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Ukubona kokucaca okuphezulu kudinga ukusampula kahle kwe-retina nokuhlanganiswa ukuze kwakhiwe kabusha izakhiwo zento.Kubalulekile ukuqaphela ukuthi lapho uxuba amasampula wendawo avela ezintweni ezahlukene, ukunemba kulahleka.Ngakho-ke, ukuhlukaniswa, ukuqoqwa kwezindawo zesithombe ukuze kucutshungulwe ngokwehlukana, kubalulekile ekuboneni.Emsebenzini odlule, izakhiwo ze-lattice ze-bistable, ezingabhekwa njengendawo eyodwa noma ngaphezulu ezihambayo, zazisetshenziselwa ukutadisha le nqubo.Lapha, sibika ubudlelwano phakathi komsebenzi kanye nokwahlulela kwesegmenti ezifundeni ezimaphakathi zendlela yokubuka yezilwane zasendle.Ikakhulukazi, sithole ukuthi ama-neuron esikhashana aqondisa ngokukhetha ayezwela kuzimpawu zokuthungwa ezisetshenziselwa ukuhlanekezela umbono we-gratings e-bistable futhi abonisa ukuhlobana okubalulekile phakathi kwezilingo kanye nombono ozimele wesisusa esiqhubekayo.Lokhu kuhlotshaniswa kukhulu kumayunithi abonisa ukunyakaza komhlaba wonke ngamaphethini anezikhombisi-ndlela eziningi zendawo.Ngakho-ke, siphetha ngokuthi isizinda sesikhathi esimaphakathi siqukethe amasignali asetshenziselwa ukuhlukanisa izigcawu eziyinkimbinkimbi zibe izinto eziyisisekelo kanye nezindawo.
Umbono awuncikile kuphela ekubandlululweni okunembile kwezici zesithombe eziyisisekelo njengokuma konqenqema nesivinini, kodwa okubaluleke kakhulu ekuhlanganisweni okulungile kwalezi zici ukuze kubalwe izici zemvelo ezifana nokuma kwento kanye ne-trajectory1.Kodwa-ke, izinkinga ziphakama lapho izithombe ze-retina zisekela amaqembu amaningana wesici esivumelana ngokulinganayo 2, 3, 4 (Fig. 1a).Isibonelo, lapho amasethi amabili amasignali esivinini esondelene kakhulu, lokhu kungahunyushwa ngokunengqondo njengento eyodwa ehambayo noma izinto ezimbalwa ezihambayo (Fig. 1b).Lokhu kukhombisa imvelo yokuhlukanisa ngokwehlukana, okungukuthi akuyona into emisiwe yesithombe, kodwa inqubo yokuhumusha.Naphezu kokubaluleka kwakho okusobala kokubona okuvamile, ukuqonda kwethu isisekelo se-neural sokuhlukaniswa kombono kuhlala kungaphelele.
Umfanekiso wekhathuni wenkinga yokuhlukaniswa kombono.Umbono womuntu obukele ngokujula kukhiyubhu ye-Necker (kwesokunxele) ushintshana phakathi kwezincazelo ezimbili ezingaba khona (kwesokudla).Lokhu kungenxa yokuthi awekho amasiginali esithombeni avumela ubuchopho ukuthi bunqume ngokuhlukile umumo we-3D wesibalo (kuhlinzekwe isignali ye-monocular occlusion kwesokudla).b Uma amasignali anyakazayo amaningi ethulwa eduze kwendawo, isistimu yombono kufanele inqume ukuthi amasampula endawo avela entweni eyodwa noma ngaphezulu.Ukungaqondakali okungokwemvelo kwamasignali okunyakaza kwasendaweni, okungukuthi ukulandelana kokunyakaza kwento kungaveza umnyakazo wasendaweni ofanayo, okuholela ekuchazeni okuningi okubambekayo ngokulinganayo kokokufaka okubukwayo, okungukuthi izinkambu ze-vector lapha zingavela kusukela ekunyakazeni okuhambisanayo kwendawo eyodwa noma ukunyakaza okusobala kwezindawo ezigqagqene.c (kwesokunxele) Isibonelo sesivuseleli segridi yethu eyenziwe ngomumo.Ama-grating angama-rectangular akhukhuleka aqonde lapho eqonde khona ("izinkomba zengxenye" ​​- imicibisholo emhlophe) ayadlulana ukuze akhe iphethini yokugaya.I-lattice ingabonwa njengokuhamba okukodwa, okujwayelekile, okuxhunyiwe kwezikhombisi-ndlela (imicibisholo ebomvu) noma ukunyakaza okusobala kwezikhombisi-ndlela ezihlanganisiwe.Umbono we-lattice uhlanekezelwa ngokwengezwa kwezimpawu zokuthungwa kwephuzu okungahleliwe.(Maphakathi) Indawo egqanyiswe ngokuphuzi iyanwetshwa futhi iboniswe njengochungechunge lwamafreyimu amasignali ahambisanayo nasobala, ngokulandelanayo.Ukunyakaza kwechashazi esimweni ngasinye kuvezwa imicibisholo eluhlaza nebomvu.(Kwesokudla) Igrafu yendawo (x, y) yephoyinti lokukhetha ngokumelene nenombolo yozimele.Endabeni ehambisanayo, wonke ama-texture akhukhuleka aye ohlangothini olufanayo.Endabeni yokubonisa ngale, ukuthungwa kuhamba ngendlela yengxenye.d Umfanekiso wekhathuni womsebenzi wethu wokuhlukanisa ukunyakaza.Izinkawu zaqala uvivinyo ngalunye ngokulungisa ichashaza elincane.Ngemva kokulibaziseka okufushane, uhlobo oluthile lwephethini yokugreyitha (ukuhambisana/ukukhanya) nosayizi wesignali wokuthungwa (isb. umehluko) uvele endaweni ye-MT RF.Phakathi nokuhlolwa ngakunye, igridi ingase ikhukhuleke iye kolunye lwezikhombisi-ndlela ezimbili zephethini.Ngemuva kokuhoxiswa kokuvuselela, okuhlosiwe kokukhetha kuvele ngaphezulu nangaphansi kwe-MT RF.Izinkawu kufanele zibonise umbono wazo wegridi kuma-saccades kuya ekukhethweni okuhlosiwe okufanelekile.
Ukucutshungulwa kokunyakaza okubukwayo kubonakala kahle futhi ngaleyo ndlela kunikeza imodeli enhle kakhulu yokufunda iziyingi ze-neural zokuhlukaniswa kombono.Ucwaningo oluningana lokubala luye lwaphawula ukusetshenziswa kwamamodeli wokucubungula ukunyakaza kwezinyathelo ezimbili lapho ukulinganisa kokuqala kokuxazulula okuphezulu kulandelwa ukuhlanganiswa okukhethiwe kwamasampula wendawo ukuze kususwe umsindo nokubuyisela i-velocity7,8.Kubalulekile ukuqaphela ukuthi izinhlelo zokubona kufanele ziqikelele ukuthi lokhu kuhlanganisa kulinganiselwe kuphela kulawo masampula endawo avela ezintweni ezijwayelekile.Ucwaningo lwezengqondo luchaze izici zomzimba ezithonya indlela amasiginali anyakazayo ahlukaniswe ngayo, kodwa ukwakheka kwama-trajectories e-anatomical namakhodi e-neural kuhlala kuyimibuzo evulekile.Imibiko eminingi iphakamisa ukuthi amaseli akhetha umumo endaweni yesikhashana (MT) ye-primate cortex angamakhandidethi we-neural substrates.
Okubalulekile, kulezi zivivinyo zangaphambilini, izinguquko emsebenzini we-neural ohambisana nezinguquko zomzimba ekushukumiseni okubonakalayo.Kodwa-ke, njengoba kushiwo ngenhla, ukuhlukaniswa empeleni kuyinqubo yokuqonda.Ngakho-ke, ukucwaninga kwe-neural substrate yayo kudinga ukuxhumanisa izinguquko emsebenzini we-neural nezinguquko ekubonweni kwe-stimuli engaguquki.Ngakho-ke, siqeqeshe izinkawu ezimbili ukuze zibike ukuthi ingabe iphethini ye-bistable grating ecatshangelwayo eyakhiwe ngamagreyiti angamanxande akhukhulekayo kwakuyindawo eyodwa noma izindawo ezimbili ezizimele.Ukutadisha ubudlelwano phakathi komsebenzi we-neural kanye nokwahlulela kwesegmentation, siqophe umsebenzi owodwa ku-MT lapho izinkawu zenza lo msebenzi.
Sithole ukuhlobana okubalulekile phakathi kocwaningo lomsebenzi we-MT nokubona.Lokhu kuhlotshaniswa bekukhona noma ngabe izisusa ziqukethe izimpawu zokuhlukanisa ezigqamile.Ngaphezu kwalokho, amandla alo mphumela ahlobene nokuzwela kumasiginali wokuhlukanisa, kanye nenkomba yephethini.Lokhu kwakamuva kulinganisa izinga lapho iyunithi ikhipha khona umhlaba wonke kunokunyakaza kwendawo ngamaphethini ayinkimbinkimbi.Nakuba ukukhethwa kokuqondisa kwemfashini sekuyisikhathi eside kuqashelwa njengesici esichazayo se-MT, futhi amangqamuzana akhetha imfashini abonisa ukulungisela izisusa eziyinkimbinkimbi ezihambisana nombono womuntu walezo zisusa, ngokwazi kwethu konke, lobu ubufakazi bokuqala bokuhlobana phakathi kwamaphethini.inkomba kanye nokuhlukaniswa kwemibono.
Siqeqeshe izinkawu ezimbili ukuze zibonise umbono wazo wezisusa zegridi ekhukhulekayo (ukunyakaza okuhambisanayo noma okusobala).Izibukeli ezingabantu ngokuvamile zibona lezi zisusa njengokunyakaza okuhambisanayo noma okusobala okucishe kube yimvamisa efanayo.Ukuze unikeze impendulo elungile kulokhu kuhlolwa futhi usethe isisekelo somvuzo osebenzayo, sidale amasiginali okuhlukanisa ngokuthumela umbhalo ku-raster yengxenye eyenza i-lattice (Fig. 1c, d).Ngaphansi kwezimo ezihambisanayo, yonke imidwebo ihamba ngendlela yephethini (Fig. 1c, "coherent").Esimweni esisobala, ukuthungwa kuhamba nge-perpendicular ukuya ekuqondeni kwe-grating lapho kubekwe khona phezulu (Fig. 1c, "transparent").Silawula ubunzima bomsebenzi ngokushintsha ukugqama kwale lebula yokuthungwa.Ezivivinyweni ezicushiwe, izinkawu zaklonyeliswa ngezimpendulo ezihambisana nezimpawu zokuthungwa, futhi imiklomelo yanikezwa ngokungahleliwe (okungenzekanga okungu-50/50) ezivivinyweni eziqukethe amaphethini ngaphandle kwezimpawu zokuthungwa (isimo sokungafani kokuthungwa okuyiziro).
Idatha yokuziphatha evela ekuhlolweni okubili okuqokelwe ukumela abanye iboniswa kuMfanekiso 2a, futhi izimpendulo zihlelwa njengengxenye yezinqumo zokuhambisana nokungafani kwamasignali wokuthungwa (ukugqama obala kucatshangwa ukuthi kubi ngencazelo) kumaphethini asuka phezulu noma phansi, ngokulandelana. Sekukonke, umbono wezinkawu wokuhambisana/ukukhanya uthintwe ngokuthembekile kukho kokubili uphawu (okungafihli lutho, okuhambisanayo) namandla (okugqamayo) kwecue yokuthungwa (ANOVA; inkawu N: isiqondiso – F = 0.58, p = 0.45, uphawu – F = 1248, p <10−0 0 = 1, 2 ; monkey <2, 2 ; 2 ;             ;        ;         ;       ;       ;        ; umehluko we- monkey S: isiqondiso - F = 0.41, p = 0.52, uphawu - F = 2876.7, p <10-10, umehluko - F = 36.5, p <10−10). Sekukonke, umbono wezinkawu wokuhambisana/ukukhanya uthintwe ngokuthembekile kukho kokubili uphawu (okungafihli lutho, okuhambisanayo) namandla (okugqamayo) kwe-texture cue (ANOVA; inkawu N: isiqondiso – F = 0.58, p = 0.45, uphawu – F = 1248, p <10−1 = 1, 2, 2, 2, 2, p <10− 3, 2−1, 2−1 = 10; umehluko; ukhiye S: isiqondiso - F = 0.41, p = 0.52, uphawu - F = 2876.7, p <10-10, umehluko - F = 36.5, p <10-10). В целом на восприятие обезьянами когерентности/прозрачности достоверно влияли как знак (прозрачность, когерентность), так исток (кон) так и простира (кон) обезьяна N: направление — F = 0,58, p = 0,45, знак — F = 1248, p < 10−10, контраст – F = 22,63, p < 10; −10 обезья – 50 = F, p, p; ризнак - F = 2876,7, p <10−10, контраст - F = 36,5, р <10-10). Ngokuvamile, umbono wokuhambisana/ukubonisa obala kwezinkawu kwathinteke kakhulu kukho kokubili uphawu (ukukhanya, ukuhambisana) namandla (ukugqama) kwesici sombhalo (ANOVA; inkawu N: isiqondiso — F = 0.58, p = 0.45, uphawu — F = 1248, p <10− = 3 <0, 10: 1, 1, 1, 1, 1, 3, 1, 1, 1, 1, 1, 1, 1, 3, p - 1, 1, 1, 1, 1, 1, 1, 1, 1, 1, 1, 1, 1, 10, 10, 10, 10, p. F = 0.41, p = 0.52, uphawu - F = 2876.7, p <10 -10, umehluko - 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: S = 22.63, p <10;−1, S = 10 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;−0010.Ngokuvamile, umbono wokuhambisana kwenkawu/ukubonisa ngale kwathinteka kakhulu uphawu (ukubonisa ngale, ukuhambisana) nokuqina (ukugqama) kwezimpawu 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: ukuma - F = 0.58, p = 0.45, uphawu - F = 1248, p <10−10, Ukugqama - 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 Inkawu S: Ukuma - F = 0.41, p = 0.52, Uphawu - F = 2876.7, p <10-10, Ukuqhathanisa - F = 36.5, p <10-10).Imisebenzi eqongelelekayo ye-Gaussian ifakwe kudatha evela kuseshini ngayinye ukuze ibonise izici zengqondo ye-psychophysical yezinkawu.Emkhiwaneni.I-2b ibonisa ukusatshalaliswa kwesivumelwano kulawa mamodeli kuzo zonke izikhathi zazo zombili izinkawu.Sekukonke, izinkawu ziqede umsebenzi ngokunemba nangokungaguquguquki, futhi senqabe ngaphansi kuka-13% wezikhathi zezinkawu ezimbili ngenxa yokungangeni kahle kwemodeli ye-Gaussian enqwabelene.
izibonelo zokuziphatha kwezinkawu kumaseshini amele (n≥ izilingo ezingu-20 ngesimo ngasinye sokuvuselela).Kumaphaneli angakwesokunxele (kwesokudla), idatha evela kuseshini eyodwa yenkawu ye-N(S) ihlelwe njengamaphuzu okukhetha ahambisanayo (okudidiyelwe) ngokuqhathanisa nezimpawu zokugqama kwamasignali wokuthungwa (abscissa).Lapha kucatshangwa ukuthi ukuthungwa okusobala (okuhambisanayo) kunamanani angemuhle (okuhle).Izimpendulo zakhiwe ngokuhlukana ngokuya ngesiqondiso sokunyakaza kwephethini (phezulu (90 °) noma phansi (270 °)) ekuhlolweni.Kuzo zombili izilwane, ukusebenza, noma ngabe impendulo ihlukaniswa ngokugqama okungu-50/50 (PSE - imicibisholo eqinile) noma inani lokungafani kombhalo elidingekayo ukuze kusekelwe izinga elithile lokusebenza (umbundu - imicibisholo evulekile), ikulezi zikhombisi-ndlela zokukhukhuleka.b Ifakelwe i-histogram yamanani angu-R2 omsebenzi wokuqongelela we-Gaussian.Idatha yeMonkey S(N) iboniswa kwesokunxele (kwesokudla).c (Phezulu) I-PSE elinganiswa kugridi ehanjiswe phansi (ukuhleleka) uma kuqhathaniswa ne-PSE inyuse igridi (i-abscissa) ehleliwe, ngemiphetho emele ukusatshalaliswa kwe-PSE kusimo ngasinye nemicibisholo ebonisa incazelo yazo zonke izimo.Idatha yawo wonke amaseshini wezinkawu ze-N(S) inikezwa kukholamu yesokunxele (kwesokudla).(Ezansi) Umhlangano ofanayo nowedatha ye-PSE, kodwa yesici esifakiwe.Bekungekho mehluko obalulekile emikhawulweni ye-PSE noma izitayela zemfashini (bona umbhalo).d I-PSE kanye ne-slope (i-odine) kuhlelwa kuye ngokuma kwe-raster evamile kwengxenye yokuhlukaniswa kwe-angular ("i-angle ye-grating eqinile" - abscissa).Imibuthano evulekile izindlela, umugqa oqinile uyimodeli yokuhlehla efaneleka kakhulu, futhi umugqa onamachashazi uyisikhawu sokuzethemba esingu-95% semodeli yokuhlehla.Kukhona ukuhlobana okubalulekile phakathi kwe-PSE kanye ne-engeli yokuhlanganisa evamile, kodwa hhayi i-slope kanye ne-engeli yokuhlanganisa evamile, okuphakamisa ukuthi umsebenzi we-psychometric uyashintsha njengoba i-engeli ihlukanisa amalethisi engxenye, kodwa hhayi ukulola noma ukucaba.(Inkawu N, n = izikhathi ezingu-32; Inkawu S, n = izikhathi ezingu-43).Kuwo wonke amaphaneli, amabha wephutha amelela iphutha elijwayelekile lencazelo.Haha.Ukuhambisana, i-PSE subjective equality score, evamile.ukulinganisa.
Njengoba kuphawuliwe ngenhla, kokubili ukugqama kwezimpawu zokuthungwa kanye nesiqondiso sokunyakaza kwephethini kuyahlukahluka kuzo zonke izivivinyo, okushukumisayo kukhuphukela phezulu noma phansi esivivinyweni esithile.Lokhu kwenzelwa ukunciphisa imiphumela eguquguqukayo ye-psychophysical11 kanye ne-neuronal28.Ukuma kwephethini kuqhathaniswa nokuchema (iphuzu lokulingana ngokwesihloko noma i-PSE) (ukuhlolwa kwesamba sezinga le-Wilcoxon; imfene N: z = 0.25, p = 0.8; imfene S: z = 0.86, p = 0.39) noma umkhawulo womsebenzi ofakiwe (inani le-Wilcoxon irenki 4, 8: 0 = monkey; z. z = 0.49, p = 0.62) (Fig. 2c).Ngaphezu kwalokho, awukho umehluko obalulekile phakathi kwezinkawu ngezinga lokungafani kokuthungwa okudingekayo ukuze kugcinwe amazinga omkhawulo wokusebenza (N monkey = 24.5% ± 3.9%, S monkey = 18.9% ± 1.9%; Wilcoxon Rank sum , z = 1.01, p = 0.31).
Kuseshini ngayinye, siguqule i-engeli ye-interlattice ehlukanisa umumo wamalethisi engxenye.Ucwaningo lwezengqondo lubonise ukuthi abantu kungenzeka kakhulu babone iseli 10 njengexhumene lapho le engeli incane.Uma izinkawu bezibika ngendlela enokwethenjelwa umbono wazo wokuhambisana/ukubonisa izinto obala, khona-ke ngokusekelwe kulokhu okutholakele, umuntu ubengalindela i-PSE, umehluko wokuthungwa ohambisana nokuhlukaniswa okufanayo phakathi kokuhambisana nokukhetha obala, ukwanda ekuhlanganyeleni.i-lattice angle. Lokhu kwakunjalo ngempela (I-Fig. 2d; ukubhidlika kuzo zonke izikhombisi-ndlela zephethini, i-Kruskal–Wallis; inkawu N: χ2 = 23.06, p <10−3; inkawu S: χ2 = 22.22, p <10−3; ukuhlobana phakathi kwe-angle ejwayelekile - 6 −3; i-engeli ye-PSE = 6 p. 9; inkawu S: r = 0.76, p <10−13). Lokhu kwakunjalo ngempela (I-Fig. 2d; ukudilika kuzo zonke izikhombisi-ndlela zephethini, i-Kruskal–Wallis; inkawu N: χ2 = 23.06, p <10−3; inkawu S: χ2 = 22.22, p <10-3; ukuhlobana phakathi kwe-engeli evamile - 6 −3 nokhiye we-PSE = 6 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; < 10–3; корреляция между нормализованными угол решетки и PSE – обезьяна N: r = 0,67, p < 10-9, обезьяна S: r = 0,76-1 <10 <10). Lokhu kwenzeka ngempela (Umdwebo 2d; ukugoqa ngaphesheya kwephethini, i-Kruskal-Wallis; inkawu N: χ2 = 23.06, p <10-3; inkawu S: χ2 = 22.22, p <10-3; ukuhlobana phakathi kwe-lettice angle evamile kanye ne-PSE: 0, ukhiye we-9 = ukhiye we-lattice kanye ne-PSE: 1: 9, 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;猴子N:χ2 = 23.06,p <10-3猴子S:χ2-3;2 = 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 ; <10-3 ; <10-3 ; <2 : 2 - 3 = 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；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; обезь,3:2, 2, 3 лизованный межрешеточный угол). Lokhu kwakunjalo ngempela (I-Fig. 2d; phinda eduze kwe-axis yemodi, i-Kruskal-Wallis; inkawu N: χ2 = 23.06, p <10-3; inkawu S: χ2 = 22.22, p <10-3; ikhoneni le-interlattice elijwayelekile). PSE-обезьяна N: r = 0,67, p < 10–9, обезьяна S: r = 0,76, p < 10–13). I-PSE imfene N: r = 0.67, p <10–9, inkawu S: r = 0.76, p <10–13).Ngokuphambene, ukushintsha i-angle ye-interlattice akuzange kube nomthelela obalulekile emthambekeni womsebenzi we-psychometric (Fig. 2d; i-cross-modal orientation fold, i-Kruskal-Wallis; inkawu N: χ2 = 8.09, p = 0.23; inkawu S χ2 = 3.18 = 3.18 , i-angle ye-interlarre = 7, p, i-engeli ye-interlatti, i-p, i-interlatti, i-p. N: r = -0.4, p = 0.2, inkawu S: r = 0.03, p = 0.76).Ngakho-ke, ngokuhambisana nedatha ye-psychophysical yomuntu, umthelela ojwayelekile wokushintsha i-engeli phakathi kwama-gratings ukushintshwa kwamaphoyinti okufuduka, hhayi ukwanda noma ukwehla kokuzwela kumasiginali wokuhlukanisa.
Okokugcina, imiklomelo inikezwa ngokungahleliwe namathuba angu-0.5 ezivivinyweni ezinokungafani kokuthungwa okunguziro.Uma zonke izinkawu bezikwazi lokhu kungenzeki okuyingqayizivele futhi zikwazi ukuhlukanisa phakathi kokungafani kokuthungwa okunguziro kanye ne-cue stimu, bezingakwazi ukuthuthukisa amasu ahlukene ezinhlobo ezimbili zokuhlola.Imibono emibili iphakamisa ngokuqinile ukuthi lokhu akunjalo.Okokuqala, ukushintsha i-engeli yokugaya kube nemiphumela efana nekhwalithi kumaphuzu wokuqhathanisa we-cue kanye ne-zero (Fig. 2d kanye ne-Supplementary Fig. 1).Okwesibili, kuzo zombili izinkawu, ukukhethwa kwesilingo se-bistable akunakwenzeka ukuthi kube impinda yokukhethwa komvuzo wakamuva (okwangaphambilini) (ukuhlolwa kwe-binomial, izinkawu ezingu-N: 0.52, z = 0.74, p = 0.22; Izinkawu ze-S: 0.51, r = 0.9, p = 0.18).
Sengiphetha, ukuziphatha kwezinkawu emsebenzini wethu wokuhlukanisa kwakungaphansi kokulawulwa okuhle kokugqugquzela.Ukuncika kwezinqumo zombono kuphawu nosayizi wezimpawu zokuthungwa, kanye nezinguquko ku-PSE nge-engeli yokugaya, kubonisa ukuthi izinkawu zibike umbono wazo oqondile wokuhambisana kwemoto/ukukhanya.Okokugcina, izimpendulo zezinkawu ezivivinyweni zokuqhathanisa zokuthungwa kweqanda azizange zithintwe umlando womvuzo wezivivinyo zangaphambilini futhi zathinteka kakhulu izinguquko ze-angular phakathi kwama-raster.Lokhu kusikisela ukuthi izinkawu ziyaqhubeka nokubika umbono wazo wokucushwa kwe-lattice surface ngaphansi kwalesi simo esibalulekile.
Njengoba kushiwo ngenhla, ukuguquka kokushintshashintsha kokuthungwa kusuka kokunegethivu kuye kokuphozithivu kulingana nokushintsha kombono we-stimu kusuka kokusobala kuya kokuhambisanayo.Ngokuvamile, kuseli elinikeziwe, impendulo ye-MT ivame ukukhula noma yehle njengoba ukuhluka kokuthungwa kushintsha kusuka kokubi kuye kokuphozithivu, futhi isiqondiso salo mphumela ngokuvamile sincike ekuqondeni kokunyakaza kwephethini/ingxenye.Isibonelo, ijika lokushuna eliqondisayo lamaseli amabili e-MT amelelayo aboniswa kuMfanekiso 3 kanye nezimpendulo zalawa maseli kuma-grating aqukethe amasiginali ahambisanayo noma aphezulu ahambisanayo noma abonisa ngale.Sizame ngandlela thize ukulinganisa kangcono lezi zimpendulo zegridi, ezingase zihlobane nokusebenza kwengqondo kwezinkawu zethu.
Indawo ezungezile yejika lokushuna eliqondisayo lemfene emele i-MT cell S ngokuphendula kumalungu afanayo e-sinusoidal.I-engeli ikhombisa isiqondiso sokunyakaza kwe-grating, ubukhulu bubonisa ukuphuma, futhi isiqondiso seseli esincanyelwayo sigqagqana cishe ngo-90° (phezulu) nesiqondisindlela sengxenye yengxenye yendlela yephethini yegridi.b I-histogram yesikhathi se-stimulus-time (PSTH) yegridi yokuphendula, ihanjiswe ohlangothini lwesifanekiso ngo-90° (eboniswa ngokohlelo kwesokunxele) kuseli eliboniswe kokuthi.Izimpendulo zihlelwa ngohlobo lwehinti yokuthungwa (okuhlangene/okusobala - iphaneli ephakathi/kwesokudla ngokulandelanayo) kanye nokugqama kwe-Michelson (ukusikisela kombala we-PSTH).Imizamo elungile kuphela eboniswa kuhlobo ngalunye lwezimpawu zokuthungwa ezinomehluko ophansi kanye nokugqama okuphezulu.Amaseli aphendule kangcono kumaphethini e-lattice akhukhulekayo anezimpawu zokuthungwa ezibonisa ngale, futhi impendulo kulawa maphethini yanda ngokungafani kokuthungwa okukhulayo.c, d yizimiso ezifanayo njengaku-a no-b, kodwa kumaseli e-MT ngaphandle kwe-monkey S, umumo wawo awuncamelayo ucishe udlule lowo wegridi edonsela phansi.Iyunithi incamela ama-grating adonsela phansi anezimpawu zokuthungwa okuhambisanayo, futhi impendulo kulawa maphethini iyanda ngokungafani kokwakheka.Kuwo wonke amaphaneli, indawo enomthunzi imelela iphutha elijwayelekile lencazelo.Uyakhuluma.Ama-spikes, imizuzwana.okwesibili.
Ukuze uhlole ubudlelwano phakathi kokulungiswa kwendawo ye-lattice (ehambisanayo noma ebonisa ngale) njengoba kuboniswa amasignali ethu okuthungwa kanye nomsebenzi we-MT, siqale sahlehlisa ukuhlobana phakathi kwamaseli okunyakaza okuhambisanayo (i-positive slope) noma ukunyakaza okusobala (i-negative slope) ngokuhlehla.inikezwe ukuhlukanisa amaseli ngokwesilinganiso sokuphendula kophawu uma kuqhathaniswa nokugqama (ngokuhlukene kundlela ngayinye yemodi).Izibonelo zalawa majika okushuna ama-lattice asuka kuseli yesibonelo esifanayo kuMfanekiso 3 aboniswa kuMfanekiso 4a.Ngemva kokuhlukaniswa, sisebenzise ukuhlaziywa kokusebenza kowamukelayo (i-ROC) ukuze silinganise ukuzwela kweseli ngalinye ekushintsheni amasiginali wokuthungwa (bona Izindlela).Imisebenzi ye-neurometric etholwe ngale ndlela ingaqhathaniswa ngokuqondile nezici ze-psychophysical zezinkawu kuseshini efanayo ukuze kuqhathaniswe ngokuqondile ukuzwela kwengqondo ye-psychophysical yama-neurons nokuthungwa kwe-lattice.Senze ukuhlaziya okubili kokutholwa kwesignali kuwo wonke amayunithi kusampula, sibala izici ezihlukene ze-neurometric ohlangothini ngalunye lwephethini (futhi, phezulu noma phansi).Kubalulekile ukuqaphela ukuthi, kulokhu kuhlaziya, sifake kuphela izivivinyo lapho (i) isisusa siqukethe inkomba yokuthungwa kanye (ii) nezinkawu zaphendula ngokuvumelana nalolo phawu (okungukuthi, ukuhlola “okulungile”).
Izilinganiso zomlilo zenziwa ngokumelene nokuhluka kophawu lokuthungwa, ngokulandelanayo, kuma-grating asuka phezulu (kwesokunxele) noma phansi (kwesokudla), umugqa oqinile umelela ukuhlehla komugqa okulingana kahle kakhulu, futhi idatha kumugqa ophezulu (phansi) ithathwa kulawo aboniswe ku-Fig.Ilayisi.3a Iseli, b (Fig. 3c, d).Izici zemithambeka yokuhlehla zisetshenziselwe ukwabela izinkomba zokuthungwa ezikhethwayo (ezihambisanayo/ezibonisa ngale) kunhlanganisela yokuma kweseli/i-lattice ngayinye (n ≥ izilingo ezingu-20 ngesimo ngasinye sokuvuselela).Amabha wephutha amele ukuchezuka okujwayelekile kwencazelo.Imisebenzi ye-neurometric yamayunithi aboniswe ku-ba ichazwa kanye nemisebenzi ye-psychometric eqoqwe phakathi neseshini efanayo.Manje, esicini ngasinye, sihlela ukukhetha kwethiphu yamathuluzi (ukudidiyela) (bona umbhalo) njengephesenti lokugqama kophawu lokuthungwa (abscissa).Ukuhluka kokwakheka kushintshiwe ukuze izeluleko zamathuluzi ezithandwayo zibe zinhle futhi amathuluzi angenalutho abe negethivu.Idatha esuka kumagridi adonsela phezulu (phansi) aboniswa kumaphaneli angakwesokunxele (kwesokudla), emigqeni engaphezulu (ephansi) - idatha evela kumaseli aboniswe ku-Fig. 3a,b (Fig. 3c,d).Izilinganiso ze-neurometric ne-psychometric threshold (N/P) ziboniswa kuphaneli ngayinye.Uyakhuluma.Ama-spikes, imizuzwana.isekhondi, inkomba.isiqondiso, isifundazwe sikhethwa, psi.Psychometry, Neurology.
Amajika okushuna ama-lattice kanye nemisebenzi ye-neurometric yamaseli amabili e-MT amele kanye nemisebenzi yawo yengqondo ehlobene, ehlanganiswe kanye nalezi zimpendulo, kuboniswa kumaphaneli aphezulu naphansi oMfanekiso 4a,b, ngokulandelana.Lawa maseli abonisa ukukhula okucishe kube okuyisidina noma ukwehla njengoba isu lokuthungwa lisuka ekubeni obala liye ekuhlanganeni.Ngaphezu kwalokho, isiqondiso namandla alesi sibopho sincike ekuqondisweni kokunyakaza kwe-lattice.Ekugcineni, imisebenzi ye-neurometric ebalwa kusukela ezimpendulweni zalawa maseli yasondela kuphela (kodwa noma kunjalo ayizange ihambisane) nezakhiwo ze-psychophysical zokunyakaza kwegridi eyodwa.Kokubili imisebenzi ye-neurometric kanye ne-psychometric yafingqwa ngama-threshold, okusho ukuthi ehambelana cishe nama-84% wokuqhathanisa okukhethwe kahle (okuhambisana nencazelo + 1 sd yomsebenzi ohlanganisiwe we-Gaussian ohlanganisiwe).Kuwo wonke amasampula, isilinganiso se-N/P, isilinganiso somkhawulo we-neurometric kuya kwese-psychometric, silinganiselwa ku-12.4 ± 1.2 enkawuni engu-N kanye no-15.9 ± 1.8 ku-monkey S, kanye nokuthi i-lattice ihambe okungenani iye ohlangothini olulodwa, kuphela Ku-~16% (18).%) amayunithi avela enkawini N (inkawu S) (Fig. 5a).Kusukela kusibonelo seseli esiboniswe esithombeni.Njengoba kubonakala ku-Figure 3 kanye ne-4, ukuzwela kwama-neurons kungathinteka ubuhlobo phakathi kokuma okuthandwayo kweseli kanye nesiqondiso sokunyakaza kwe-lattice okusetshenziswe ekuhloleni.Ikakhulukazi, amajika okulungisa umumo ku-Fig. 3a,c abonisa ubudlelwano phakathi kwesilungiselelo se-neuron yohlelo olulodwa lwe-sinusoidal kanye nokuzwela kwayo ukunyakaza okusobala/okuhambisanayo kuhlelo lwethu oluthungwe. Kwakunjalo kuzo zombili izinkawu (ANOVA; izikhombisi-ndlela ezithandwayo eziboshwe ngokulungiswa okungu-10°; inkawu N: F = 2.12, p <0.01; imfene S: F = 2.01, p <0.01). Kwakunjalo kuzo zombili izinkawu (ANOVA; izikhombisi-ndlela ezithandwayo eziboshwe ngokulungiswa okungu-10°; inkawu N: F = 2.12, p <0.01; imfene S: F = 2.01, p <0.01). Это имело место для обеих обезьян (ANOVA; относительные предпочтительные направления объединены в группы с разрешением, 10° 1; обезьяна S: F = 2,01, p <0,01). Kwakunjalo kuzo zombili izinkawu (ANOVA; izikhombisi-ndlela ezithandwayo eziqoqwe ngo-10° ukulungiswa; inkawu N: F=2.12, p<0.01; imfene 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 以 10 ° 分辨率 合并 的 相对 方向 : 猴子 n: : f = 10 p2.; f = 10 p2. <0.01........）) Это имело место для обеих обезьян (ANOVA; относительная предпочтительная ориентация объединена при разрешении 10°; обезREшении 10°; обез:1, обезья = 10°; обез:1, на S: F = 2,01, p <0,01). Kwakunjalo kuzo zombili izinkawu (ANOVA; umumo okhethwayo wesihlobo uhlanganiswe ngokulungiswa okungu-10°; inkawu N: F=2.12, p<0.01; imfene S: F=2.01, p<0.01).Uma kubhekwa izinga elikhulu lokuhlukahluka kokuzwela kwe-neuron (Fig. 5a), ukuze ubone ngeso lengqondo ukuncika kokuzwela kwe-neuron ekumakiseni okuthandwayo okuhlobene, siqale sajwayeza umumo okhethwayo weseli ngalinye ube umumo “ongcono kakhulu” wokunyakaza kwephethini yegridi (okungukuthi isiqondiso).lapho igridi yakha i-engeli encane kakhulu phakathi kokuma kweseli okuncanyelwayo kanye nokuma kwephethini yokugaya).Sithole ukuthi ama-threshold ahlobene ama-neuron (umbundu wokuma/umbundu we-lattice “okubi kakhulu” wokuma kwelathisi “okungcono kakhulu”) ayahluka ngokwale ndlela evamile yokuma ekhethwayo, neziqongo zalesi silinganiso sombundu ezenzeka kumaphethini noma umumo wengxenye (Umfanekiso 5b).)). Lo mphumela awukwazanga ukuchazwa ngokuchema ekusabalaliseni izikhombisi-ndlela ezikhethwayo kumayunithi kusampula ngayinye ukuya kweyodwa yephethini ye-plaid noma izikhombisi-ndlela zengxenye (I-Fig. 5c; ukuhlolwa kwe-Rayleigh; imfene N: z = 8.33, p <10−3, isilinganiso esiyindilinga = 190.13 deg 30: 9 = 190.13 deg 30: 9 = 40 = 190 = 30 ± 4; 9. 5) futhi yayingaguquguquki kuwo wonke ama-engeli we-plaid inter-grating (I-Supplementary Fig. 2). Lo mphumela awukwazanga ukuchazwa ngokuchema ekusabalaliseni izikhombisi-ndlela ezikhethwayo kumayunithi kusampula ngayinye ukuya kweyodwa yephethini ye-plaid noma izikhombisi-ndlela zengxenye (Fig. 5c; Ukuhlolwa kwe-Rayleigh; imfene N: z = 8.33, p <10−3, indilinga incazelo = 190.13 demon 80 p. ± 9; 45) futhi yayingaguquguquki kuwo wonke ama-engeli we-plaid inter-grating (I-Supplementary Fig. 2). Этот эффект нельзя было объяснить смещением распределения предпочтительных направлений в единицах в каждой выборкей выборке выборкей й или направлений компонентов (рис. 5в; критерий Рэлея; обезьяна N: z = 8,33, p < 10–3). Lo mphumela awukwazanga ukuchazwa ngokushintsha ekusabalaliseni kwezikhombisi-ndlela ezikhethwayo kumayunithi kusampula ngayinye ukuya kweyodwa yezikhombisi-ndlela ezihloliwe noma izikhombisi-ndlela zengxenye (I-Fig. 5c; ukuhlolwa kwe-Rayleigh; imfene N: z = 8.33, p <10–3)., isilinganiso esiyindilinga = 190.13 degrees ± 9.83 degrees;inkawu S: z = 0.79, p = 0.45) futhi yayifana kuwo wonke amakhona egridi yeplaid (Umfanekiso Owengeziwe 2).這种效应不能通过每个樣本中单元中的优选方向分布偏向格子图案或组件方向之一來解）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 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 元图 图 瑞利 测试 ； 子子 n : z = 8.33 , p <10-3 , 平均值 平均值 圆形 圆形 圆墢 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). Lo mphumela awukwazi ukuchazwa iqiniso lokuthi ukusatshalaliswa kokuma okuthandwayo kumaseli kusampula ngayinye kuchemile noma kusakhiwo se-lattice noma kuya kokunye kokuma kwengxenye (Umfanekiso 5c; ukuhlolwa kuka-Rayleigh; inkawu N: z = 8.33, p <10–3)., isilinganiso esiyindilinga) = 190.13 degrees ± 9.83 degrees;inkawu S: z = 0.79, p = 0.45) futhi ayelingana ngama-engeli e-lattice phakathi kwamagridi (I-Fig. 2 eyengeziwe).Ngakho-ke, ukuzwela kwama-neurons kumagridi aqoshiwe kuncike, okungenani ngokwengxenye, ezintweni eziyisisekelo zokushuna kwe-MT.
Iphaneli engakwesokunxele ibonisa ukusatshalaliswa kwezilinganiso ze-N/P (umkhawulo we-neuron/psychophysiological);iseli ngalinye lihlinzeka ngamaphoyinti edatha amabili, elilodwa ngesiqondiso ngasinye lapho iphethini ihamba khona.Iphaneli yesokudla ihlela amathreshold e-psychophysical (i-ordinate) ngokumelene nama-neuronal thresholds (abscissa) kuwo wonke amayunithi kusampula.Idatha esemugqeni ophezulu (phansi) isuka kwimfene engu-N (S).b Ama-threshold ratios ajwayelekile ahlelwa ngokuqhathaniswa nobukhulu bomehluko phakathi kokuma kwe-lattice okulungile nokuma kweseli okuthandwayo.Isiqondiso "esingcono kakhulu" sichazwa njengesiqondiso sesakhiwo sokugaya (kukalwa nge-grating eyodwa ye-sinusoidal) eseduze nesiqondiso seseli esithandwayo.Idatha yaqale yaboshwa ngomumo okhethwayo ojwayelekile (imigqomo engu-10°), kwase kuthi izilinganiso ze-threshold zenziwa zaba ngokwejwayelekile zaba senani eliphezulu futhi zalinganiswa phakathi komgqomo ngamunye.Amaseli anomumo okhethwayo amakhudlwana kancane noma amancane kunomumo wezingxenye ze-lattice abe nomehluko omkhulu kakhulu wokuzwela ukuma kwephethini ye-lattice.c I-histogram ephinki yokusatshalaliswa kokuma okuthandwayo kwawo wonke amayunithi e-MT aqoshwe enkawini ngayinye.
Ekugcineni, impendulo ye-MT ilungiswa ngokuqondiswa kokunyakaza kwe-grating kanye nemininingwane yezimpawu zethu zokuhlukanisa (ukwakheka).Ukuqhathaniswa kokuzwela kwe-neuronal kanye ne-psychophysical kubonise ukuthi, ngokuvamile, amayunithi e-MT ayengazweli kangako kumasiginali wokuthungwa ahlukile kunezinkawu.Nokho, ukuzwela kwe-neuron kushintshile kuye ngomehluko phakathi kokuma okuncanyelwayo kweyunithi kanye nesiqondiso sokunyakaza kwegridi.Amaseli azwela kakhulu avamise ukuba nokuncamelayo kokuma okucishe kufake iphethini ye-lattice noma umumo wokukodwa wengxenye, futhi isethi encane yamasampuli ethu abezwela noma azwela kakhulu kunombono wezinkawu womehluko ohlukile.Ukuze sinqume ukuthi amasiginali asuka kulawa mayunithi azwela kakhulu ahlotshaniswa kakhulu nombono ezimfeneni, sihlole ukuhlobana phakathi kokubona nezimpendulo ze-neuronal.
Isinyathelo esibalulekile ekusunguleni ukuxhumana phakathi komsebenzi we-neural nokuziphatha ukusungula ukuhlobana phakathi kwama-neurons kanye nezimpendulo zokuziphatha ezishukumisayo njalo.Ukuze uxhumanise izimpendulo ze-neural nezahlulelo zokuhlukaniswa, kubalulekile ukudala isikhuthazo, nakuba sifana, sibonwa ngendlela ehlukile ezivivinyweni ezihlukene.Ocwaningweni lwamanje, lokhu kuvezwa ngokusobala ukuthungwa okungafani kwe grating.Nakuba sigcizelela ukuthi, ngokusekelwe emisebenzini ye-psychometric yezilwane, amagreyidi anokwahluka okuncane kombhalo (ngaphansi kuka-20%) ngokuvamile abhekwa njengokuhambisanayo noma okusobala.
Ukuze silinganise izinga izimpendulo ze-MT ezihlobana ngalo nemibiko yombono, senze ukuhlaziya okungenzeka ukuthi okukhethiwe (CP) kwedatha yethu yegridi (bona 3).Ngamafuphi, i-CP iyi-non-parametric, isilinganiso esingajwayelekile esilinganisa ubudlelwano phakathi kwezimpendulo ze-spike nezinqumo zokuqonda30.Ukukhawulela ukuhlaziya ezivivinyweni kusetshenziswa amagridi anokungafani kombhalo okuyiziro namaseshini lapho izinkawu zenze khona izinketho okungenani ezinhlanu kuhlobo ngalunye lwalezi zilingo, sibale i-SR ngokuhlukene ohlangothini ngalunye lokunyakaza kwegridi. Kuzo zonke izinkawu, sibone inani le-CP elisho ukuthi likhulu kakhulu kunalokho ebesingalilindela ngenhlanhla (Umfanekiso 6a, d; imfene N: i-CP eqondile: 0.54, 95% CI: (0.53, 0.56), ukuhlolwa kwe-t okuzinhlangothi ezimbili ngokumelene ne-null ye-CP = 0.5, t = 6.5 mon: 9; I-5% CI: (0.54, 0.57), i-t-test enezinhlangothi ezimbili, t = 9.4, p <10−13). Kuzo zonke izinkawu, sibone inani le-CP elisho ukuthi likhulu kakhulu kunalokho ebesingalilindela ngenhlanhla (Umfanekiso 6a, d; imfene N: i-CP eqondile: 0.54, 95% CI: (0.53, 0.56), ukuhlolwa kwe-t okuzinhlangothi ezimbili ngokumelene ne-null ye-CP = 0.5, t = 6.5 mon: 9; 5% CI: (0.54, 0.57), i-t-test enezinhlangothi ezimbili, t = 9.4, p <10−13).Ezimfeneni, sibone i-CP emaphakathi ephakeme kakhulu kunokulindelekile ngokungahleliwe (Fig. 6a, d; monkey N: mean CP: 0.54, 95% CI: (0.53, 0.56), T-test enemisila emibili vs. null values).I-CP = 0,5, t = 6,7, p <10–9; I-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: i-CP emaphakathi: 0.55, 95% CI: (0.54, 0.57), ukuhlolwa kwe-t enemisila emibili, 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:3,600.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), 双边t 检验 1, 4 − 1, 4.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: : ; 坳 5: 5 , 9, 9 .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, 5,5-5, 55% критерий, t = 9,4, p < 10- 13) . Ezimfeneni, sibone amanani ashoyo e-CP ngaphezu kwalokho esingase sikulindele ngenhlanhla (Umfanekiso 6a, d; imfene N: i-CP esho: 0.54, 95% CI: (0.53, 0.56), ukuhlolwa kwe-t enemisila emibili iqhudelana noziro = 0.5, t = CP mon 10-5: 9, % <p. I-CI: (0.54, 0.57), umbandela we-t-tailed two, t = 9.4, p <10-13).Ngakho-ke, ama-neuron e-MT athambekele ekuvutheleni umlilo ngamandla ngisho nalapho kungekho zimpawu zokuhlukaniswa okusobala, lapho umbono wesilwane wokunyakaza kwe-lattice ufana nokuthandwa yiseli.
ukusatshalaliswa kwamathuba okukhethwa kwamagridi angenawo amasiginali wokuthungwa kwamasampuli aqoshwe enkawini N. Iseli ngalinye linganikela ngamaphoyinti edatha amabili (elilodwa ohlangothini ngalunye lokunyakaza kwegridi).Inani elilinganiselwe le-CP ngaphezu kokungahleliwe (imicibisholo emhlophe) libonisa ukuthi sekukonke kunobudlelwano obubalulekile phakathi komsebenzi we-MT nokubona.b Ukuze sihlole umthelela wanoma ikuphi ukukhetha okungaba khona ukuchema, sibale i-CP ngokuhlukana nganoma yisiphi isikhuthazo izinkawu zenze okungenani iphutha elilodwa.Amathuba okukhetha ahlelwe njengomsebenzi wesilinganiso sokukhetha (pref/null) sazo zonke izisusa (kwesokunxele) kanye namanani aphelele wokugqama kophawu lokuthungwa (kwesokudla, idatha esuka kumaseli angawodwana angu-120).Ulayini oqinile nendawo enomthunzi kuphaneli yesokunxele imelela isilinganiso esingu-± sem yesilinganiso esinyakazayo samaphoyinti angu-20.Amathuba okukhetha abalwe izisusa ezinezilinganiso zokukhetha ezingalinganiseli, njengamagridi anokugqama kwesignali ephezulu, ahluke kakhulu futhi aye ahlanganiswa ngamathuba.Indawo enomthunzi ompunga kuphaneli engakwesokudla igcizelela umehluko wezici ezifakwe ekubalweni kwamathuba aphezulu okukhetha.c Amathuba okukhetha okukhulu (okudidiyelwe) akhiwe ngokumelene nomngcele we-neuron (abscissa).Amathuba okukhethwa ahlotshaniswa kabi kakhulu ne-threshold.I-convention df iyafana ne-ac kodwa isebenza kudatha engu-157 eyodwa evela ku-monkey S ngaphandle kwalapho kuphawulwe ngenye indlela.g Amathuba aphezulu okukhetha (okudidiyelwe) kuhlelwa ngokumelene nendawo evamile ethandwayo (i-abscissa) yenkawu ngayinye kwezimbili.Iseli ngalinye le-MT linikele ngamaphoyinti edatha amabili (elilodwa endaweni ngayinye yesakhiwo se-lattice).h Iziqephu zamabhokisi amakhulu zamathuba okukhetha e-engeli ngayinye ye-inter-raster.Umugqa oqinile uphawula i-median, imiphetho engezansi nangaphezulu yebhokisi imelela ama-percentiles angu-25 kanye nama-75 ngokulandelanayo, amadevu anwetshwa izikhathi ezingu-1.5 ububanzi be-interquartile, futhi okuphuma ngaphandle kwalo mkhawulo kuyaphawulwa.Idatha kumaphaneli angakwesokunxele (kwesokudla) isuka kumaseli wezinkawu angama-N(S) angama-120 (157).i Amathuba aphezulu okukhetha (okudidiyelwe) ahlelwa ngesikhathi sokuqala kwesisusa (abscissa).I-CP enkulu ibalwe konxande abaslayidayo (ububanzi obungu-100 ms, isinyathelo esingu-10 ms) phakathi nokuhlola konke kwase kwenziwa i-avareji phezu kwamayunithi.
Olunye ucwaningo lwangaphambilini lubike ukuthi i-CP incike enanini elihlobene lezilingo ekusabalaliseni isilinganiso se-basal, okusho ukuthi lesi silinganiso asithembeki kangako kuzinto ezishukumisayo ezibangela umehluko omkhulu engxenyeni yenketho ngayinye.Ukuze sihlole lo mphumela kudatha yethu, sibale i-CP ngokuhlukene yazo zonke izisusa, kungakhathaliseki ukuthi isibonakaliso sihluke kanjani, futhi izinkawu zenze okungenani isilingo esisodwa samanga.I-CP ihlelwe ngokumelene nesilinganiso sokukhetha (pref/null) sesilwane ngasinye kuMfanekiso 6b no-e (iphaneli yesokunxele), ngokulandelana.Uma ubheka ama-avareji anyakazayo, kuyacaca ukuthi i-CP ihlala ingaphezu kwamathuba phezu kwenani elibanzi lokukhetha, yehla kuphela lapho amathuba ehla (akhuphuka) ngezansi (ngenhla) 0.2 (0.8).Ngokusekelwe kuzici ze-psychometric zezilwane, singalindela ama-coefficients okukhetha alo bukhulu ukuthi asebenze kuphela kusisusa esinezimpawu zokuthungwa ezihluke kakhulu (ezihambisanayo noma ezibonisa ngale) (bona izibonelo zezici ze-psychometric ku-Fig. 2a, b).Ukuze sinqume ukuthi lokhu kwakunjalo kanye nokuthi i-PC ebalulekile iphikelele ngisho nasekushukumiseni ngezimpawu ezicacile zokuhlukanisa, sihlole umthelela wamanani wokungafani wombhalo aphelele ku-PC (Fig. 6b, e-right).Njengoba bekulindelekile, i-CP ibiphezulu kakhulu kunethuba lezisusa eziqukethe kufika kokumaphakathi (~20% wokungafani noma okuphansi) izimpawu zesegimenti.
Emisebenzini yokuqondisa, isivinini, kanye nokungaboni ngasolinye, i-MT CP ivame ukuba phezulu kuma-neuron azwela kakhulu, mhlawumbe ngenxa yokuthi la ma-neurons aphethe amasignali afundisa kakhulu30,32,33,34.Ngokuvumelana nalokhu okutholakele sibone ukuhlobana okuthobekile kodwa okubalulekile phakathi kwe-CP enkulu, ebalwe kusukela kumazinga okudubula anezikolo ezingu-z kuwo wonke umehluko we-texture cue ogqanyiswe kuphaneli elingakwesokudla kakhulu le-Fig.I-6b, e, ne-neuronal threshold (Fig. 6c, f; geometric mean regression; monkey N: r = -0.12, p = 0.07 monkey S: r = -0.18, p <10-3). I-6b, e, ne-neuronal threshold (Fig. 6c, f; geometric mean regression; monkey N: r = -0.12, p = 0.07 monkey S: r = -0.18, p <10-3).Ngokuvumelana nalokhu okutholakele, sibone ukuhlobana okuthobekile kodwa okubalulekile phakathi kwe-CP enkulu ebalwa kusukela ku-excitation frequency z-score kusukela ekuhlukeni kwesignali yokuthungwa okugqanyiswe kuphaneli engakwesokudla ye-Fig. 6b, e, ne-neuronal threshold (Fig. 6c, f; geometric).ukuhlehla kwejiyomethri;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，我們观察到大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 amayunithiS: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 子S:r = -0.18,p <10-3）.Ngokuvumelana nalokhu okutholakele, sibone ukuhlobana okuthobekile kodwa okuphawulekayo phakathi kwama-CV amakhulu njengoba kuboniswe ku-Fig. 6b, e kanye nemingcele ye-neuron (Umfanekiso 6c, f; ukwehla kwesilinganiso sejometri; inkawu N: r = -0.12, p = 0.07).Обезьяна S: г = -0,18, р < 10-3). Imfene S: r = -0.18, p <10-3).Ngakho-ke, amasu avela kumayunithi afundisa kakhulu avame ukukhombisa ukuhambisana okukhulu nezinqumo zokuhlukaniswa kwezigaba ezimfeneni, okubalulekile kungakhathaliseki ukuthi yiziphi izimpawu zombhalo ezingezwe ekukhetheni kombono.
Njengoba sake sasungula ubudlelwano phakathi kokuzwela kumasiginali wokuthungwa kwegridi kanye ne-neuronal orientation ekhethwayo, sizibuze ukuthi ngabe bukhona yini ubudlelwano obufanayo phakathi kwe-CP nokuma okuncanyelwayo (Fig. 6g).Lokhu kuhlotshaniswa kwakubaluleke kancane kuphela kumkawu u-S (ANOVA; imfene N: 1.03, p=0.46; inkawu S: F=1.73, p=0.04).Asibonanga mehluko ku-CP wama-engeli e-lattice phakathi kwama-lattice kunoma yisiphi isilwane (Fig. 6h; ANOVA; inkawu N: F = 1.8, p = 0.11; inkawu S: F = 0.32, p = 0. 9).
Okokugcina, umsebenzi wangaphambilini ubonise ukuthi i-CP iyashintsha phakathi novivinyo.Ezinye izifundo zibike ukwanda okubukhali okulandelwa umphumela wokukhetha obushelelezi, i-30 kanti abanye baye babika ukwanda okuqhubekayo kwesignali yokukhetha phakathi nesivivinyo31.Ngemfene ngayinye, sibale i-CP yeyunithi ngayinye esivivinyweni ngokungafani kombhalo okuyiziro (ngokulandelana, ngokuya ngomumo wephethini) kumaseli angu-100 ms anyathela njalo ngo-20 ms ukusuka ekuqaleni kwesivuseleli ukuze athumele asho ukususa ngaphambi kokuvuselela.I-avareji ye-CP dynamics yezinkawu ezimbili ikhonjiswe ku-Fig. 6i.Kuzo zombili izimo, i-CP yahlala isezingeni elingahleliwe noma isondelene kakhulu nayo kuze kube yilapho icishe ibe ngu-500 ms ngemva kokuqala kwesikhuthazo, ngemva kwalokho i-CP yanda kakhulu.
Ngokungeziwe ekushintsheni ukuzwela, i-CP iboniswe nokuthi ithintwa izimfanelo ezithile zezici zokushuna kwamaseli.Isibonelo, u-Uka no-DeAngelis34 bathole ukuthi i-CP emsebenzini wokuqaphela ukungafani kwebhinobholo kuncike ekufaneni kwejika lokushuna lokungafani kwebhinobhore yedivayisi.Kulesi simo, umbuzo ohlobene uthi ingabe amaseli e-Pattern direction selective (PDS) azwela kakhulu kunamaseli akhethiwe wengxenye (CDS).Amaseli e-PDS encode umumo ojwayelekile wamaphethini aqukethe umumo wendawo eminingi, kuyilapho amaseli e-CDS ephendula ukunyakaza kwezingxenye zephethini yokuqondisa (Fig. 7a).
ukumelwa okuhleliwe kwemodi yokushuna kwengxenye yokuvuselela kanye nokugawula okucatshangelwayo (kwesokunxele) kanye namajika okushuna ama-grating (kwesokudla) (bona Izinto Ezibalulekile Nezindlela).Ngamafuphi, uma iseli lihlanganisa izingxenye zegridi ukuze libonise ukunyakaza kwephethini, umuntu angalindela amajika afanayo okushuna egridi ngayinye kanye nesisusa segridi (ikholomu yokugcina, ijika eliqinile).Ngokuphambene, uma iseli lingazihlanganisi izikhombisi-ndlela zezingxenye ekunyakazeni kwephethini yesignali, umuntu angalindela ijika lokushuna le-bipartite elinenani eliphakeme ohlangothini ngalunye lokunyakaza kwe-grating okuhumusha ingxenye eyodwa iye endaweni ekhethwayo yeseli (ikholomu yokugcina, ijika elidayishiwe)..b (kwesokunxele) amajika okulungisa ukuqondiswa kwe-sinusoidal array yamaseli aboniswe ku-Figure 1 no-2. 3 no-4 (umugqa ophezulu - amaseli ku-Fig. 3a, b no-4a, b (phezulu); iphaneli engezansi - amaseli we-Fig. 3c, d no-4a, b (phansi)).(Phakathi) Izibikezelo zephethini nengxenye zibalwe kusukela kumaphrofayili wokushuna ama-lattice.(Kwesokudla) Ilungisa igridi yalawa maseli.Amaseli ephaneli ephezulu (phansi) ahlukaniswa njengamaseli esifanekiso (ingxenye).Qaphela ukuthi akukho ukuxhumana phakathi kwezigaba zezingxenye zephethini kanye nezintandokazi zokunyakaza kweseli okuhambisanayo/okusobala (bona izimpendulo ze-lattice yokuthungwa kwalawa maseli ku-Fig. 4a).c I-coefficient yokuhlanganisa ingxenye yemodi engu-z-score (ukudidiyela) ehlelwe ngokumelene ne-coefficient yokuhlobanisa ingxenye yengxenye engu-z-score (abscissa) yawo wonke amaseli arekhodwe kuzinkawu ezingu-N (kwesokunxele) kanye no-S (kwesokudla).Imigqa ewugqinsi ibonisa indlela yokubaluleka esetshenziselwa ukuhlukanisa amaseli.d Isakhiwo samathuba aphezulu okukhetha (okudidiyelwe) kuqhathaniswa nenkomba yemodi (Zp – Zc) (abscissa).Idatha kumaphaneli angakwesokunxele (kwesokudla) ibhekisela ku-monkey N(S).Imibuthano emnyama ibonisa idatha kumayunithi alinganiselwe.Kuzo zombili izilwane, kube nokuhlobana okubalulekile phakathi kwamathuba okukhethwa okuphezulu kanye nenkomba yephethini, okuphakamisa ukuhlobana okukhulu kokuqonda kwamaseli anomumo wephethini yesignali kusisusa esinomumo wezingxenye eziningi.
Ngakho-ke, kusethi yokuhlola ehlukile, silinganise izimpendulo kumagridi we-sinusoidal namagridi ukuze sihlukanise ama-neurons kumasampuli ethu njenge-PDS noma i-CDS (bona Izindlela).Amajika okushuna ama-lattice, izibikezelo zengxenye yesifanekiso ezakhiwe kusukela kule datha yokushuna, namajika okushuna ama-lattice amaseli aboniswe kuMfanekiso 1 no-3. Umfanekiso 3 no-4 kanye noMfanekiso 3 Owengeziwe kuboniswa kuMfanekiso 7b.Ukusatshalaliswa kwephethini nokukhetha kwengxenye, kanye nokuma kweseli okuncanyelwayo esigabeni ngasinye, kuboniswa inkawu ngayinye kuFig. 7c kanye nekhiwane elingeziwe.4 ngokulandelana.
Ukuhlola ukuncika kwe-CP ekulungisweni kwezingxenye zephethini, siqale sibala inkomba yephethini engu-35 (PI), amanani amakhulu (amancane) abonisa ukuziphatha okufanayo kwe-PDS (CDS) enkulu.Njengoba kunikezwe ukuboniswa okungenhla ukuthi: (i) ukuzwela kwe-neuronal kuyahlukahluka ngomehluko phakathi kokuma kweseli okuthandwayo kanye nesiqondiso sokunyakaza kwe-stimulus, futhi (ii) kukhona ukuhlobana okubalulekile phakathi kokuzwela kwe-neuronal nethuba lokukhetha kusampula yethu, sithole ubudlelwano phakathi kwe-PI ne-CP isiyonke kwafundelwa isiqondiso "esingcono kakhulu" sokunyakaza kweseli ngalinye (bona ngenhla). Sithole ukuthi i-CP yayihlotshaniswa kakhulu ne-PI (Umfanekiso 7d; ukuhlehla kwejiyomethri; inkawu enkulu ye-CP N: r = 0.23, p <0.01; inkawu ye-CP eqinile N r = 0.21, p = 0.013; imfene ye-CP enkulu S: r = 4 , 1-CP monkey S: r = 4 , 30 CP 0.29, p <10−3), okubonisa ukuthi amaseli ahlukaniswa njenge-PDS abonise umsebenzi omkhulu ohlobene nokukhethwa kune-CDS namaseli angahlukaniswanga. Sithole ukuthi i-CP yayihlotshaniswa kakhulu ne-PI (Umfanekiso 7d; ukuhlehla kwejiyomethri; inkawu enkulu ye-CP N: r = 0.23, p <0.01; inkawu ye-CP eqinile N r = 0.21, p = 0.013; imfene ye-CP enkulu S: r = 4 , 1-CP monkey S: r = 4 , 30 CP 0.29, p <10−3), okubonisa ukuthi amaseli ahlukaniswa njenge-PDS abonise umsebenzi omkhulu ohlobene nokukhethwa kune-CDS namaseli angahlukaniswanga. Мы обнаружили, что CP значительно коррелирует с PI (рис. 7d; регрессия среднего геометрического; большая обезьяна CP N:3, r0; 0,2, p = 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 ne-неклассифицированные клетки. Sithole ukuthi i-CP yayihlotshaniswa kakhulu ne-PI (Umfanekiso 7d; ukuhlehla kwejiyomethri; inkawu enkulu CP N: r = 0.23, p <0.01; inkawu ebistable CP N r = 0.21, p = 0.013; imfene enkulu CP S: r = 0.10: CP = 0.10 9, p <10-3), okubonisa ukuthi amaseli ahlukaniswa njenge-PDS abonise umsebenzi omningi, ohlotshaniswa nokukhetha kune-CDS namaseli angahlukaniswanga.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 = 猴0 CP: 猴0 CP = ; 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 和未分类的细类的分类的细类诚活性. 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 обезьяна CP N:3, r01; 0,2, p = зьяна CP N r = 0,21, p = 0,013; большая обезьяна CP S: r = 0,013) 0,30, p < 10-4; Sithole ukuthi i-CP yayihlotshaniswa kakhulu ne-PI (Umfanekiso 7d; ukuhlehla kwejiyomethri; inkawu enkulu i-CP N: r = 0.23, p <0.01; inkawu ebistable CP N r = 0.21, p = 0.013; imfene 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, прояотинский ть, чем клетки, классифицированные как CDS и неклассифицированные. monkey S bistable CP: r = 0.29, p <10-3), okubonisa ukuthi amaseli ahlukaniswa njenge-PDS abonise umsebenzi wokukhetha omkhulu kunamaseli ahlukaniswa njengama-CDS futhi awahlukaniswanga.Ngenxa yokuthi kokubili ukuzwela kwe-PI ne-neuron kuhlotshaniswa ne-CP, senze ukuhlaziya okuningilizayo kokuhlehla (nge-PI kanye nokuzwela kwe-neuron njengokuguquguqukayo okuzimele kanye ne-CP enkulu njengokuhlukahluka okuncikile) ukuze sikhiphe ukuhlobana phakathi kwezinyathelo ezimbili eziholela ekubeni nokwenzeka komphumela.. Womabili ama-coefficients okuxhumana ayingxenye abebalulekile (inkawu N: i-threshold vs. CP: r = −0.13, p = 0.04, PI vs. CP: r = 0.23, p < 0.01; imfene S: i-threshold vs. CP: r = −0 , p. 2, p. −3), okuphakamisa ukuthi i-CP inyuke ngokuzwela futhi ngendlela ezimele inyuke nge-PI. Womabili ama-coefficients okuxhumana ayingxenye abebalulekile (inkawu N: i-threshold vs. CP: r = −0.13, p = 0.04, PI vs. CP: r = 0.23, p < 0.01; imfene S: i-threshold vs. CP: r = −0 , p. 2, p. −3), okuphakamisa ukuthi i-CP inyuke ngokuzwela futhi ngendlela ezimele inyuke nge-PI. I-Obas частных коэффициента корреляции были значимыми (обезьяна N: порог против CP: r = -0,13, p = 0,04, PI против CP: 3, r 1 = 0, p; ротив CP: r = -0,16, p = 0,03, PI iqhudelana ne-CP: 0,29, p < 10-3), предполагая, что CP увеличивается с чувствительностью и независимзом обетский обетский обеличас обувствительностью и независимзом обетский обетский обегим. Womabili ama-coefficients okuxhumana ayingxenye abebalulekile (inkawu N: i-threshold vs. CP: r=-0.13, p=0.04, PI vs. CP: r=0.23, p<0.01; imfene S: i-threshold vs. CP: r = -0.16, p = 0.03, CP vs. CP-03: phakamisa. ngokuzwela futhi ikhuphuka ngokuzimela nge-PI.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: 阈值与 = ，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， Isithembiso чыеацх Кыыеы ыы п ко э кыы озьга s: против CP: R = -0,16, P = 0,03, PI о про тив Cp и ебьчилиныымы »нвзом. Womabili ama-coefficients wokuhlobanisa ingxenye abebalulekile (inkawu N: umkhawulo iqhudelana ne-CP: r=-0.13, p=0.04, PI vs. CP: r=0.23, p<0.01; imfene S: umkhawulo iqhudelana ne-CP: r = -0.16, p = 0.03, CPdic ku-CP. kwanda ngokuzwela futhi kwanda nge-PI ngendlela ezimele.
Sirekhode umsebenzi owodwa endaweni ye-MT, futhi izinkawu zabika umbono wazo wamaphethini angase abonakale njengokunyakaza okuhambisanayo noma okusobala.Ukuzwela kwama-neurons kuma-segment cues anezelwe emibonweni echemile kuyahluka kakhulu futhi kunqunywa, okungenani ngokwengxenye, ubudlelwano phakathi kokuma okuncanyelwayo kweyunithi kanye nesiqondiso somnyakazo wokuvuselela.Kuso sonke isibalo sabantu, ukuzwela kwe-neuronal bekuphansi kakhulu kunokuzwela kwengqondo, nakuba amayunithi azwela kakhulu afane noma eqe ukuzwela kokuziphatha kumasiginali wokuhlukanisa.Ngaphezu kwalokho, kukhona i-covariance ebalulekile phakathi kwemvamisa yokudubula kanye nokubona, okuphakamisa ukuthi ukusayina kwe-MT kudlala indima ekuhlukaniseni.Amaseli anomumo okhethwayo athuthukise ukuzwela kwawo kumehluko kumasiginali wesegimenti ye-lattice futhi athambekele ekuboniseni ukunyakaza komhlaba wonke ngesisusa esinezimo eziningi zendawo, okubonisa ukuhlobana okuphakeme kakhulu kokubona.Lapha sibheka ezinye izinkinga ezingase zibe khona ngaphambi kokuqhathanisa le miphumela nomsebenzi wangaphambilini.
Inkinga enkulu ngocwaningo olusebenzisa i-bistable stimuli kumamodeli ezilwane ukuthi izimpendulo zokuziphatha zingase zingasekelwe kubukhulu bentshisekelo.Isibonelo, izinkawu zethu zingabika umbono wazo womumo wokuthungwa ngaphandle kombono wazo wokuhlangana kwe-lattice.Izingxenye ezimbili zedatha ziphakamisa ukuthi lokhu akunjalo.Okokuqala, ngokuvumelana nemibiko yangaphambilini, ukushintsha i-engeli yokuma ehlobene yezingxenye zamalungu afanayo ahlukanisayo ngokuhlelekile kuguqule ithuba lokubona okuhambisanayo.Okwesibili, ngokwesilinganiso, umphumela uyafana kumaphethini aqukethe noma angaqukethe amasiginali wokuthungwa.Sekuhlanganiswe ndawonye, ​​lokhu kubheka kusikisela ukuthi izimpendulo zezinkawu zihlala zibonisa umbono wazo wokuxhumeka/okusobala.
Enye inkinga engaba khona ukuthi asikawathuthukisi kahle amapharamitha wokunyakaza kwe-grating esimweni esithile.Emisebenzini eminingi yangaphambilini eqhathanisa ukuzwela kwe-neuronal kanye ne-psychophysical, izisusa zakhethwa ngabanye kuyunithi ngayinye ebhalisiwe [31, 32, 34, 36, 37, 38, 39, 40, 41, 42, 43, 44, 45].Lapha sisebenzise izikhombisi-ndlela ezimbili ezifanayo zokuhamba kwephethini ye-lattice kungakhathaliseki ukulungiswa kokuma kweseli ngalinye.Lo mklamo usivumele ukuthi sifunde ukuthi ukuzwela kushintshe kanjani ngokugqagqana phakathi kokunyakaza kwe-lattice nokuma okuthandwayo, nokho, akuzange kunikeze isisekelo sokuqala sokunquma ukuthi amaseli akhetha ama-lattice ahambisanayo noma abonisa ngale.Ngakho-ke, sincika kumbandela wokusebenzisa amandla, sisebenzisa impendulo yeseli ngalinye kumanetha abhaliwe, ukunikeza okuthandwayo kanye namalebula anguziro esigabeni ngasinye sokunyakaza kwemeshi.Nakuba kungenakwenzeka, lokhu kungase kuhlanekezele ngokuhlelekile imiphumela yokuhlaziya kwethu ukuzwela nokutholwa kwesignali ye-CP, okungenzeka kulinganisele ngokweqile noma yisiphi isilinganiso.Kodwa-ke, izici eziningana zokuhlaziya kanye nedatha okuxoxwe ngayo ngezansi zibonisa ukuthi lokhu akunjalo.
Okokuqala, ukunikeza amagama ancanyelwayo (angenalutho) kusisusa esivuse umsebenzi omningi (ongaphansi) akuthinti ukuhlukaniswa kwalokhu kusatshalaliswa kwezimpendulo.Esikhundleni salokho, iqinisekisa kuphela ukuthi imisebenzi ye-neurometric kanye ne-psychometric inophawu olufanayo, ngakho-ke ingaqhathaniswa ngokuqondile.Okwesibili, izimpendulo ezisetshenziselwa ukubala i-CP (izilingo "ezingalungile" zegrated textured kanye nazo zonke izivivinyo zamagreyithi ngaphandle kokungafani kokwakheka) azizange zifakwe ekuhlaziyweni kokuhlehla okunquma ukuthi iseli ngalinye "likhetha" imidlalo exhunyiwe noma esobala.Lokhu kuqinisekisa ukuthi imiphumela yokukhetha ayichemi ekuqokweni okuthandwayo/okungavumelekile, okuholela emathubeni abalulekile okukhethwa.
Izifundo ze-Newsom nozakwabo [36, 39, 46, 47] zaba ngabokuqala ukunquma indima ye-MT ekulinganisweni okulinganiselwe kokuqondiswa kokunyakaza.Imibiko elandelayo iqoqe idatha yokubamba iqhaza kwe-MT ekujuleni34,44,48,49,50,51 kanye nesivinini32,52, umumo omuhle33 kanye nombono wesakhiwo se-3D kusukela ekuhambeni31,53,54 (3D amahlathi aqhubekayo).ukubusa).Sinweba le miphumela ngezindlela ezimbili ezibalulekile.Okokuqala, sinikeza ubufakazi bokuthi izimpendulo ze-MT zifaka isandla ekuhlukaniseni okubonwayo kwamasignali e-visuomotor.Okwesibili, sibone ubudlelwano phakathi kokukhetha kokuma kwemodi ye-MT kanye nalesi siginali yokukhetha.
Ngokomqondo, imiphumela yamanje ifana kakhulu nomsebenzi we-3-D SFM, njengoba yomibili iwumbono oyinkimbinkimbi we-bistable ohlanganisa ukunyakaza nokuhleleka kokujula.U-Dodd et al.31 bathole amathuba amakhulu okukhetha (0.67) kumsebenzi wenkawu obika umumo ojikelezayo wesilinda se-bistable 3D SFM.Sithole umphumela wokukhetha omncane kakhulu we-bistable grid stimuli (cishe u-0.55 kuzo zombili izinkawu).Njengoba ukuhlolwa kwe-CP kuncike ku-coefficient yokukhetha, kunzima ukuhumusha i-CP etholwe ngaphansi kwezimo ezahlukene emisebenzini ehlukene.Kodwa-ke, ubukhulu bomphumela wokukhetha esiwubonile ubufana kuziro nokuthungwa okuphansi kokugqama kwe gratings, nalapho sihlanganisa izisusa ezihlukile zokuthungwa okuphansi/akukho ukuze kwandiswe amandla.Ngakho-ke, lo mehluko ku-CP cishe ngeke ube ngenxa yomehluko kuzilinganiso zokukhetha phakathi kwamasethi edatha.
Izinguquko ezincane kuzinga lokudubula le-MT elihambisana nombono esimweni sakamuva zibonakala zidida uma kuqhathaniswa nezimo zokuqonda eziqinile nezihlukile ngenxa yokukhuthazwa kwe-3-D SFM kanye nezinhlaka zegridi ebhistable.Okunye okungenzeka ukuthi siwuthathe kancane umphumela wokukhetha ngokubala izinga lokudubula phakathi naso sonke isikhathi sesikhuthazo.Ngokuphambene necala le-31 3-D SFM, lapho umehluko emsebenzini we-MT uthuthuke cishe ku-250 ms ekuhlolweni futhi wabe usukhula kancane kancane kulo lonke uvivinyo, ukuhlaziya kwethu amandla esikhashana ezimpawu zokukhetha (bona i-500 ms ngemva kokuqala kokuvuselela kuzo zombili izinkawu. Ngaphezu kwalokho, ngemva kokukhuphuka okubukhali phakathi nalesi sikhathi sombiko, i-Rubin ihlale ibonwa ngesikhathi sokuhlolwa kwe-Hubin5 yomuntu ngamunye. ukwakheka kwamalungu afanayo angama-bistable angama-rectangular ngokuvamile kuyashintsha phakathi novivinyo olude. Nakuba ukuvuselela kwethu kwethulwe imizuzwana engu-1.5 kuphela, umbono wezinkawu zethu ungase uhluke kusukela ekuhlanganeni kuye obala phakathi novivinyo (izimpendulo zazo zibonise umbono wazo wokugcina ekukhetheni i-cue.) Ngakho-ke, inguqulo yesikhathi sokusabela yomsebenzi wethu, noma uhlelo lapho izinkawu zizoba khona umphumela olindelekile ukuthi umbono wabo ube MT ngokuqhubekayo. funda ngendlela ehlukile kule misebenzi emibili.Nakuba sekuyisikhathi eside kucatshangwa ukuthi amasignali e-CPU abangelwa ukuqoshwa kwezinzwa nomsindo ohlotshaniswayo,56 UGu kanye nozakwabo57 bathole ukuthi kumamodeli ezibalo, amasu ahlukene okuhlanganisa, kunamazinga okuhlukahluka okuhlobene, angakwazi ukuchaza kangcono i-CPU kuma-neuron e-dorsal media-superior temporal.ishidi lokushintsha i-orientation recognition task (MSTd).Umphumela omncane wokukhetha esiwubonile ku-MT cishe ubonisa ukuhlanganiswa okubanzi kwama-neuron amaningi anolwazi oluphansi ukuze kwakhe imibono yokuhambisana noma ukubeka izinto obala.Kunoma yikuphi lapho izinkomba zokunyakaza zendawo kufanele ziqoqwe zibe into eyodwa noma ezimbili (i-bistable gratings) noma izindawo ezihlukene zezinto ezivamile (3-D SFM), ubufakazi obuzimele bokuthi izimpendulo ze-MT zazihlotshaniswa kakhulu nezahlulelo zokuqonda, kwakukhona izimpendulo eziqinile ze-MT.kuhlongozwa ukuthi kudlale indima ekuhlukaniseni izithombe eziyinkimbinkimbi zibe izigcawu zezinto eziningi kusetshenziswa ulwazi lokunyakaza okubukwayo.
Njengoba kushiwo ngenhla, sibe ngabokuqala ukubika ukuhlangana phakathi komsebenzi weselula wephethini ye-MT kanye nokubona.Njengoba kwakhiwe kumodeli yasekuqaleni yezigaba ezimbili ngu-Movshon nozakwabo, iyunithi yemodi iyisigaba sokuphumayo se-MT.Nokho, umsebenzi wakamuva ubonise ukuthi imodi namaseli engxenye amelela iziphetho ezihlukene zokuqhubeka nokuthi umehluko wepharamitha esakhiweni senkambu yokwamukela ubophezelekile kububanzi bokushuna bezingxenye zemodi.Ngakho-ke, sithole ukuhlobana okubalulekile phakathi kwe-CP ne-PI, okufana nobudlelwano phakathi kokulinganisa kokungafani kokulinganisa kwebhinoli kanye ne-CP emsebenzini wokuqaphela ukujula noma ukulungiselelwa kwesilungiselelo sokuma emsebenzini wokubandlulula umumo omuhle.Ubudlelwano phakathi kwemibhalo ne-CP 33.U-Wang no-Movshon62 bahlaziye inani elikhulu lamaseli anokukhethwa kokuma kokuma kwe-MT futhi bathola ukuthi, ngokwesilinganiso, inkomba yemodi yayihlotshaniswa nezakhiwo eziningi zokulungisa, iphakamisa ukuthi ukukhetha kwemodi kukhona kwezinye izinhlobo eziningi zezimpawu ezingafundwa kubantu be-MT..Ngakho-ke, ezifundweni zesikhathi esizayo zobudlelwano phakathi komsebenzi we-MT kanye nombono oqondile, kuzobaluleka ukunquma ukuthi inkomba yephethini ihlobana ngokufanayo nomunye umsebenzi kanye namasiginali wokukhetha isikhuthazo, noma ukuthi lobu budlelwano buqondile yini esimweni sokuhlukaniswa kombono.
Ngokufanayo, i-Nienborg kanye ne-Cumming 42 bathola ukuthi nakuba amaseli aseduze nakude akhetha ukungafani kwebinocular ku-V2 ayezwela ngokulinganayo emsebenzini wobandlululo olujulile, isibalo samangqamuzana asondelene ancamelayo kuphela abonise i-CP ebalulekile.Kodwa-ke, ukuqeqesha kabusha izinkawu ekukhetheni umehluko okude wesisindo kubangele ama-CP abalulekile emakhejini athandwa kakhulu.Ezinye izifundo ziphinde zabika ukuthi umlando wokuqeqesha uncike ekuxhumaneni kombono34,40,63 noma ubudlelwano obuyimbangela phakathi komsebenzi we-MT nokubandlululwa okuhlukile48.Ubudlelwano esibubonile phakathi kwe-CP nokukhethwa kokuqondisa kwerejimeni kungenzeka kubonisa isu elithile izinkawu ezilisebenzise ukuxazulula inkinga yethu, hhayi indima ethile yamasiginali wokukhetha imodi ekuboneni kwemoto ebonakalayo.Emsebenzini wesikhathi esizayo, kuzobaluleka ukunquma ukuthi umlando wokufunda unomthelela obalulekile ekunqumeni ukuthi yimaphi amasiginali we-MT anesisindo esikhethekile futhi esivumelana nezimo ukuze enze izahlulelo zokuhlukaniswa.
U-Stoner nozakwabo14,23 babe ngabokuqala ukubika ukuthi ukuguqula ukukhanya kwezifunda zegridi ezeqayo kuthinte ngokubikezela ukuhambisana nokungafihli kwemibiko yezibukeli zomuntu kanye nokulungiswa kokuqondisa kuma-macaque MT neurons.Ababhali bathole ukuthi lapho ukukhanya kwezifunda ezigqagqene kufanelana nokubonakala obala, izibukeli zabika umbono osobala, kuyilapho ama-neurons e-MT ebonisa ukunyakaza kwezingxenye ze-raster.Ngokuphambene, lapho ukukhanya okugqagqene nokugqagqana okusobala kungqubuzana ngokomzimba, isibukeli sibona ukunyakaza okuhambisanayo, futhi ama-neuron e-MT abonisa ukunyakaza komhlaba wonke kwephethini.Ngakho-ke, lezi zifundo zibonisa ukuthi izinguquko ezingokomzimba ekushukumiseni okubonakalayo okuthonya ngokuthembekile imibiko yokuhlukaniswa futhi zenza izinguquko ezibikezelwe ekuvukeni kwe-MT.Umsebenzi wakamuva kule ndawo uhlole ukuthi yiziphi izimpawu ze-MT ezilandelela ukubukeka komqondo we-stimuli eyinkimbinkimbi18,24,64.Isibonelo, isethi engaphansi ye-MT neurons iye yaboniswa ukubonisa ukushuna kwe-bimodal kumephu enyakazayo yephoyinti engahleliwe (RDK) enezikhombisi-ndlela ezimbili ezihlukaniswe kancane kune-RDK engaqondile.Umkhawulokudonsa wokushuna kwamaselula 19, 25 .Izibukeli zihlala zibona iphethini yokuqala njengokunyakaza okusobala, nakuba ama-neuron amaningi e-MT ebonisa ukuguquguquka okungajwayelekile ekuphenduleni lezi zisusa, futhi isilinganiso esilula sawo wonke amaseli e-MT sinikeza impendulo yabantu abaningi.Ngakho-ke, isethi engaphansi yamaseli abonisa i-bimodal tuning ingase yakhe i-neural substrate yalo mbono.Kuyathakazelisa ukuthi kuma-marmosets, lesi sibalo sabantu sifane namaseli e-PDS lapho sihlolwa kusetshenziswa igridi evamile kanye nezisusa zegridi.
Imiphumela yethu ihamba ngesinyathelo esisodwa ngaphezu kwalokhu okungenhla, okubalulekile ekusunguleni indima ye-MT ekuhlukaniseni imibono.Eqinisweni, ukuhlukaniswa kuyisimo se-subjective.Izibonisi eziningi ze-polystable ezibukwayo zibonisa ikhono lesistimu yokubuka ukuhlela nokuhumusha izisusa eziphikelelayo ngezindlela ezingaphezu kweyodwa.Ukuqoqa ngesikhathi esifanayo izimpendulo ze-neural kanye nemibiko yokuqonda ocwaningweni lwethu kusivumele ukuthi sihlole ukuhlangana phakathi kwezinga lokudubula kwe-MT kanye nokuchazwa komqondo kokugqugquzelwa njalo.Njengoba sibonise lobu buhlobo, siyavuma ukuthi isiqondiso se-causality asikasungulwa, okungukuthi, ukuhlolwa okwengeziwe kuyadingeka ukuze kutholakale ukuthi isignali yokuhlukaniswa kombono obonwa yithi, njengoba abanye bephikisana [65, 66, 67], okuzenzakalelayo.Inqubo iphinde imele izimpawu ezehlayo ezibuyela ku-cortex yezinzwa ezivela ezindaweni eziphakeme 68, 69, 70 (Fig. 8).Imibiko yengxenye enkulu yamaseli akhetha iphethini ku-MSTd71, okukodwa kokuphokophelwe kwekhorotiki eyinhloko ye-MT, iphakamisa ukuthi ukunweba lokhu kuhlolwa ukuze kufake ukurekhodwa ngesikhathi esisodwa kwe-MT ne-MSTd kungaba isinyathelo sokuqala esihle ekuqondeni okwengeziwe izindlela ze-neural zokubona.ukuhlukaniswa.
Imodeli yezigaba ezimbili yengxenye kanye nemodi yokukhetha ukuma kwayo kanye nomthelela ongaba khona wempendulo eya phezulu emsebenzini ohlobene nokukhetha ekuhumusheni komshini.Lapha, ukukhetha kokuqondisa kwemodi (PDS – “P”) esinyathelweni se-MT kudalwe (i) isampula enkulu yedatha yokufaka ekhethiwe ehambisana nesivinini semodi ethile, kanye (ii) nokucindezela okuqinile kokushuna.Ingxenye ekhetha ngokuqondisa (CDS) yesiteji se-MT (“C”) inobubanzi besampula obuncane endaweni yokokufaka futhi ayinakho ukucindezelwa kokushuna okuningi.Ukuvinjelwa okungahleliwe kunikeza ukulawula kuzo zombili izigaba zabantu.Imicibisholo enemibala ibonisa umumo wedivayisi okhethwayo.Ukuze kucace, isethi encane kuphela yoxhumo lwe-V1-MT kanye nemodi yengxenye eyodwa nebhokisi lokukhetha umumo kukhonjiswa.Esimweni sokutolika imiphumela yethu yokuphakela phambili (FF), ukulungiselelwa okokufaka okubanzi kanye nokuvimbela ukushuna okuqinile (okugqanyiswe ngokubomvu) kumaseli e-PDS kubangele umehluko omkhulu emsebenzini ekuphenduleni amaphethini amaningi okunyakaza.Enkingeni yethu yokuhlukanisa, leli qembu liqhuba amaketanga ezinqumo futhi lihlanekezela umbono.Ngokuphambene, esimweni sempendulo (i-FB), izinqumo zokuqonda zenziwa kumasekethe akhuphukayo ngedatha yezinzwa kanye nokuchema kokuqonda, futhi umphumela omkhulu we-FB engezansi kumaseli e-PDS (imigqa ewugqinsi) ikhiqiza amasignali okukhetha.b Ukumelwa okuhleliwe kwamanye amamodeli emishini ye-CDS ne-PDS.Lapha amasignali e-PDS ku-MT akhiqizwa hhayi kuphela ngokufaka okuqondile kwe-V1, kodwa futhi ngokufaka okungaqondile kwendlela ye-V1-V2-MT.Izindlela ezingaqondile zemodeli ziyalungiswa ukuze zinikeze ukukhetha emingceleni yokuthungwa (izindawo ezigqagqene igridi).Imojuli ye-CDS yesendlalelo se-MT yenza isamba esinesisindo sokufakwayo okuqondile nokungaqondile futhi ithumele okukhiphayo kumojula ye-PDS.I-PDS ilawulwa ukuvinjelwa kokuncintisana.Futhi, lezo zixhumanisi kuphela eziboniswayo ezidingekayo ukudweba izakhiwo eziyisisekelo zemodeli.Lapha, indlela ye-FF ehlukile kunaleyo ehlongozwayo ku-a ingaholela ekuhlukeni okukhulu ekuphenduleni kwe-lattice yeselula ku-PDS, kuphinde kuholele ekuchemaneni emaphethini ezinqumo.Kungenjalo, i-CP enkulu kumaseli e-PDS ingase ibe umphumela wokuchema emandleni noma ekusebenzeni kahle kokunamathiselwe kwe-FB kumaseli e-PDS.Ubufakazi busekela amamodeli we-MT PDS wezigaba ezimbili nezintathu kanye nezincazelo ze-CP FF kanye ne-FB.
Ama-macaque amabili amadala (i-macaca mulatta), owesilisa oyedwa kanye nowesifazane oyedwa (iminyaka engu-7 kanye ne-5 ubudala ngokulandelana), anesisindo esisuka ku-4.5 kuya ku-9.0 kg, asetshenziswe njengezinto zokufunda.Ngaphambi kwakho konke ukuhlolwa kokuhlinzwa okungenanyumba, izilwane zafakwa negumbi lokuqopha elenziwe ngokwezifiso lama-electrode aqondile asondela endaweni ye-MT, indawo yokubeka ikhanda yensimbi engagqwali (Crist Instruments, Hagerstown, MD), kanye nokuma kweso ngekhoyili yokusesha elinganisiwe.(Cooner Wire, San Diego, California).Wonke amaphrothokholi ahambisana nemithetho yoMnyango Wezolimo wase-United States (i-USDA) kanye Neziqondiso Zezikhungo Zezempilo Zikazwelonke (NIH) Zokunakekelwa Kwabantu Nokusetshenziswa Kwezilwane Zaselabhorethri futhi zigunyazwe yi-University of Chicago Institutional Animal Care and Use Committee (IAUKC).
Zonke izisusa ezibukwayo zethulwe endaweni eyindilinga ngokumelene nengemuva elimnyama noma elimpunga.Ngesikhathi sokurekhoda, indawo kanye nobubanzi bale mbobo kwalungiswa ngokuhambisana nenkambu yakudala yokwamukela yama-neurons ethiphu ye-electrode.Sisebenzise izigaba ezimbili ezibanzi zezisusa ezibonwayo: izisusa ze-psychometric kanye nezisusa zokushuna.
I-psychometric stimulus iyiphethini yokugaya (20 cd / m2, 50% umehluko, 50% duty cycle, 5 degrees / sec) adalwe ngokuphakamisa ama-grating amabili angama-rectangular akhukhuleka aqonde ekuqondeni kwawo (Fig. 1b).Kuye kwaboniswa ngaphambilini ukuthi izibukeli ezingabantu zibona lawa maphethini egridi njengezisusa ze-bistable, ngezinye izikhathi njengephethini eyodwa ehamba ngendlela efanayo (ukunyakaza okuhambisanayo) futhi ngezinye izikhathi njengezindawo ezimbili ezihlukene ezihamba ngezindlela ezahlukene (ukunyakaza okusobala).izingxenye zephethini ye-lattice, i-oriented symmetrically - i-engeli phakathi kwama-lattices isuka ku-95 ° kuya ku-130 ° (ethathwe kusethi: 95 °, 100 °, 105 °, 115 °, 120 °, 125 °, 130 ° °, kuso sonke isikhathi seseshini Isolation 1 - ama-neurons agcinwe lapha, cishe ama-neurons angu-9 agcina lapha. ° noma 270° (iphethini yokuma).Esigabeni ngasinye, kusetshenziswe ikhoneni elilodwa kuphela le-interlattice lattice;phakathi neseshini ngayinye, ukuma kwephethini yesilingo ngasinye kukhethwe ngokungahleliwe emandleni amabili.
Ukuze singaqondi kahle umbono wegridi futhi sinikeze isisekelo sokuqina somvuzo wesenzo, sethula ukuthungwa kwephoyinti okungahleliwe kubha yokukhanya isinyathelo 72 sengxenye yegridi ngayinye.Lokhu kufinyelelwa ngokwandisa noma ukunciphisa (ngenani elinqunyiwe) ukukhanya kwesethi engaphansi ekhethwe ngokungahleliwe yamaphikseli (Fig. 1c).Isiqondiso sokunyakaza kokuthungwa kunikeza isignali eqinile egudluza umbono wombukeli ekunyakazeni okuhambisanayo noma okusobala (Fig. 1c).Ngaphansi kwezimo ezihambisanayo, yonke imidwebo, kungakhathaliseki ukuthi iyiphi ingxenye yesembozo se-texture lattice, ihunyushwa ngendlela yephethini (Fig. 1c, coherent).Esimeni esisobala, ukuthungwa kuhamba ngendlela eqondile ukuya ohlangothini lwe-grating elisimbozayo (Fig. 1c, transparent) (Supplementary Movie 1).Ukuze ulawule ubunkimbinkimbi bomsebenzi, kumaseshini amaningi umehluko we-Michelson (Lmax-Lmin/Lmax+Lmin) walolu phawu lokuthungwa beluhluka kusethi ethi (-80, -40, -20, -10, -5, 0, 5)., 10, 20, 40, 80).Ukugqama kuchazwa njengokukhanya okuhlobene kwe-raster (ngakho inani lokungafani elingu-80% lingabangela ukuthungwa okungu-36 noma 6 cd/m2).Kumaseshini angu-6 enkawuni engu-N kanye neseshini engu-5 ku-monkey S, sisebenzise ububanzi obuncane bokuqhathanisa bombhalo (-30, -20, -15, -10, -5, 0, 5, 10, 15, 20, 30), lapho izici ze-psychophysical zilandela iphethini efanayo njengokugqama kobubanzi obugcwele, kodwa ngaphandle kokugcwalisa.
Izinto ezishukumisayo zokushuna zingamagridi e-sinusoidal (qhathanisa no-50%, umjikelezo/idigri engu-1, amadigri angu-5/isekhondi) ehamba endaweni eyodwa kweziyi-16 ezinezikhala ngokulinganayo, noma amagridi e-sinusoidal ahamba ngalezi zindlela (okuhlanganisa ama-engeli amabili aphikisanayo angu-135° abekwe phezulu ama-sinusoidal gratings phezu komunye nomunye).ohlangothini olufanayo lwephethini.