoko N: chikumbaridzo ichipikisana neCP: r=-0.13, p=0.04, PI vs. CP: r=0.23, p<0.01; tsoko S: chikumbaridzo ichipikisana neCP: r = -0.16, p = 0.03 , CP- 16, p = 0.03, p nekunzwa uye inowedzera yakazvimiririra nePI.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:阈值与CP:r = -0.13,p = 0.04,PI 与CP:r = 0.23，p <0.01: 阈值值 = S. ,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漉， Об б б б о О ч тив CP: R = -0,16, p = 0,03, Pi ♥ 0,29, p <10-3), что CP увеливиливи Ом. Ose ari maviri epakati coefficients akakosha (tsoko N: chikumbaridzo ichipikisana neCP: r=-0.13, p=0.04, PI vs. CP: r=0.23, p<0.01; tsoko S: chikumbaridzo ichipikisana. CP: r = -0.16, p = 0.03 PIdic mu 19, p yakawedzera nekunzwa uye yakawedzera nePI nenzira yakazvimirira.
Takanyora chiitiko chimwe chete munharaunda yeMT, uye tsoko dzakashuma maonero adzo emaitiro ayo anogona kuratidzika seanowirirana kana mafambiro akajeka.Kunzwa kwemaneuroni kumasegmentation cues akawedzerwa kune akarerekera maonero anosiyana zvakanyanya uye anotemerwa, kanenge muchidimbu, nehukama pakati peiyo unit inofarirwa kutaridzika uye kutungamira kwekufambisa kwekusimudzira.Muhuwandu hwese, neuronal senitivity yaive yakaderera zvakanyanya pane psychophysical senitivity, kunyangwe iwo akanyanya kuomarara mauniti akafananidzwa kana kudarika maitiro ekunzwa kune zvikamu zvezviratidzo.Uye zvakare, pane yakakosha covariance pakati pekupfura frequency uye maonero, zvichiratidza kuti kusaina MT kunotora chikamu muzvikamu.Masero ane maitiro aanofarira akakwenenzvera kunzwisiswa kwawo kune misiyano mumasaini echikamu chelatisi uye aiwanzoratidza kufamba kwepasirese mukukurudzira nekwakawanda kwenzvimbo, zvichiratidza kuwirirana kwepamusoro-soro.Pano tinoona mamwe matambudziko anogona kuitika tisati taenzanisa izvi zvabuda nebasa rekare.
Dambudziko guru nekutsvaga uchishandisa bistable stimuli mumhando dzemhuka nderekuti maitiro emhinduro anogona kunge asina kubva pachiyero chekufarira.Semuyenzaniso, tsoko dzedu dzinokwanisa kutaura maonere adzo ezvekumisikidzwa kwemavara pasina maonero adzo ekubatana kweretisi.Zvikamu zviviri zve data zvinoratidza kuti izvi handizvo.Chekutanga, maringe nemishumo yapfuura, kushandura nhanho yekutarisisa yezvikamu zvakapatsanurwa zvakarongeka zvakashandura mukana wekuona kwakabatana.Chechipiri, paavhareji, mhedzisiro yacho yakafanana kune mapatani ane kana asina masaini masaini.Zvakatorwa pamwechete, zvakaonekwa izvi zvinoratidza kuti mhinduro dzetsoko dzinogara dzichiratidza maonero adzo ekubatana/kubudirana pachena.
Rimwe dambudziko rinogona kuitika nderekuti isu hatina kukwidziridza maparamita ekufamba kweiyo mamiriro chaiwo.Mumabasa akawanda apfuura achienzanisa neuronal uye psychophysical sensitivity, zvinokurudzira zvakasarudzwa mumwe nomumwe kune imwe neimwe yakanyoreswa unit [31, 32, 34, 36, 37, 38, 39, 40, 41, 42, 43, 44, 45].Pano tashandisa nzira mbiri dzakafanana dzekufamba kweretiti patani zvisinei nekugadziriswa kwekutarisa kwesero rega rega.Iyi dhizaini yakatibvumira kuti tidzidze kuti kunzwisiswa kwakashanduka sei nekupindirana pakati pekufamba kwelatisi uye kutaridzika kwaanofarira, zvisinei, hazvina kupa hwaro hwekutanga hwekuona kana masero anofarira akabatana kana akajeka lattices.Naizvozvo, isu tinovimba nemaitiro eempirical, tichishandisa mhinduro yesero kune yakashongedzwa mesh, kugovera zvaunofarira uye zero label kuchikamu chega chega chekufamba mesh.Kunyangwe zvisingaite, izvi zvinogona kukanganisa zvakarongeka mhedzisiro yekuongorora kwedu kwekunzwa uye kuonekwa kwechiratidzo cheCP, zvichigona kuwedzeredza chero chiyero.Zvisinei, zvimwe zvikamu zvekuongorora uye data inokurukurwa pasi apa inoratidza kuti izvi hazvisizvo.
Chekutanga, kugovera mazita akasarudzika (asina maturo) kune zvinokurudzira izvo zvakaita kuti zvimwe (zvishoma) zviitwe hazvina kukanganisa kusiyanisa kwekugoverwa kwemhinduro idzi.Pane kudaro, inongovimbisa kuti iyo neurometric uye psychometric mabasa ane chiratidzo chimwe chete, saka anogona kuenzaniswa zvakananga.Chechipiri, mhinduro dzakashandiswa kuverenga CP (iyo "yakashata" miedzo yezvinyorwa zvinyorwa uye miedzo yose ye gratings isina mavara akasiyana) yakanga isingabatanidzwi mukudzokororwa kwekuongorora kwakaratidza kana sero rimwe nerimwe "rinoda" mitambo yakabatana kana yakajeka.Izvi zvinova nechokwadi chekuti sarudzo hadzina kurerekera kune yakasarudzika / isina kufanira mazita, zvichikonzera yakakosha sarudzo.
Zvidzidzo zveNewsom nevamwe vake [36, 39, 46, 47] ndivo vakatanga kuona basa reMT mukufungidzira kunofungidzirwa kwekutungamira kwekufamba.Mishumo yakatevera yakaunganidza dhiyabhorosi paMT kubatanidzwa mukudzika34,44,48,49,50,51 uye speed32,52, yakanaka orientation33 uye maonero e3D chimiro kubva kufamba31,53,54 (3D masango akasimba).kutonga).Tinowedzera mhedzisiro iyi nenzira mbiri dzakakosha.Chekutanga, isu tinopa humbowo hwekuti mhinduro dzeMT dzinobatsira mukunzwisisa chikamu chezviratidzo zve visuomotor.Chechipiri, takaona hukama pakati peMT mode orientation selectivity uye iyi chiratidzo chekusarudza.
Sezvineiwo, mhedzisiro iripo yakanyanya kufanana nebasa re3-D SFM, sezvo ese ari maviri akaomesesa bistable maonero anosanganisira kufamba uye kudzika kurongeka.Dodd et al.31 vakawana mukana wakakura wekusarudza (0.67) mubasa retsoko richirondedzera kutenderera kutenderera kwebistable 3D SFM silinda.Isu takawana diki diki sarudzo yekuita bistable grid stimuli (inenge 0.55 yetsoko dzese).Sezvo kuongororwa kweCP kunoenderana nesarudzo yekusarudza, zvakaoma kududzira CP yakawanikwa pasi pemamiriro akasiyana mumabasa akasiyana.Zvisinei, ukuru hwesarudzo yatakaona yakanga yakafanana ne zero uye yakaderera texture kusiyana gratings, uyewo pataibatanidza yakaderera/pasina manyorerwo musiyano stimuli kuwedzera simba.Naizvozvo, musiyano uyu muCP haugone kuve nekuda kwekusiyana kwemitengo yekusarudza pakati pe dataset.
Shanduko dzine mwero muMT yekupfura mwero inoperekedza maonero mune yekupedzisira nyaya inoita seinokatyamadza kana ichienzaniswa neyakaomesesa uye yemhando dzakasiyana mamiriro ekufunga anokonzerwa ne3-D SFM kukurudzira uye bistable grid zvimiro.Imwe mukana ndeyekuti isu takarerutsa maitiro ekusarudza nekuverenga mwero wekupfura pamusoro penguva yese yekukurudzira.Kusiyana nenyaya ye31 3-D SFM, apo kusiyana kweMT basa rakagadzirwa zvakapoteredza 250 ms mumiedzo uye zvakabva zvawedzera kuwedzera mukati memiedzo yose, kuongorora kwedu kwemaitiro echinyakare ezviratidzo zvekusarudza (ona 500 ms mushure mekukurudzira kunotanga mutsoko mbiri. Mukuwedzera , mushure mekukwira kwakanyanya munguva iyi mushumo muhupfenyuro muhupfenyuro hwe5 uye Rubin mushumo weRubin panguva iyi, isu takacherechedza 500 ms mushure mekukurudzira. Kuona kwebistable rectangular arrays kazhinji kunochinja panguva yekuedza kwenguva refu.Kunyange zvazvo kukurudzira kwedu kwakaunzwa kwemasekonzi 1.5 chete, maonero etsoko dzedu anogonawo kusiyana kubva pakubatana kusvika pakujeka panguva yekutongwa (mhinduro dzadzo dzakaratidza maonero avo ekupedzisira pakusarudzwa kwe cue.) Naizvozvo, vhezheni yenguva yekupindura yebasa redu, kana hurongwa umo tsoko dzinogona kuve nemhedzisiro inotarisirwa kuenderera mberi. verenga zvakasiyana mumabasa maviri.Kunyange zvazvo zvagara zvichifungidzirwa kuti zviratidzo zveCPU zvinokonzerwa nekugadzirisa kwekunzwa uye ruzha rwakabatanidzwa,56 Gu nevamwe vaaishanda navo57 vakawana kuti mumakomputa emhando, nzira dzakasiyana dzekubatanidza, pane mazinga ekusiyana kwakabatana, zvinogona kutsanangura zviri nani CPU mudorsal media-superior temporal neurons.sheet shanduko yekutarisa kuzivikanwa basa (MSTd).Kusarudzwa kuduku kwatakaona muMT zvichida kunoratidza kuunganidzwa kwakakura kwemaneuron akawanda asina ruzivo kuti agadzire maonero ekubatana kana pachena.Chero mamiriro ezvinhu apo maitiro ekufambisa emunharaunda aifanira kuiswa muchinhu chimwe kana zviviri (bistable gratings) kana nzvimbo dzakasiyana dzezvinhu zvakajairika (3-D SFM), uchapupu hwakazvimirira hwekuti mhinduro dzeMT dzaive dzakabatanidzwa zvakanyanya nekutonga kwekunzwisisa, pakanga pane mhinduro dzakasimba dzeMT.inorongedzerwa kuita basa mukukamurwa kwemifananidzo yakaoma muzvikamu zvezvinhu zvakawanda uchishandisa ruzivo rwekufamba kwekuona.
Sezvambotaurwa pamusoro apa, isu taiva vekutanga kuzivisa kushamwaridzana pakati peMT pateni cellular chiitiko uye maonero.Sezvakagadzirwa mune yekutanga-matanho maviri modhi naMovshon uye vaanoshanda navo, iyo mode unit ndiyo yekubuda nhanho yeMT.Nekudaro, basa razvino rakaratidza kuti modhi uye masero echikamu anomiririra magumo akasiyana ekuenderera uye kuti parametric misiyano muchimiro chendima yekugamuchira inokonzeresa iyo tuning spectrum yezvikamu zvemodhi.Naizvozvo, takawana kuwirirana kwakakosha pakati peCP nePI, yakafanana nehukama pakati pebinocular mismatch adjustment symmetry uye CP mune yakadzama yekuziva basa kana orientation yekumisikidza gadziriso mune yakanaka orientation kusarura basa.Hukama pakati pezvinyorwa uye CP 33.Wang naMovshon62 vakaongorora nhamba huru yemasero ane MT orientation selectivity uye akawana kuti, paavhareji, iyo mode index yakabatanidzwa nehuwandu hwekugadzirisa zvinhu, zvichiratidza kuti sarudzo yemodhi iripo mune mamwe marudzi akawanda ezviratidzo zvinogona kuverengwa kubva kuvanhu veMT..Nokudaro, kune zvidzidzo zvenguva yemberi zvehukama pakati pebasa reMT uye maonero ekuona, zvichave zvakakosha kuti uone kana iyo index index inopindirana zvakafanana nemamwe mabasa uye zviratidzo zvekusarudza zvinokurudzira, kana kuti hukama uhwu hwakananga kune imwe nyaya yekuparadzanisa kwemaonero.
Saizvozvo, Nienborg naCumming 42 vakawana kuti kunyangwe maseru ari pedyo uye kure anosarudza kusawirirana kwebinocular muV2 aive akaenzana mukusarura kwakadzama basa, chete masero ari pedyo-anofarira airatidza yakakosha CP.Nekudaro, kudzidzisazve tsoko kuti dziite uremu kure nekusiyana kwakakonzera maCP akakosha mumakeji anofarirwa.Zvimwe zvidzidzo zvakashumawo kuti nhoroondo yekudzidzisa inobva pakuona kuwirirana34,40,63 kana hukama hunokonzera pakati pebasa reMT uye kusarura kwakasiyana48.Hukama hwatakacherekedza pakati peCP nerejimendi yekusarudzika ingangoratidza zano rakashandiswa netsoko kugadzirisa dambudziko redu, uye kwete basa chairo remasaini ekusarudza masaini mukuona-motor kuona.Mubasa remunguva yemberi, zvichave zvakakosha kuona kana nhoroondo yekudzidza ine zvakakosha pakuona kuti ndeapi masaini eMT anoyerwa zvakanyanya uye achichinjika kuita segmentation mitongo.
Stoner nevamwe vaaishanda navo14,23 ndivo vaive vekutanga kutaura kuti kushandura kupenya kwenzvimbo dzegridi dzakafanofungidzira kwakakanganisa kubatana uye kubuda pachena kwemishumo yevacherechedzi vevanhu uye kugadzirisa kwegwara mumacaque MT neurons.Vanyori vakaona kuti kana kupenya kwenzvimbo dzakatenderedza muviri kunoenderana nekujeka, vacherechedzi vakashuma maonero akajeka, nepo MT neurons yairatidza kufamba kwezvikamu zve raster.Zvakasiyana, apo kupenya kunopindirana uye kupindirana kwakajeka kusingaenderani mumuviri, mucherechedzi anoona kufamba kwakabatana, uye MT neurons inoratidza kufamba kwepasi rose kwepateni.Nokudaro, zvidzidzo izvi zvinoratidza kuti kuchinja kwemuviri mukukurudzira kwekuona kunopesvedzera zvakavimbika mishumo yezvikamu zvakare inokonzera shanduko dzinofanotaurwa mukumutsa MT.Basa richangoburwa munzvimbo ino rakaongorora kuti ndeapi masaini eMT anoteedzera kutaridzika kwekunaka kwekukurudzira18,24,64.Semuenzaniso, chikamu chidiki cheMT neurons chakaratidzwa kuratidza bimodal tuning kune isina kujairika point inofamba mepu (RDK) ine nzira mbiri dzakapatsanurwa zvishoma pane unidirectional RDK.Bandwidth yeserura tuning 19, 25 .Vacherechedzi vanogara vachiona maitiro ekutanga sekufamba kwakajeka, kunyange zvazvo MT neurons dzakawanda dzichiratidzira unimodal adaptation mukupindura kune izvi zvinokurudzira, uye chiyero chiri nyore chemasero ose eMT anopa unimodal huwandu hwevanhu.Saka, chikamu chemasero anoratidza bimodal tuning chinogona kuumba neural substrate yekuona uku.Sezvineiwo, mumarmosets, huwandu uhu hwakafananidza maseru ePDS paakayedzwa uchishandisa akajairwa grid uye grid stimuli.
Mhedzisiro yedu inoenda nhanho imwe kupfuura iyo iri pamusoro, iyo yakakosha kumisa basa reMT muchikamu chekunzwisisa.Muchokwadi, segmentation chinhu chinozvimiririra.Mazhinji mapolystable ekuona anoratidza kugona kweiyo yekuona system kuronga nekududzira zvinoramba zvichikurudzira munzira dzinopfuura imwe.Panguva imwecheteyo kuunganidza mhinduro dzeneural uye mishumo yekunzwisisa mukudzidza kwedu kwakatibvumira kuti tiongorore covariance pakati peMT kupfura chiyero uye kududzirwa kwekunzwisisa kwekukurudzira nguva dzose.Taratidza hukama uhwu, tinobvuma kuti kutungamirirwa kwechikonzero hachina kusimbiswa, ndiko kuti, zvimwe zviedzo zvinodiwa kuti zvione kana chiratidzo chekunzwisisa chikamu chinocherechedzwa nesu, sekupokana kunoita vamwe [65, 66, 67], otomatiki.Iyo nzira zvakare inomiririra kudzika zviratidzo zvinodzokera kune sensory cortex kubva kunzvimbo dzakakwirira 68, 69, 70 (Fig. 8).Mishumo yechikamu chikuru chemasero-anosarudza masero muMSTd71, chimwe chezvinangwa zvikuru zvekortical yeMT, inoratidza kuti kuwedzera zviedzo izvi kuti zvibatanidze kurekodha panguva imwe chete yeMT neMSTd ingava nhanho yakanaka yekutanga yekuenderera mberi nekunzwisisa nzira dzemaneural dzekuona.segmentation.
Muenzaniso wematanho maviri echikamu uye maitiro ekugadzirisa sarudzo uye zvinogona kuitika zvepamusoro-pasi mhinduro pamusoro pechisarudzo-chine chekuita nebasa mushanduro yemuchina.Pano, modhi inotungamira sarudzo (PDS - "P") muMT nhanho inogadzirwa ne (i) sampuli hombe yekutungamira yakasarudzika data yekuisa inoenderana neyakajeka modhi velocities, uye (ii) yakasimba tuning kudzvanywa.Iyo inotungamirwa inosarudza (CDS) chikamu cheMT ("C") nhanho ine nhete yesampling renji munzira yekupinza uye haina kudzvanyirira kwakawanda kwekugadzirisa.Untuned inhibition inopa kutonga pamusoro pehuwandu hwevanhu.Miseve ine mavara inoratidza yaidiwa mudziyo wakatarisa.Kuti ujekeswe, chete chikamu chidiki cheV1-MT chinongedzo uye chimwe chechikamu modhi uye yekumisikidza sarudzo bhokisi ndiyo inoratidzwa.Muchirevo chekududzira yedu feed-forward (FF) mhedzisiro, yakafararira yekuisa yekuisa uye yakasimba tuning inhibition (yakasimudzwa mutsvuku) mumaseru ePDS yakakonzera misiyano yakakura muchiitiko mukupindura kune akawanda mafambiro maitiro.Mudambudziko redu rezvikamu, boka iri rinotyaira maketani esarudzo uye kukanganisa maonero.Kusiyana neizvi, munyaya yemhinduro (FB), zvisarudzo zvepfungwa zvinogadzirwa mumasekete ekumusoro ne data sensory uye cognitive biases, uye simba guru rekudzika kweFB pamasero ePDS (mitsetse yakakora) inogadzira zviratidzo zvekusarudza.b Schematic inomiririra mamwe mamodheru eCDS nePDS zvishandiso.Pano zviratidzo zvePDS muMT zvinogadzirwa kwete chete nekuisa zvakananga kweV1, asiwo nekupinza kusina kunanga kweV1-V2-MT nzira.Nzira dzisina kunanga dzemuenzaniso dzinogadziriswa kuti dzipe kusarudzwa kumiganhu yezvinyorwa (grid inopindirana nzvimbo).Iyo MT layer CDS module inoita huremu hwezvakananga uye zvisina kunanga zvekuisa uye inotumira zvinobuda kune iyo PDS module.PDS inodzorwa nemakwikwi inhibition.Zvekare, izvo chete zvinongedzo zvinoratidzwa izvo zvinodikanwa kudhirowa yekutanga dhizaini yemuenzaniso.Pano, imwe nzira yeFF yakasiyana pane iyo yakarongwa mune inogona kutungamirira kumutsauko mukuru mukupindura kwemaserura kuPDS, zvakare zvichitungamirira kune kusarura mumatongerwo ezvisarudzo.Neimwe nzira, yakakura CP mumaseru ePDS inogona kunge ichiri mhedzisiro yekusarura musimba kana kugona kweFB kunamatira kumaseru ePDS.Humbowo hunotsigira maviri- uye matatu-nhanho MT PDS modhi uye iyo CP FF uye FB kududzira.
Macaques maviri evakuru (macaca mulatta), mumwe murume uye mukadzi (7 uye 5 wemakore zvichiteerana), anorema kubva 4.5 kusvika 9.0 kg, akashandiswa sezvinhu zvekudzidza.Pasati paitwa ongororo dzese dzekuvhiya dzisina mbereko, mhuka dzaisimwa nekamuri yekurekodha yakaitwa yemagetsi epamhepo achiswedera kunharaunda yeMT, simbi isina tsvina yemusoro (Crist Instruments, Hagerstown, MD), uye nzvimbo yeziso ine yakayerwa scleral yekutsvaga coil.(Cooner Wire, San Diego, California).Mitemo yose inoenderana neUnited States Department of Agriculture (USDA) mitemo uye National Institutes of Health (NIH) Mirayiridzo yeHumane Care uye Kushandiswa kweMhuka dzeLabhoritari uye yakagamuchirwa neYunivhesiti yeChicago Institutional Animal Care and Use Committee (IAUKC).
Zvese zvinokurudzira zvinoonekwa zvakaunzwa mune yakatenderedza aperture pakatarisana nedema kana grey kumashure.Panguva yekurekodha, nzvimbo uye dhayamita yegomba iri zvakagadziridzwa zvinoenderana neiyo classical receptive field ye neurons pa electrode tip.Isu takashandisa zvikamu zviviri zvakafara zvekukurudzira zvinoonekwa: psychometric stimuli uye tuning stimuli.
The psychometric stimulus is a grating pateni (20 cd/m2, 50% kusiyana, 50% duty cycle, 5 madhigirii/sec) yakagadzirwa superimposing maviri rectangular gratings kukukurwa mugwara perpendicular kwaakananga (Fig. 1b).Zvakagara zvichiratidzwa kuti vacherechedzi vevanhu vanoona aya mapatani egidhi senge bistable stimuli, dzimwe nguva sepateni imwe chete inofamba munzira imwechete (yakabatana kufamba) uye dzimwe nguva senzvimbo mbiri dzakaparadzana dzinofamba munzira dzakasiyana (transparent movement).zvikamu zvelattice pateni, yakatarisana nesymmetrically - kona iri pakati pema lattices kubva pa95 ° kusvika 130 ° (yakatorwa kubva pakaseti: 95 °, 100 °, 105 °, 115 °, 120 °, 125 °, 130 ° °, mukati mesesheni Isolation1 angle 5 ° isu takanga tisingabatanidzi data paIsolation1 angle - 9 data ° kana 270 ° (pattern orientation).Muchikamu chimwe nechimwe, kona imwe chete ye interlattice lattice yakashandiswa;panguva yega yega yega, kutaridzika kwemaitiro ekuedzwa kwega kwega kwakasarudzwa kubva pane zvingangoita zviviri.
Kuitira kusanzwisisa maonero egridi uye nekupa hwaro hwesimba hwemubairo wechiito, isu tinosuma zvisina tsarukano magadzirirwo mubhara rechiedza nhanho 72 yegidhi yega yega chikamu.Izvi zvinowanikwa nekuwedzera kana kuderera (nechiyero chakatarwa) kupenya kwechikamu chakasarudzwa chepixels (Fig. 1c).Nhungamiro yekufamba kwemavara inopa chiratidzo chakasimba chinoshandura maonero emucherechedzi kuenda kune yakabatana kana kujeka kufamba (Fig. 1c).Pasi pemamiriro ezvinhu akabatana, zvinyorwa zvose, pasinei nokuti ndezvipi chikamu chemafukidziro emucheka, zvinoshandurwa nenzira yemuenzaniso (Fig. 1c, inobatana).Mumamiriro ezvinhu akajeka, maitiro anofambisa perpendicular kune nzira ye grating iyo inovhara (Fig. 1c, yakajeka) (Supplementary Movie 1).Kudzora kuomarara kwebasa, mumasesheni mazhinji mutsauko weMichelson (Lmax-Lmin/Lmax+Lmin) weiyi mucherechedzo wekuumbwa wakasiyana kubva kune seti ye (-80, -40, -20, -10, -5, 0, 5)., 10, 20, 40, 80).Musiyano unotsanangurwa sekupenya kunoenderana kweraster (saka kukosha kwekusiyana kwe80% kunoguma nekugadzirwa kwe36 kana 6 cd/m2).Kwezvikamu zvitanhatu mutsoko N uye 5 zvikamu mutsoko S, takashandisa yakamanikana yemavara akasiyana siyana (-30, -20, -15, -10, -5, 0, 5, 10, 15, 20, 30), apo psychophysical maitiro anotevera maitiro akafanana sekuzara-range kusiyana, asi pasina saturation.
Tuning stimuli is sinusoidal grids (kusiyanisa 50%, 1 cycle/degree, 5 degrees/sec) ichifamba mune imwe ye16 yakaenzana nzira dzakapatsanurwa, kana sinusoidal grids inofamba munzira idzi (inosanganisira maviri akatarisana 135° makona akaiswa superimposed sinusoidal gratings pamusoro peumwe neumwe).munzira imwechete yemuenzaniso.