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Umbono ogqibeleleyo ufuna isampuli entle ye-retina kunye nokuhlanganiswa ukuze kwakhiwe kwakhona iipropati zezinto. Kubalulekile ukuqaphela ukuba xa kuhlanganiswa iisampuli zasekuhlaleni ezivela kwizinto ezahlukeneyo, ukuchaneka kuyalahleka. Ke ngoko, ukwahlulwahlulwa, ukuhlelwa kweendawo zomfanekiso ukuze kucutshungulwe ngokwahlukeneyo, kubalulekile ekuqondeni. Kumsebenzi wangaphambili, izakhiwo ze-lattice ezi-bistable, ezinokuthathwa njengomphezulu omnye okanye ngaphezulu oshukumayo, zasetyenziswa ukufunda le nkqubo. Apha, sixela ubudlelwane phakathi kwezenzo kunye nokugweba ukwahlulwahlulwa kwiindawo eziphakathi zendlela yokubona ye-primate. Ngokukodwa, sifumanise ukuba ii-neurons ze-median temporal ezijolise ngokukhethayo zazinovelwano kwiindlela zokuthungwa ezisetyenziselwa ukugqwetha ukuqonda kwee-bistable gratings kwaye zibonise ulwalamano olubalulekileyo phakathi kwezilingo kunye nokuqonda okuzimeleyo kwezinto eziqhubekayo. Olu lwalamano lukhulu kwiiyunithi ezibonisa intshukumo yehlabathi kwiipateni ezineendlela ezininzi zasekuhlaleni. Ke ngoko, sigqiba kwelokuba indawo yexesha eliphakathi iqulethe imiqondiso esetyenziselwa ukwahlula izigcawu ezintsonkothileyo kwizinto kunye neendawo ezizimeleyo.
Umbono awuxhomekekanga kuphela kulwahlulo oluchanekileyo lweempawu zomfanekiso ezisisiseko ezifana nokujongwa komphetho kunye nesantya, kodwa okubaluleke ngakumbi ekudityanisweni ngokuchanekileyo kwezi mpawu ukuze kubalwe iipropati zokusingqongileyo ezifana nokuma kwento kunye nendlela elandelwayo1. Nangona kunjalo, iingxaki zivela xa imifanekiso ye-retina ixhasa amaqela amaninzi afanelekileyo ngokulinganayo 2, 3, 4 (Umzobo 1a). Umzekelo, xa iiseti ezimbini zemiqondiso yesantya zisondelelene kakhulu, oku kunokutolikwa ngokufanelekileyo njengento enye ehambayo okanye izinto ezininzi ezihambayo (Umzobo 1b). Oku kubonisa uhlobo oluzimeleyo lokwahlulwahlulwa, oko kukuthi, akusiyo ipropathi esisigxina yomfanekiso, kodwa yinkqubo yokutolika. Nangona ibalulekile ngokucacileyo kwimbono eqhelekileyo, ukuqonda kwethu isiseko se-neural sokwahlulwahlulwa kokuqonda akukapheleli kakuhle.
Umfanekiso wekhathuni wengxaki yokwahlulahlula kwembono. Umbono wombukeli ngobunzulu kwi-Necker cube (ekhohlo) uyatshintshana phakathi kweenkcazo ezimbini ezinokwenzeka (ekunene). Oku kungenxa yokuba akukho zimpawu kumfanekiso ezivumela ingqondo ukuba ichonge ngokukodwa indlela ebekwe ngayo umfanekiso kwi-3D (ebonelelwa yisignali yokuvaleka kwe-monocular ngasekunene). b Xa imiqondiso emininzi yokuhamba iboniswa kufutshane kwendawo, inkqubo yombono kufuneka imisele ukuba iisampulu zasekuhlaleni zivela kwinto enye okanye ezingaphezulu. Ukungacaci okukhoyo kwemiqondiso yokuhamba yendawo, oko kukuthi ulandelelwano lweentshukumo zezinto lunokuvelisa intshukumo efanayo yendawo, nto leyo ekhokelela kwiingcaciso ezininzi ezinokwenzeka ngokulinganayo zongeniso olubonakalayo, oko kukuthi amasimi evektha apha anokuvela kwintshukumo ehambelanayo yomphezulu omnye okanye intshukumo ecacileyo yeendawo ezigqumayo. c (ekhohlo) Umzekelo we-textured grid stimulus yethu. Ii-rectangular gratings ezirhuqa zijikeleze kwicala lazo ("izalathiso zecandelo" - iintolo ezimhlophe) ziyagqubana ukuze zenze ipateni ye-grating. I-lattice inokubonwa njengentshukumo enye, eqhelekileyo, edibeneyo yezikhokelo (iintolo ezibomvu) okanye intshukumo ecacileyo yezikhokelo ezidibeneyo. Umbono we-lattice ugqwethwa ngokongezwa kweempawu zokuthungwa kwenqaku elingacwangciswanga. (Embindini) Indawo egqanyiswe ngombala otyheli iyandiswa kwaye iboniswe njengoluhlu lweefreyimu zemiqondiso ehambelanayo necacileyo, ngokulandelelana. Intshukumo yechaphaza kwimeko nganye imelwe ziintolo eziluhlaza nezibomvu. (Ekunene) Igrafu yendawo (x, y) yendawo yokukhetha ngokuchasene nenani leefreyimu. Kwimeko ehambelanayo, zonke iitextures zihamba kwicala elinye. Kwimeko yokucaca, itexture ihamba kwicala lecandelo. d Umzobo wekhathuni womsebenzi wethu wokwahlulahlula intshukumo. Iinkawu zaqala uvavanyo ngalunye ngokulungisa ichaphaza elincinci. Emva kokulibaziseka okufutshane, uhlobo oluthile lwepateni yokuthungwa (ukuhambelana/ukucaca) kunye nobukhulu besignali yokuthungwa (umz. umahluko) kwavela kwindawo ye-MT RF. Ngexesha lovavanyo ngalunye, i-grating inokuthungwa kwenye yeendlela ezimbini ezinokwenzeka zepateni. Emva kokurhoxiswa kwe-stimulus, iithagethi zokukhetha zavela ngaphezulu nangaphantsi kwe-MT RF. Iinkawu kufuneka zibonise indlela ezibona ngayo igridi kwiisaccade ukuya kwindawo efanelekileyo yokukhetha.
Ukucubungula iintshukumo ezibonakalayo kuchazwe kakuhle kwaye ngaloo ndlela kubonelela ngemodeli egqwesileyo yokufunda iisekethe ze-neural zokwahlulwa kwe-perceptual. Izifundo ezininzi zekhompyutha ziphawule ukusetyenziswa kweemodeli zokucubungula intshukumo ezinamanqanaba amabini apho uqikelelo lokuqala olunesisombululo esiphezulu lulandelwa kukuhlanganiswa okukhethiweyo kweesampuli zasekuhlaleni ukususa ingxolo nokubuyisela isantya sento7,8. Kubalulekile ukuqaphela ukuba iinkqubo zokubona kufuneka ziqaphele ukunciphisa olu luhlu kuphela kwezo sampuli zasekuhlaleni ezivela kwizinto eziqhelekileyo. Izifundo ze-Psychophysical zichaze izinto zomzimba ezichaphazela indlela imiqondiso yentshukumo yasekuhlaleni eyahlulwahlulwa ngayo, kodwa imilo yeendlela ze-anatomical kunye neekhowudi ze-neural zihlala zivulekile imibuzo. Iingxelo ezininzi zibonisa ukuba iiseli ezikhetha ukujongwa kwindawo yexesha (MT) ye-primate cortex zizicelo ze-neural substrates.
Okubalulekileyo kukuba, kwezi zilingo zangaphambili, utshintsho kwimisebenzi ye-neural luhambelana notshintsho lomzimba kwi-visutical stimuli. Nangona kunjalo, njengoko kukhankanyiwe ngasentla, ukuhlukaniswa kwe-neural ngokuyintloko yinkqubo yokuqonda. Ke ngoko, uphononongo lwe-neural substrate yayo lufuna ukudibanisa utshintsho kwimisebenzi ye-neural kunye notshintsho kwindlela yokubona i-stimuli ezinzileyo. Ke ngoko, siqeqeshe iinkawu ezimbini ukuba zixele ukuba ipateni ye-bistable grating ebonwayo eyenziwe yi-superimposed drifting rectangular gratings yayingumphezulu omnye okanye iindawo ezimbini ezizimeleyo. Ukuze sifunde ulwalamano phakathi komsebenzi we-neural kunye nokugweba ukuhlukaniswa kwe-segmentation, sirekhode umsebenzi omnye kwi-MT xa iinkawu zenza lo msebenzi.
Sifumene ulwalamano olubalulekileyo phakathi kophando lomsebenzi we-MT kunye nokuqonda. Olu lwalamano belukho nokuba izikhuthazo ziqulathe iimpawu zokuhlukaniswa okucacileyo okanye akunjalo. Ukongeza, amandla esi siphumo anxulumene novakalelo kwiimpawu zokuhlukaniswa, kunye nesalathiso sepateni. Esi sibini silinganisa inqanaba apho iyunithi ikhupha khona intshukumo yehlabathi kunokuba ihambe kwindawo ethile kwiipateni ezinzima. Nangona ukukhetha kwicala lefashoni sele kuqatshelwe njengophawu oluchazayo lwe-MT, kwaye iiseli ezikhetha imfashini zibonisa ukulungiswa kwezikhuthazo ezinzima ezihambelana nombono womntu wezo zikhuthazo, ngokolwazi lwethu, obu bubungqina bokuqala bokunxibelelana phakathi kweepateni, isalathiso kunye nokwahlulwa kwembono.
Siqeqeshe iinkawu ezimbini ukubonisa indlela ezibona ngayo izinto ezishukumisa igridi eshukumayo (iintshukumo ezihambelanayo okanye ezicacileyo). Abantu ababukeleyo badla ngokuzibona ezi zinto zishukumisayo njengeentshukumo ezihambelanayo okanye ezicacileyo ezimalunga nesantya esifanayo. Ukuze sinike impendulo echanekileyo kolu vavanyo kwaye simise isiseko somvuzo osebenzayo, senze imiqondiso yokwahlulahlula ngokulungisa i-raster yecandelo elenza i-lattice (Umzobo 1c, d). Phantsi kweemeko ezihambelanayo, zonke izinto eziguqukayo zihamba kwicala lepateni (Umzobo 1c, “ezihambelanayo”). Kwimeko ecacileyo, izinto eziguqukayo zihamba ngokuthe nkqo kwicala le-grating apho zibekwe khona (Umzobo 1c, “ezicacileyo”). Silawula ubunzima bomsebenzi ngokutshintsha umahluko wale lebhile yento eguqukayo. Kwizilingo ezixutyiweyo, iinkawu zavuzwa ngeempendulo ezihambelana neempawu eziguqukayo, kwaye imivuzo yanikezelwa ngokungacwangciswanga (amathuba angama-50/50) kwizilingo eziqulethe iipateni ezingenazo iimpawu eziguqukayo (imeko yokungafani kwento eguqukayo).
Idatha yokuziphatha evela kuvavanyo olumeleyo olubini iboniswe kuMfanekiso 2a, kwaye iimpendulo zidweliswe njengenxalenye yezigqibo zokuvumelana ngokuchaseneyo nomahluko weempawu zokuthungwa (umahluko ocacileyo uthathwa njengongalunganga ngokwenkcazo) kwiipateni ezitshintshayo ziye phezulu okanye ezantsi, ngokulandelelana. Ngokubanzi, indlela ezibona ngayo iinkawu ngokuhambelana/ukucaca ichaphazeleke ngokuthembekileyo luphawu (olucacileyo, oluhambelanayo) kunye namandla (umahluko) lwe-texture cue (ANOVA; inkawu N: ulwalathiso – F = 0.58, p = 0.45, uphawu – F = 1248, p < 10−10, umahluko – F = 22.63, p < 10;−10 inkawu S: ulwalathiso – F = 0.41, p = 0.52, uphawu – F = 2876.7, p < 10−10, umahluko – F = 36.5, p < 10−10). Ngokubanzi, indlela ezibona ngayo iinkawu ngokuhambelana/ukucaca ichaphazeleke ngokuthembekileyo luphawu (olucacileyo, oluhambelanayo) kunye namandla (umahluko) lwe-texture cue (ANOVA; inkawu N: ulwalathiso – F = 0.58, p = 0.45, uphawu – F = 1248, p < 10−10, umahluko – F = 22.63, p < 10; −10 inkawu S: ulwalathiso – F = 0.41, p = 0.52, uphawu – F = 2876.7, p < 10−10, umahluko – F = 36.5, p < 10−10). В целом на восприятие обезьянами когерентности/прозрачности достоверно влияли как знак (прозрачность, когерентность), так истоверные влияли как знак (ANOVA; обезьяна N: направление — F = 0,58, p = 0,45, знак — F = 1248, p < 10−10, контраст – F = 22,63, p < 10; −10 обни, F p = 0,52, признак – F = 2876,7, p <10−10, контраст - F = 36,5, р <10-10). Ngokubanzi, imbono yokuhambelana/ukucaca kweenkawu ichaphazeleke kakhulu luphawu (ukucaca, ukunamathelana) kunye namandla (umahluko) ophawu lombhalo (ANOVA; inkawu N: ulwalathiso — F = 0.58, p = 0.45, uphawu — F = 1248, p < 10−10, umahluko – F = 22.63, p < 10; -10 inkawu S: ulwalathiso – F = 0.41, p = 0.52, uphawu – F = 2876.7, p < 10 −10, umahluko – F = 36.5, p < 10-10).总体而言，猴子对连贯性/透明度的感知受到纹理提示（ANOVA）的符号（透明、连贯）和强度（对比度）的可靠影响；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 猴子S: 方向– F = 0.41, p = 0.41, p = 0.41 0.52, 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（透明、连贯）和强度（对比度）的可靠影响；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 36.5,p <10-10）.Ngokubanzi, imbono yokuhambelana/ukucaca kwenkawu ichaphazeleke kakhulu luphawu (ukucaca, ukunamathelana) kunye nobunzulu (umahluko) beempawu 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: ulwalathiso – F = 0.58, p = 0.45, uphawu – F = 1248, p < 10−10, Umahluko – 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: Ukuqhelanisa – F = 0.41, p = 0.52, Uphawu – F = 2876.7, p < 10-10, Umahluko – F = 36.5, p < 10-10).Imisebenzi yokuqokelelana kweGaussian ifakelwe kwidatha evela kwiseshoni nganye ukuze ichaze iimpawu zengqondo zenkawu. Kumfanekiso 2b ubonisa ukusasazwa kwesivumelwano sale modeli kuzo zonke iiseshoni zeenkawu zombini. Ngokubanzi, iinkawu ziwugqibile umsebenzi ngokuchanekileyo nangokungaguquguqukiyo, kwaye sazigatya ngaphantsi kwe-13% zeeseshoni zeenkawu ezimbini ngenxa yokungalingani kakuhle kwimodeli yokuqokelelana yeGaussian.
a Imizekelo yokuziphatha kweenkawu kwiiseshoni ezimeleyo (n ​​≥ izilingo ezingama-20 ngemeko nganye yokukhuthaza). Kwiiphaneli zasekhohlo (ekunene), idatha evela kwiseshoni enye yenkawu ye-N(S) idwetshwe njengamanqaku okukhetha ahambelanayo (ordinate) ngokuchasene nomahluko weempawu zokuthungwa (abscissa). Apha kucingelwa ukuba ukuthungwa okucacileyo (okudibeneyo) kunexabiso elibi (elihle). Iimpendulo zakhiwe ngokwahlukeneyo ngokwendlela yokuhamba kwepateni (phezulu (90°) okanye ezantsi (270°)) kuvavanyo. Kuzo zombini izilwanyana, ukusebenza, nokuba impendulo yahlulwe yi-50/50 contrast (PSE - iintolo eziqinileyo) okanye ubungakanani bomahluko wokuthungwa okufunekayo ukuxhasa inqanaba elithile lokusebenza (umda - iintolo ezivulekileyo), zikwiindlela zokushukuma. b I-histogram efakiweyo yamaxabiso e-R2 omsebenzi we-Gaussian cumulative. Idatha ye-Monkey S(N) iboniswe ngasekhohlo (ekunene). c (Phezulu) I-PSE elinganisiweyo yegridi eshukunyiswe phantsi (i-ordinate) xa ithelekiswa ne-PSE eshukunyiswe phezulu kwigridi (i-abscissa) edwetshiweyo, kunye nemiphetho emele usasazo lwe-PSE kwimeko nganye kunye neentolo ezibonisa i-mean kwimeko nganye. Idatha yazo zonke iiseshoni ze-N(S) zenkawu zinikwe kwikholamu ekhohlo (ekunene). (Ezantsi) Isivumelwano esifanayo nesedatha ye-PSE, kodwa kwimida yeempawu ezifakelweyo. Akukho mahluko abalulekileyo kwimida ye-PSE okanye kwiindlela zefashoni (jonga umbhalo). d I-PSE kunye ne-slope (ordinate) zidwetshiwe ngokuxhomekeke kwi-raster orientation eqhelekileyo ye-angular separation component ("i-integral grating angle" - i-abscissa). Izangqa ezivulekileyo zezona ndlela, umgca oqinileyo yeyona modeli ifanelekileyo yohlengahlengiso, kwaye umgca onamachaphaza yi-95% confidence interval yemodeli yohlengahlengiso. Kukho ulwalamano olubalulekileyo phakathi kwe-PSE kunye ne-normalized integration angle, kodwa hayi i-slope kunye ne-normalized integration angle, nto leyo ebonisa ukuba umsebenzi we-psychometric uyatshintsha njengoko i-angle yahlula ii-lattices ze-component, kodwa ayiloli okanye i-flatting. (Iiseshoni zeMonkey N, n = iiseshoni ezingama-32; iMonkey S, n = iiseshoni ezingama-43). Kuzo zonke iiphaneli, iibha zempazamo zimele impazamo eqhelekileyo ye-mean. Haha. Ukuhambelana, amanqaku okulingana ngokwe-PSE, umgangatho oqhelekileyo.
Njengoko kuphawuliwe apha ngasentla, umahluko weempawu zokuthungwa kunye nendlela yokuhamba kwepateni yahlukile kuzo zonke iimvavanyo, kunye nezikhuthazo ezinyukayo okanye ezisezantsi kuvavanyo oluthile. Oku kwenzelwa ukunciphisa iziphumo eziguquguqukayo ze-psychophysical11 kunye ne-neuronal28. Ulwalathiso lwepateni ngokuchasene ne-bias (inqaku lokulingana elizimeleyo okanye i-PSE) (uvavanyo lwe-Wilcoxon rank sum; inkawu N: z = 0.25, p = 0.8; inkawu S: z = 0.86, p = 0.39) okanye umda womsebenzi ofanelekileyo (isimbuku see-Wilcoxon ranks; inkawu N: z = 0.14, p = 0.89, inkawu S: z = 0.49, p = 0.62) (Umzobo 2c). Ukongeza, bekungekho mahluko abalulekileyo phakathi kweenkawu kwinqanaba lokungafani kobume obufunekayo ukugcina amanqanaba omgangatho wokusebenza (inkawu i-N = 24.5% ± 3.9%, inkawu i-S = 18.9% ± 1.9%; i-Wilcoxon rank sum , z = 1.01, p = 0.31).
Kwiseshoni nganye, sitshintshe i-engile ye-interlattice eyahlula iindawo ezisetyenziswa kuzo ii-lattices ze-component. Izifundo ze-Psychophysical zibonise ukuba abantu banokubona iseli 10 njengedibeneyo xa le engile incinci. Ukuba iinkawu bezixela ngokuthembekileyo indlela ezibona ngayo ukuhambelana/ukucaca, ngoko ke ngokusekelwe kwezi ziphumo, umntu angalindela i-PSE, umahluko wobume obuhambelana nokwahlukana okufanayo phakathi kokuhambelana kunye nokukhetha okucacileyo, ukuba inyuke xa kusebenzisana. Oku bekunjalo ngokwenene (Umzobo 2d; ukudilika kwiindlela zepateni, iKruskal–Wallis; inkawu N: χ2 = 23.06, p < 10−3; inkawu S: χ2 = 22.22, p < 10−3; ulwalamano phakathi kwe-angle ye-inter-grating eqhelekileyo kunye ne-PSE – inkawu N: r = 0.67, p < 10−9; inkawu S: r = 0.76, p < 10−13). Oku bekunjalo ngokwenene (Umzobo 2d; ukudilika kwiindlela zepateni, iKruskal–Wallis; inkawu N: χ2 = 23.06, p < 10−3; inkawu S: χ2 = 22.22, p < 10−3; ulwalamano phakathi kwe-angle edibeneyo eqhelekileyo kunye ne-PSE – inkawu N: r = 0.67, p < 10−9; 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 <10:–3; 22, 22, p < 10–3 Oku kwenzeke ngokwenene (Umzobo 2d; ukuwohloka kwicala lephethini, iKruskal-Wallis; inkawu N: χ2 = 23.06, p < 10–3; inkawu S: χ2 = 22.22, p < 10–3; ulwalamano phakathi kwe-angle ye-lattice eqhelekileyo kunye ne-PSE – inkawu N: r = 0.67, p < 10-9, inkawu S: r = 0.76, p < 10-13).情况确实如此（图2d；跨模式方向折叠，Kruskal-Wallis；子子N:χ2 = 23.06,p <10-3猴子S:χ2 =p2. 10-3；标准化间光栅角和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 ;光栅角 和 pse-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; 10-3 нормализованный межрешеточный угол). Oku bekunjalo ngokwenene (Umzobo 2d; phinda ecaleni kwe-axis yemo, iKruskal-Wallis; inkawu N: χ2 = 23.06, p < 10-3; inkawu S: χ2 = 22.22, p < 10-3; ikona ye-interlattice eqhelekileyo). PSE-обезьяна N: r = 0,67, p < 10–9, обезьяна S: r = 0,76, p < 10–13). Inkawu yePSE N: r = 0.67, p < 10–9, inkawu S: r = 0.76, p < 10–13).Ngokwahlukileyo koko, ukutshintsha i-angle ye-interlattice akuzange kube nefuthe libalulekileyo kwi-slope yomsebenzi we-psychometric (Umzobo 2d; i-cross-modal orientation fold, iKruskal-Wallis; i-monkey N: χ2 = 8.09, p = 0.23; i-monkey S χ2 = 3.18, p = 0.67, ulwalamano phakathi kwe-angle ye-interlattice eqhelekileyo kunye ne-slope – i-monkey N: r = -0.4, p = 0.2, i-monkey S: r = 0.03, p = 0.76). Ngoko ke, ngokuhambelana nedatha yengqondo yomntu, isiphumo esiqhelekileyo sokutshintsha i-angle phakathi kwe-gratings kukutshintsha kwamanqaku okufuduka, kwaye akukokunyuka okanye ukwehla kovakalelo kwiimpawu zokwahlulwa.
Okokugqibela, imivuzo inikwa ngokungacwangciswanga kunye namathuba angama-0.5 kwizilingo ezine-zero texture contrast. Ukuba zonke iinkawu bezisazi ngale randomness ikhethekileyo kwaye zikwazile ukwahlula phakathi kwe-zero texture contrast kunye ne-cue stimuli, zinokuphuhlisa amaqhinga ahlukeneyo kwiintlobo ezimbini zezilingo. Iimbono ezimbini zibonisa ngamandla ukuba oku akunjalo. Okokuqala, ukutshintsha i-grating angle kunemiphumo efanayo ngokwemigangatho kwi-cue kunye ne-zero texture contrast scores (Umzobo 2d kunye nomzobo ongezelelweyo 1). Okwesibini, kuzo zombini iinkawu, ukhetho lwe-bistable trial alunakwenzeka ukuba lube luphinda-phindo lokhetho lwamva nje (lwangaphambili) lomvuzo (uvavanyo lwe-binomial, iinkawu ze-N: 0.52, z = 0.74, p = 0.22; iinkawu ze-S: 0.51, r = 0.9, p = 0.18).
Ukuqukumbela, indlela eziziphatha ngayo iinkawu kumsebenzi wethu wokwahlulahlula yayiphantsi kolawulo olufanelekileyo lokukhuthaza. Ukuxhomekeka kwezigqibo zokuqonda kwisibonakaliso kunye nobukhulu beempawu zokuthungwa, kunye notshintsho kwi-PSE ene-grating angle, kubonisa ukuba iinkawu zibike umbono wazo wokuhambelana kwemoto/ukucaca. Okokugqibela, iimpendulo zeenkawu kwizilingo zokungafani kothungo olungenanto azichaphazelekanga yimbali yomvuzo yezilingo zangaphambili kwaye zichaphazeleke kakhulu lutshintsho lwe-angular phakathi kwe-raster. Oku kuthetha ukuba iinkawu ziyaqhubeka nokuxela umbono wazo wobuqu wobume bomphezulu we-lattice phantsi kwale meko ibalulekileyo.
Njengoko kukhankanyiwe apha ngasentla, utshintsho lomahluko phakathi kwento ethile ukusuka kweyona imbi ukuya kweyona ilungileyo lulingana notshintsho olubonakalayo lwezisusa ukusuka kweyona icacileyo ukuya kweyona ihambelanayo. Ngokubanzi, kwiseli ethile, impendulo ye-MT idla ngokwanda okanye ukwehla njengoko umahluko phakathi kwento ethile utshintsha ukusuka kweyona ingalunganga ukuya kweyona ilungileyo, kwaye indlela yale mpembelelo idla ngokuxhomekeka kwicala lokuhamba kwepateni/inxalenye. Umzekelo, ii-directional tuning curves zeeseli ezimbini ze-MT ezimeleyo ziboniswe kuMfanekiso 3 kunye neempendulo zezi seli kwi-gratings eziqulethe imiqondiso ye-texture ecacileyo okanye ecacileyo engaphantsi okanye ephezulu. Sizame ngandlela thile ukulinganisa ngcono ezi mpendulo zegridi, ezinokunxulunyaniswa nokusebenza kwengqondo kweenkawu zethu.
Iploti ye-polar ye-directional tuning curve ye-monkey MT cell S emele i-monkey ephendula kwi-sinusoidal array enye. I-engile ibonisa indlela yokuhamba kwe-grating, ubukhulu bubonisa i-emissivity, kwaye indlela yeseli ekhethwayo idlula malunga ne-90° (phezulu) kunye necala lenye yeenxalenye kwicala le-grating pattern. b I-Weekly stimulus-time histogram (PSTH) yegridi yempendulo, itshintshelwe kwicala letemplate nge-90° (iboniswe ngokwesikimu ngasekhohlo) kwiseli eboniswe ku-a. Iimpendulo zihlelwa ngohlobo lwe-texture hint (i-cohesive/transparent – ​​i-middle/right panel ngokulandelelanayo) kunye ne-Michelson contrast (PSTH color hint). Kuboniswa imizamo echanekileyo kuphela kuhlobo ngalunye lweempawu ze-texture eziphantsi kunye ne-high-contrast. Iiseli ziphendule ngcono kwiipateni ze-lattice ezinyukayo ezine-transparent texture cues, kwaye impendulo kwezi pateni yanda ngokwanda kwe-texture contrast. c, d ziindlela ezifanayo njengakwi-a kunye ne-b, kodwa kwiiseli ze-MT ngaphandle kwe-monkey S, ukujongwa kwazo okuthandwayo phantse kudlula kwegridi ehlayo. Le yunithi ikhetha iigrating ezijonge ezantsi ezineempawu zokuthungwa ezihambelanayo, kwaye impendulo kwezi patheni iyanda ngokwanda kokungafani kokuthungwa. Kuzo zonke iiphaneli, indawo enomthunzi imele impazamo eqhelekileyo ye-mean. Iispokes. Iispikes, imizuzwana. yesibini.
Ukuze sihlolisise ubudlelwane phakathi koqwalaselo lomphezulu we-lattice (oluhambelanayo okanye olucacileyo) njengoko kuboniswe ziimpawu zethu zokuthungwa kunye nomsebenzi we-MT, siqale sabuyisela ulwalamano phakathi kweeseli kwintshukumo ehambelanayo (i-slope entle) okanye intshukumo ecacileyo (i-slope engalunganga) ngokubuyela umva. sinikwe ukwahlulahlula iiseli ngesantya sempendulo yesibonakaliso xa kuthelekiswa nomahluko (ngokwahlukileyo kwicala ngalinye lemo). Imizekelo yale migca yokulungisa i-lattice evela kwiseli yomzekelo ofanayo kuMfanekiso 3 iboniswe kuMfanekiso 4a. Emva kokwahlulahlula, sisebenzise uhlalutyo lokusebenza kwe-receiver (ROC) ukulinganisa uvakalelo lweseli nganye ekuguquleni imiqondiso yokuthungwa (jonga Iindlela). Imisebenzi ye-neurometric efunyenwe ngale ndlela inokuthelekiswa ngokuthe ngqo neempawu zengqondo ze-kawu kwiseshoni enye ukuze sithelekise ngokuthe ngqo uvakalelo lwe-psychophysical lwee-neurons kwi-lattice textures. Senze uhlalutyo lwe-signal find ezimbini kuzo zonke iiyunithi kwisampulu, sibala iimpawu ze-neurometric ezahlukeneyo kwicala ngalinye lepatheni (kwakhona, phezulu okanye ezantsi). Kubalulekile ukuqaphela ukuba, kolu hlalutyo, siquke kuphela iimvavanyo apho (i) izinto ezikhuthazayo zazine-texture cue kwaye (ii) iinkawu zaphendula ngokuhambelana naloo cue (oko kukuthi, uvavanyo "oluchanekileyo").
Amanqanaba omlilo acwangciswa ngokuchasene nomahluko weempawu zokwakheka, ngokulandelelana, kwiigratings ezitshintshayo (ekhohlo) okanye ezantsi (ekunene), umgca oqinileyo umele uhlengahlengiso oluchanekileyo oluhambelanayo, kwaye idatha ekumqolo ophezulu (osezantsi) ithathwe kwezo ziboniswe kwiFig. . Rice. 3a Cell, b (Umzobo 3c, d). Iimpawu ze-Regression slope zisetyenzisiwe ukunika ii-texture cues ezikhethwayo (ezihambelanayo/ezicacileyo) kwindibaniselwano nganye yokujonga iseli/i-lattice (n ≥ izilingo ezingama-20 ngemeko nganye yokukhuthaza). Iibha zempazamo zimele ukuphambuka okuqhelekileyo kwe-mean. Imisebenzi ye-neurometric yeeyunithi eziboniswe kwi-ba ichazwe kunye nemisebenzi ye-psychometric eqokelelwe ngexesha leseshoni enye. Ngoku, kwisici ngasinye, sidweba ukhetho lwe-tooltip olukhethwayo (ordinate) (jonga umbhalo) njengepesenti yomahluko we-texture's sign (abscissa). Umahluko we-texture utshintshiwe ukuze i-tooltips ezikhethwayo zibe ntle kwaye i-tooltips ezingenanto zibe negative. Idatha evela kwiigridi ezijikelezayo eziya phezulu (ezantsi) iboniswa kwiiphaneli zasekhohlo (ekunene), kwimigca ephezulu (esezantsi) - idatha evela kwiiseli eziboniswe kuMfanekiso 3a, b (Umfanekiso 3c, d). Iireyithi ze-neurometric kunye ne-psychometric threshold (N/P) ziboniswa kwiphaneli nganye. Ii-Spokes. Ii-Spikes, imizuzwana. i-sec, i-directory. ulwalathiso, iphondo elikhethwayo, i-psi. I-Psychometry, i-Neurology.
Ii-Lattice tuning curves kunye nemisebenzi ye-neurometric yeeseli ezimbini ze-MT ezimeleyo kunye nemisebenzi yazo ye-psychometric enxulumene nazo, zihlanganiswe kunye nezi mpendulo, ziboniswe kwiiphaneli eziphezulu nezisezantsi zeMfanekiso 4a, b, ngokwahlukeneyo. Ezi seli zibonisa ukwanda okanye ukwehla okungaqhelekanga njengoko isuntswana lokwakheka lisuka ekukhanyeni ukuya ekuhlanganeni. Ukongeza, ulwalathiso kunye namandla olu qhagamshelo luxhomekeke kwicala lentshukumo ye-lattice. Okokugqibela, imisebenzi ye-neurometric ebalwe kwiimpendulo zezi seli isondele kuphela (kodwa ayizange ihambelane) neepropati zengqondo zentshukumo yegridi engacwangciswanga. Zombini imisebenzi ye-neurometric kunye neye-psychometric ishwankathelwe ngemida, oko kukuthi ihambelana malunga ne-84% yomahluko okhethwe ngokuchanekileyo (ohambelana ne-mean + 1 sd yomsebenzi we-Gaussian ohlanganisiweyo). Kuyo yonke isampuli, umlinganiselo we-N/P, umlinganiselo womda we-neurometric ukuya kwi-psychometric, ube ngumyinge we-12.4 ± 1.2 kwi-monkey N kunye ne-15.9 ± 1.8 kwi-monkey S, kwaye ukuba i-lattice ihambe ubuncinci kwicala elinye, kuphela kwi-~16% (18). %) iiyunithi ezivela kwi-monkey N (monkey S) (Umzobo 5a). Kumzekelo weseli oboniswe kumfanekiso. Njengoko kubonwe kwiMifanekiso 3 kunye no-4, uvakalelo lwee-neurons lunokuchaphazeleka lulwalamano phakathi kolwalathiso olukhethwayo lweseli kunye nolwalathiso lwentshukumo ye-lattice esetyenzisiweyo kwizilingo. Ngokukodwa, ii-curve zohlengahlengiso lwe-orientation kwiMifanekiso 3a, c zibonisa ubudlelwane phakathi kokuseta i-neuron orientation ye-single sinusoidal array kunye novakalelo lwayo kwintshukumo ecacileyo/ehambelanayo kwi-textured array yethu. Oku bekunjalo kwiinkawu zombini (i-ANOVA; izikhokelo ezikhethwayo ezifakwe kwi-10° resolution; inkawu N: F = 2.12, p < 0.01; inkawu S: F = 2.01, p < 0.01). Oku bekunjalo kwiinkawu zombini (i-ANOVA; izikhokelo ezikhethwayo ezifakwe kwi-10° resolution; inkawu N: F = 2.12, p < 0.01; inkawu S: F = 2.01, p < 0.01). Это имело место для обеих обезьян (ANOVA; относительные предпочтительные направления объединены в группы с разрешениение, 12 °, p <0,01 обезьяна S: F = 2,01, p <0,01). Oku bekunjalo kwiinkawu zombini (i-ANOVA; izikhokelo ezikhethwayo eziqokelelwe kwisisombululo se-10°; inkawu N: F=2.12, p<0.01; inkawu S: F=2.01, p<0.01).两只猴子都是這种情况（方差分析；以10° 分辨率合并的相对首选方向；猴子N:F = 2.10=S = 2.10,p = 10; 0.01两 只 猴子 都 是 這 种 （方差 分析 以 10 ° 分辨率 合并 的 相对 方向 : : f = 2; f = 20. 2.01, p <0.01....... Это имело место для обеих обезьян (ANOVA; относительная предпочтительная ориентация объединена при разрешении 10°; обезрешении 10°; обезьяна S: F = 2,01, p <0,01). Oku bekunjalo kuzo zombini iinkawu (i-ANOVA; ukujongwa okuthandwayo okudityaniswe kwisisombululo se-10°; inkawu N: F=2.12, p<0.01; inkawu S: F=2.01, p<0.01).Ngenxa yokuguquguquka okukhulu kovakalelo lwe-neuron (Umzobo 5a), ukuze sibone ukuxhomekeka kovakalelo lwe-neuron kwiindawo ezikhethwayo, siqale salungisa indlela ekhethwayo yeseli nganye ukuya kwindawo "eyona" ilungileyo yokuhamba kwepateni yegridi (oko kukuthi, kwicala). apho i-grating yenza i-engile encinci phakathi kwendlela ekhethwayo yeseli kunye nendlela ekhethwayo yepateni ye-grating). Sifumanise ukuba imida ehambelanayo yee-neurons (umda wendlela "eyona" imbi kakhulu ye-lattice orientation/umda we-lattice orientation "eyona" ilungileyo) yahluka ngolu hlobo olukhethwayo oluqhelekileyo, kunye neencopho kulo mlinganiselo womda ezenzeka ngeenxa zonke zeepateni okanye iindawo ezikhethwayo (Umfanekiso 5b). )). Esi siphumo asikwazanga ukuchazwa kukucalucalulwa kwendlela ezikhethwayo kwiiyunithi kwisampulu nganye ukuya kwenye yeepateni zeplaid okanye iindlela zecandelo (Umzobo 5c; uvavanyo lweRayleigh; inkawu N: z = 8.33, p < 10−3, i-circular mean = 190.13 deg ± 9.83 deg; inkawu S: z = 0.79, p = 0.45) kwaye yayihambelana kuzo zonke ii-angles zeplaid inter-grating (Umzobo ongezelelweyo 2). Esi siphumo asinakuchazwa kukucalucalulwa kwendlela ezikhethwayo kwiiyunithi kwisampulu nganye ukuya kwenye yeepateni zeplaid okanye iindlela zecandelo (Umzobo 5c; uvavanyo lweRayleigh; inkawu N: z = 8.33, p < 10−3, i-circular mean = 190.13 deg ± 9.83 deg; inkawu S: z = 0.79, p = 0.45) kwaye yayihambelana kuzo zonke ii-angles zeplaid inter-grating (Umzobo ongezelelweyo 2). Этот эффект нельзя было объяснить смещением распределения предпочтительных направлений в единицах в каждой выборке здорке направлений или направлений компонентов (рис. 5в; критерий Рэлея; обезьяна N: z = 8,33, p < 10–3). Esi siphumo asinakuchazwa kukutshintsha kokusasazwa kweendlela ezikhethwayo kwiiyunithi kwisampulu nganye ukuya kwenye yeendlela ezitshekishwe okanye iindlela zecandelo (Umzobo 5c; uvavanyo lweRayleigh; inkawu N: z = 8.33, p < 10–3)., i-avareji ejikelezayo = 190.13 degrees ± 9.83 degrees; inkawu S: z = 0.79, p = 0.45) kwaye yayifana kuzo zonke iikona zegridi yeplaid (Umfanekiso Ongezelelweyo 2).這种效应不能通过每个样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释（图5c；瑞利测试；猴子N：z = 8.33,p <10-3 ，圆形平均值= 190.13 ± 9.83 ii-子S:z = 0.79，p = 0.45）并且在格子间光栅角上是一致的（补充图2）.7 （图 图 图瑞利测试 ；猴子 n : z = 8.33 , p <10-3 , 平均值 平均值 圆形 圆墢 圆墢圆形 umfanekiso weskrini 圆形 圆形 Этот эффект не может быть объяснен тем, что распределение предпочтительных ориентаций в клетках в каждом образце смеще решетки, либо в сторону одной из ориентаций компонентов (рис. 5в; критерий Рэлея; обезьяна N: z = 8,33, p <10–3). Esi siphumo asinakuchazwa kukuba ukusasazwa kweendlela ezikhethwayo kwiiseli kwisampuli nganye kuthambekele kwisakhiwo selatisi okanye kwenye yeendlela ezisetyenziswayo (Umzobo 5c; uvavanyo lukaRayleigh; inkawu N: z = 8.33, p < 10–3)., umndilili ojikelezayo) = 190.13 degrees ± 9.83 degrees; inkawu S: z = 0.79, p = 0.45) kwaye zazilingana kwii-engile ze-lattice phakathi kweegridi (Umzobo Ongezelelweyo 2).Ngoko ke, uvakalelo lwee-neurons kwiigridi ezithungelweyo luxhomekeke, ubuncinane ngokuyinxenye, kwiimpawu ezisisiseko ze-MT tuning.
Iphaneli yasekhohlo ibonisa usasazo lwee-N/P ratios (neuron/psychophysiological threshold); iseli nganye inika amanqaku amabini edatha, enye kwicala ngalinye apho ipateni ihamba khona. Iphaneli yasekunene ibonisa imida ye-psychophysical (ordinate) ngokuchasene nemida ye-neuronal (abscissa) yazo zonke iiyunithi kwisampulu. Idatha ekumqolo ophezulu (osezantsi) ivela kwinkawu N (S). b Imida yemida eqhelekileyo idwetshwe ngokuchasene nobukhulu bomahluko phakathi kolwalathiso lwe-lattice oluchanekileyo kunye nolwalathiso lweseli olukhethiweyo. Ulwalathiso "olungcono" luchazwa njengolwalathiso lwesakhiwo se-grating (olulinganiswa nge-sinusoidal grating enye) ekufutshane nolwalathiso lweseli olukhethiweyo. Idatha yaqala yahluzwa nge-normalized preferred orientation (10° bins), emva koko imida ye-threshold yahlengahlengiswa ukuya kwixabiso eliphezulu kwaye yalinganiswa ngaphakathi kwe-bin nganye. Iiseli ezine-orientation ekhethwayo enkulu kancinci okanye encinci kunolwalathiso lwe-lattice components zazinomahluko omkhulu kulwazelelo lolwalathiso lwe-lattice pattern. c I-Pink histogram yokusasazwa kwe-orientation ekhethwayo yazo zonke iiyunithi ze-MT ezirekhodwe kwinkawu nganye.
Okokugqibela, impendulo ye-MT ilawulwa yindlela ehamba ngayo intshukumo ye-grating kunye neenkcukacha zeempawu zethu zokwahlulwa (ubume). Ukuthelekiswa kobuntununtunu be-neuronal kunye ne-psychophysical kubonise ukuba, ngokubanzi, iiyunithi ze-MT bezingenabuntununtunu kangako kwizibonakaliso zokuthungwa ezichaseneyo kuneenkawu. Nangona kunjalo, ubuntununtunu be-neuron butshintshile ngokuxhomekeke kumahluko phakathi kwendlela ekhethwayo yeyunithi kunye nendlela ehamba ngayo intshukumo yegridi. Iiseli ezinobuntununtunu kakhulu bezisoloko zinezinketho zokuthotyelwa eziphantse zigubungele ipateni ye-lattice okanye enye yeendlela ezisetyenziswayo, kwaye icandelo elincinci leesampuli zethu lalinobuntununtunu okanye linobuntununtunu ngakumbi kunendlela iinkawu ezibona ngayo umahluko womahluko. Ukufumanisa ukuba iimpawu ezivela kwezi yunithi zinobuntununtunu zinxulumene ngakumbi nombono kwiinkawu, sihlolisise unxulumano phakathi kokuqonda kunye neempendulo ze-neuronal.
Inyathelo elibalulekileyo ekusekeni unxibelelwano phakathi komsebenzi we-neural kunye nokuziphatha kukumisela ulwalamano phakathi kwee-neurons kunye neempendulo zokuziphatha kwi-stimuli eziqhubekayo. Ukuze kuqhagamshelwe iimpendulo ze-neural kwizigqibo zokwahlulwahlulwa, kubalulekile ukudala i-stimuli ethi, nangona ifana, ibonwe ngokwahlukileyo kwizilingo ezahlukeneyo. Kolu phononongo lwangoku, oku kuboniswe ngokucacileyo yi-zero texture contrast grating. Nangona sigxininisa ukuba, ngokusekelwe kwimisebenzi ye-psychometric yezilwanyana, ii-gratings ezine-minimum (ngaphantsi kwe-~20%) textural contrast zihlala zithathwa njenge-coherent okanye ecacileyo.
Ukuze silinganisele ubungakanani beempendulo ze-MT ezihambelana ngayo neengxelo ze-perceptual, senze uhlalutyo lwe-selection probability (CP) lwedatha yethu yegridi (jonga 3). Ngamafutshane, i-CP sisilinganiselo esingesiso i-parametric, esingesiso isiqhelo esilinganisa ubudlelwane phakathi kweempendulo ze-spike kunye nezigwebo ze-perceptual30. Ukukhawulela uhlalutyo kwizilingo kusetyenziswa iigridi ezingenayo i-textural contrast kunye neeseshoni apho iinkawu zenze ubuncinane ukhetho oluhlanu kuhlobo ngalunye lwezi zilingo, sibale i-SR ngokwahlukileyo kwicala ngalinye lentshukumo yegridi. Kuzo zonke iinkawu, sibone ixabiso eliphakathi le-CP likhulu kakhulu kunokuba besinokulindela ngengozi (Umzobo 6a, d; inkawu N: i-mean CP: 0.54, 95% CI: (0.53, 0.56), uvavanyo lwe-t olunecala ezimbini ngokuchasene ne-null ye-CP = 0.5, t = 6.7, p < 10−9; inkawu S: i-mean CP: 0.55, 95% CI: (0.54, 0.57), uvavanyo lwe-t olunecala ezimbini, t = 9.4, p < 10−13). Kuzo zonke iinkawu, sibone ixabiso eliphakathi le-CP likhulu kakhulu kunokuba besinokulindela ngengozi (Umzobo 6a, d; inkawu N: i-mean CP: 0.54, 95% CI: (0.53, 0.56), uvavanyo lwe-t olunecala ezimbini ngokuchasene ne-null ye-CP = 0.5, t = 6.7, p < 10−9; inkawu S: i-mean CP: 0.55, 95% CI: (0.54, 0.57), uvavanyo lwe-t olunecala ezimbini, t = 9.4, p < 10−13).Kwiinkawu, sibone i-avareji ye-CP iphezulu kakhulu kunokuba bekulindelwe ngokungacwangciswanga (Umzobo 6a, d; inkawu N: i-avareji ye-CP: 0.54, 95% CI: (0.53, 0.56), uvavanyo lwe-t olunemisila emibini vs. amanani angenanto).CP = 0,5, t = 6,7, p <10-9; CP = 0.5, t = 6.7, p <10-9; обезьяна S: среднее CP: 0,55, 95% ДИ: (0,54, 0,57), двусторонний t-критерий, t = 9,4, p < 10–13) . inkawu S: umyinge we-CP: 0.55, 95% CI: (0.54, 0.57), uvavanyo lwe-t olunemisila emibini, 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: (0.53，0.56），针对空值的双边t 检验CP = 0.5, t = 6.7, p <10−9; 猴子S: 平均CP: 0.55, 05, 95%. I-双边t 检验, t = 9.4, p <10−13) .在 猴子 中 , 我們 观察 平均 平均 值 显着 大于 我們 偶然 的值0.53, 0.56） , 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: d: d; 0,54, 95% ДИ: (0,53, 0,56), двусторонний t- тест CP против нуля = 0,5, t = 6,7, p < 10-9, обезьяна S: средний CP5, И5, :0, 5 0,57), двусторонний t-критерий, t = 9,4, p < 10- 13). Kwiinkawu, sibone amaxabiso e-CP aphakathi kakhulu ngaphezu koko besinokukulindela ngengozi (Umzobo 6a, d; inkawu N: umyinge we-CP: 0.54, 95% CI: (0.53, 0.56), uvavanyo lwe-t olunemisila emibini vs. zero = 0.5, t = 6.7, p < 10-9, inkawu S: umyinge we-CP: 0.55, 95% CI: (0.54, 0.57), i-t-criterion enemisila emibini, t = 9.4, p < 10-13).Ngoko ke, ii-neurons ze-MT zidla ngokudubula ngamandla nangona kungekho zimpawu zokwahlulwa okucacileyo, xa indlela isilwanyana esibona ngayo intshukumo yelattice ihambelana nezinto ezikhethwa yiseli.
a Ukusasazwa kwamathuba okukhetha kwiigridi ngaphandle kwemiqondiso yokwakheka kweesampuli ezirekhodwe kwinkawu u-N. Iseli nganye inokufaka isandla ukuya kuthi ga kwiingongoma ezimbini zedatha (enye kwicala ngalinye lentshukumo yegridi). Ixabiso eliphakathi le-CP elingaphezulu kwe-random (iintolo ezimhlophe) libonisa ukuba ngokubanzi kukho ubudlelwane obubalulekileyo phakathi komsebenzi we-MT kunye nokuqonda. b Ukuhlola impembelelo yayo nayiphi na i-potential selection bias, sibalele i-CP ngokwahlukeneyo kuyo nayiphi na i-stimulus apho iinkawu zenze impazamo enye ubuncinane. Amathuba okukhetha adwetshwe njengomsebenzi womlinganiselo wokukhetha (pref/null) kuzo zonke i-stimuli (ekhohlo) kunye namaxabiso apheleleyo omahluko wophawu lokwakheka (ekunene, idatha evela kwiiseli ezili-120 ngazinye). Umgca oqinileyo kunye nendawo enombala kwiphaneli yasekhohlo imele i-mean ± sem yomndilili ohambayo wama-20-point. Amathuba okukhetha abalwe kwi-stimuli enemilinganiselo yokukhetha engalinganiyo, njengeegridi ezine-high signal contrast, zahluke ngakumbi kwaye zahlanganiswa malunga namathuba. Indawo enombala ongwevu kwiphaneli yasekunene igxininisa umahluko weempawu ezibandakanyiweyo ekubalweni kwamathuba okukhetha aphezulu. c Amathuba okukhetha okukhulu (i-ordinate) adwetshwe ngokuchasene nomda we-neuron (i-abscissa). Amathuba okukhetha adibene kakubi kakhulu nomda. I-convention df iyafana ne-ac kodwa isebenza kwi-157 idatha enye evela kwi-monkey S ngaphandle kokuba kuphawulwe ngenye indlela. g Amathuba okukhetha aphezulu (i-ordinate) adwetshwe ngokuchasene nolwalathiso olukhethwayo oluqhelekileyo (i-abscissa) kwinkawu nganye kwezi zimbini. Iseli nganye ye-MT inegalelo lamanqaku amabini edatha (enye kwicala ngalinye lesakhiwo se-lattice). h Iiploti ezinkulu zebhokisi zokhetho olunokwenzeka kwi-engile nganye ye-inter-raster. Umgca oqinileyo uphawula i-median, imiphetho esezantsi nephezulu yebhokisi imele i-25th kunye ne-75th percentile ngokulandelelana, ii-whiskers zolulelwe kwi-1.5 amaxesha kuluhlu lwe-interquartile, kwaye ii-outliers ezingaphezulu kwalo mda ziyaphawulwa. Idatha kwiiphaneli zasekhohlo (ekunene) ivela kwiiseli zenkawu ze-N(S) ezili-120 (157) zomntu ngamnye. i Amathuba aphezulu okukhetha (i-ordinate) adwetshwe ngokuchasene nexesha lokuqala kwe-stimulus (i-abscissa). I-CP enkulu ibalwe kwiingxande ezityibilikayo (ububanzi yi-100 ms, inyathelo yi-10 ms) kulo lonke uvavanyo yaze yajoliswa kwi-avareji kwiiyunithi.
Ezinye izifundo zangaphambili zibike ukuba i-CP ixhomekeke kwinani elihambelanayo lezilingo kulwabiwo lwesantya esisisiseko, oko kuthetha ukuba olu linganiselo aluthembekanga kangako kwi-stimuli ezibangela umahluko omkhulu kumlinganiselo wokhetho ngalunye. Ukuvavanya esi siphumo kwidatha yethu, sibale i-CP ngokwahlukeneyo kuzo zonke i-stimuli, nokuba i-sign texture contrast injani, kwaye iinkawu zenze ubuncinane uvavanyo olunye olungeyonyani. I-CP idwetshwe ngokuchasene nomlinganiselo wokukhetha (pref/null) kwisilwanyana ngasinye kuMfanekiso 6b kunye no-e (iphaneli yasekhohlo), ngokwahlukeneyo. Ukujonga i-avareji ehambayo, kuyacaca ukuba i-CP ihlala ingaphezulu kwe-probability kuluhlu olubanzi lwe-selection odds, iyancipha kuphela xa i-odds iwela (inyuka) ngaphantsi (ngasentla) 0.2 (0.8). Ngokusekelwe kwiimpawu ze-psychometric zezilwanyana, singalindela ukuba ii-select coefficients zalo bukhulu zisebenze kuphela kwi-stimuli ezine-high-contrast texture cues (ezihambelanayo okanye ezicacileyo) (jonga imizekelo yeempawu ze-psychometric kuMfanekiso 2a, b). Ukuze sifumanise ukuba oku bekunjalo na kwaye ingaba i-PC ebalulekileyo iqhubeke nokuba kukho izinto ezikhuthazayo ezineempawu ezicacileyo zokwahlulwahlulwa, sihlolisise isiphumo sexabiso elipheleleyo lokwahlulwahlulwa kombhalo kwi-PC (Umzobo 6b, e-right). Njengoko bekulindelekile, i-CP yayiphezulu kakhulu kunethuba lokuba izinto ezikhuthazayo zibe neempawu zokwahlulwahlulwa ezifikelela kwi-moderate (~20% yokwahlulwahlulwa okanye ezingaphantsi).
Kwimisebenzi yokuqondisa, isantya, kunye nokuqonda okungafaniyo, i-MT CP idla ngokuba yeyona iphezulu kwii-neurons ezibuthathaka kakhulu, mhlawumbi kuba ezi neurons zithwala imiqondiso enolwazi kakhulu30,32,33,34.Ngokuhambelana nezi ziphumo sibonile ulwalamano oluncinci kodwa olubalulekileyo phakathi kwe-grand CP, olubalwe ukusuka kwizinga lokudubula elino-z-scored kulo lonke ulwahlulo lwe-texture cue oluboniswe kwiphaneli esekunene yeFig.6b, e, kunye nomgca we-neuronal (Umzobo 6c, f; i-geometric mean regression; inkawu N: r = −0.12, p = 0.07 inkawu S: r = −0.18, p < 10−3). 6b, e, kunye nomgca we-neuronal (Umzobo 6c, f; i-geometric mean regression; inkawu N: r = −0.12, p = 0.07 inkawu S: r = −0.18, p < 10−3).Ngokuhambelana nezi ziphumo, sibone ulwalamano oluncinci kodwa olubalulekileyo phakathi kwe-CP enkulu ebalwe ukusuka kwi-excitation frequency z-score ukusuka kwi-texture signal contrast eboniswe kwipaneli esekunene ye-Fig. 6b, e, kunye ne-neuronal threshold (Fig. 6c, f; geometric). geometric mean regression;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和神经元阈值（图6c,f;於是這些 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 .图 6c, f ; 回归 ； 猴子 n : r = -0.12 , p = 0.07 子S:r = -0.18,p <10-3）.Ngokuhambelana nezi ziphumo, sibone ulwalamano oluncinci kodwa olubalulekileyo phakathi kwee-CV ezinkulu njengoko kubonisiwe kwiFig. 6b,e kunye nemida ye-neuron (Umfanekiso 6c,f; i-geometric mean regression; inkawu N: r = -0.12, p = 0.07).Обезьяна S: г = -0,18, р < 10-3). Inkawu S: r = -0.18, p < 10-3).Ke ngoko, izikhokelo ezivela kwiiyunithi ezinolwazi kakhulu zibonise ukuguquguquka okukhulu kunye nokugweba ukwahlulwahlulwa kweempawu kwiinkawu, nto leyo ibalulekileyo nokuba kukho naziphi na izikhokelo zombhalo ezongeziweyo kucalucalulo lokuqonda.
Ngenxa yokuba besikhe saseka ubudlelwane phakathi kokuva kwimiqondiso yegridi kunye nolwalathiso olukhethwayo lwe-neuronal, sizibuze ukuba ngaba kukho ubudlelwane obufanayo phakathi kwe-CP kunye nolwalathiso olukhethwayo (Umzobo 6g). Olu nxulumano lwalubaluleke kancinci kwinkawu S (ANOVA; inkawu N: 1.03, p=0.46; inkawu S: F=1.73, p=0.04). Asibonanga mahluko kwi-CP kwii-angles ze-lattice phakathi kwe-lattices nakwesiphi na isilwanyana (Umzobo 6h; ANOVA; inkawu N: F = 1.8, p = 0.11; inkawu S: F = 0.32, p = 0. 9).
Okokugqibela, umsebenzi wangaphambili ubonise ukuba i-CP iyatshintsha kulo lonke uvavanyo. Ezinye izifundo zibike ukunyuka okukhulu okulandelwa yimpembelelo yokukhetha egudileyo,30 ngelixa ezinye zibike ukunyuka okuzinzileyo kwisignali yokukhetha ngexesha lovavanyo31. Kwinkawu nganye, sibale i-CP yeyunithi nganye kwizilingo ezingenayo i-textural contrast (ngokulandelelana, ngokwendlela yokujonga ipateni) kwiiseli ze-100 ms sinyuka rhoqo emva kwe-20 ms ukusuka ekuqaleni kwe-pre-stimulus ukuya kwi-post mean pre-stimulus offset. I-avareji ye-CP dynamics yeenkawu ezimbini iboniswe kwiFig. 6i. Kuzo zombini iimeko, i-CP yahlala kwinqanaba elingacwangciswanga okanye ikufutshane kakhulu nayo de kwaba phantse yi-500 ms emva kokuqala kwe-stimulus, emva koko i-CP yanda kakhulu.
Ukongeza ekutshintsheni kobuntununtunu, i-CP ibonakalisiwe ukuba ichaphazeleka ziimpawu ezithile zeempawu zokulungisa iiseli. Umzekelo, u-Uka no-DeAngelis34 bafumanise ukuba i-CP kumsebenzi wokuqaphela ukungalingani kwe-binocular ixhomekeke kulungelelwaniso lwe-binocular mismatch tuning curve yesixhobo. Kule meko, umbuzo onxulumeneyo kukuba ingaba iiseli ezikhetha i-pattern direction (PDS) zinobuntununtunu ngakumbi kuneeseli ezikhetha i-component direction (CDS). Iiseli ze-PDS zibeka i-general orientation yeepateni eziqulethe i-local orientation ezininzi, ngelixa iiseli ze-CDS zisabela kwintshukumo yee-directional pattern components (Umzobo 7a).
ukubonakaliswa kweSchematic kwe-mode component tuning stimulus kunye ne-hypothetical grating (ekhohlo) kunye ne-grating orientation tuning curves (ekunene) (jonga izixhobo kunye neendlela). Ngamafutshane, ukuba iseli idibanisa kuzo zonke iinxalenye zegridi ukuze ishukume ipatheni yesignali, umntu angalindela ii-tuning curves ezifanayo zegridi nganye kunye ne-grid stimuli (ikholamu yokugqibela, i-solid curve). Ngokwahlukileyo koko, ukuba iseli ayihlanganisi imiyalelo yeenxalenye kwintshukumo yepatheni yesignali, umntu angalindela i-bipartite tuning curve enencopho kwicala ngalinye le-grating motion eguqulela icandelo elinye kwicala elikhethwayo leseli (ikholamu yokugqibela, i-dashed curve). . b (ekhohlo) ii-curves zokulungisa ulungelelwaniso lwe-sinusoidal array yeeseli eziboniswe kwiMifanekiso 1 kunye no-2. 3 kunye no-4 (umqolo ophezulu - iiseli kwiMifanekiso 3a,b kunye no-4a,b (phezulu); iphaneli esezantsi - iiseli zeMifanekiso 3c, d kunye no-4a, b (ezantsi)). (Ephakathi) Uqikelelo lwePateni kunye neenxalenye lubalwa kwiiprofayili ze-lattice tuning. (Ekunene) Ukulungisa igridi yezi seli. Iiseli zephaneli ephezulu (ezantsi) zahlulwe njengeeseli zetemplate (icandelo). Qaphela ukuba akukho nxibelelwano phakathi kohlulo lweenxalenye zepateni kunye nezinto ezikhethwayo zokuhamba kweeseli ezihambelanayo/ezicacileyo (jonga iimpendulo ze-texture lattice zezi seli kwiFig. 4a). c I-coefficient yolwalamano oluyinxenye yemo ye-z-score (i-ordinate) edwetshwe ngokuchasene ne-partial correlation coefficient ye-z-score component (i-abscissa) kuzo zonke iiseli ezirekhodwe kwiinkawu ze-N (ekhohlo) kunye ne-S (ekunene). Imigca engqindilili ibonisa iikhrayitheriya zokubaluleka ezisetyenziselwa ukwahlula iiseli. d Iploti ye-high selection probability (i-ordinate) ngokuchasene ne-mode index (Zp - Zc) (abscissa). Idatha kwiiphaneli zasekhohlo (ekunene) ibhekisa kwinkawu i-N(S). Izangqa ezimnyama zibonisa idatha kwiiyunithi eziqikelelweyo. Kuzo zombini izilwanyana, bekukho ulwalamano olubalulekileyo phakathi kwe-high selection probability kunye ne-pattern index, nto leyo ebonisa ulwalamano olukhulu lweeseli ezine-signal pattern orientation kwi-stimuli ene-multiple component orientation.
Ngoko ke, kwiseti yovavanyo eyahlukileyo, silinganise iimpendulo kwiigridi ze-sinusoidal kunye neegridi ukuze sihlele ii-neurons kwiisampulu zethu njenge-PDS okanye i-CDS (jonga iindlela). Ii-curve ze-Lattice tuning, uqikelelo lwe-template component olwakhiwe kule datha ye-tuning, kunye nee-curve ze-lattice tuning zeeseli eziboniswe kwiMifanekiso 1 kunye ne-3. Imifanekiso 3 kunye ne-4 kunye noMfanekiso oNcedisayo 3 ziboniswe kwiMifanekiso 7b. Ukusasazwa kweepateni kunye nokukhetha kwe-component, kunye nolwalathiso lweeseli olukhethwayo kudidi ngalunye, ziboniswa kwinkawu nganye kwiMifanekiso 7c kunye nomfanekiso oNcedisayo 4 ngokwahlukeneyo.
Ukuvavanya ukuxhomekeka kwe-CP ekulungisweni kwezixhobo zepateni, siqale sabala i-pattern index 35 (PI), ixabiso elikhulu (elincinci) elibonisa ukuziphatha okufanayo kwe-PDS (CDS) enkulu. Ngenxa yobungqina obungentla bokuba: (i) uvakalelo lwe-neuronal luyahluka ngokwahluka phakathi kolwalathiso lweseli olukhethwayo kunye necala lentshukumo ye-stimulus, kwaye (ii) kukho ulwalamano olubalulekileyo phakathi kovakalelo lwe-neuronal kunye namathuba okukhetha kwisampulu yethu, sifumene ubudlelwane phakathi kwe-PI kunye ne-CP iyonke efundwe kwicala "elilungileyo" lentshukumo yeseli nganye (jonga ngasentla). Sifumanise ukuba i-CP yayinxulumene kakhulu ne-PI (Umzobo 7d; i-geometric mean regression; i-grand CP monkey N: r = 0.23, p < 0.01; i-bi-stable CP monkey N = 0.21, p = 0.013; i-grand CP monkey S: r = 0.30, p < 10−4; i-bi-stable CP monkey S: r = 0.29, p < 10−3), nto leyo ebonisa ukuba iiseli ezihlelwe njenge-PDS zibonise umsebenzi omkhulu onxulumene nokukhetha kune-CDS kunye neeseli ezingahlelwanga. Sifumanise ukuba i-CP yayinxulumene kakhulu ne-PI (Umzobo 7d; i-geometric mean regression; i-grand CP monkey N: r = 0.23, p < 0.01; i-bi-stable CP monkey N = 0.21, p = 0.013; i-grand CP monkey S: r = 0.30, p < 10−4; i-bi-stable CP monkey S: r = 0.29, p < 10−3), nto leyo ebonisa ukuba iiseli ezihlelwe njenge-PDS zibonise umsebenzi omkhulu onxulumene nokukhetha kune-CDS kunye neeseli ezingahlelwanga. Мы обнаружили, что CP значительно коррелирует с PI (рис. 7d; регрессия среднего геометрического; большая обезьяна CP N:3, r = 0,20; бистабильная обезьяна CP N r = 0,21, p = 0,013 большая обезьяна CP S: r = 0,30, p < 10-4; 10-3), ndenze into enje, into enye, классифицированные как PDS, проявляли большую активность, связанную с выбором, чем CDS и неклассифицированные клетки. Sifumanise ukuba i-CP yayinxulumene kakhulu ne-PI (Umfanekiso 7d; i-geometric mean regression; i-great monkey CP N: r = 0.23, p < 0.01; i-bistable monkey CP N r = 0.21, p = 0.013; i-great monkey CP S: r = 0.30, p < 10-4; i-monkey S bistable CP: r = 0.29, p < 10-3), nto leyo ebonisa ukuba iiseli ezihlelwe njenge-PDS zibonise umsebenzi omninzi, onxulunyaniswa nokukhetha kune-CDS kunye neeseli ezingahlelwanga.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 = 3 CP = 3.02r CP = 0.02r S: 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 和未分类的细胞表现出更大的选择相关活性. CP 与PI 显着相关（图7d；几何平均回归；大CP猴N:r = 0.23，p <0.01；双稳态CP 猴N r = 0.20,r = 0.20; Мы обнаружили, что CP был значительно связан с PI (рис. 7d; регрессия среднего геометрического; большая обезьяна CP N:3, r0; 0,21; бистабильная обезьяна CP N r = 0,21, p = 0,013 большая обезьяна CP S: r = 0,013) 0,30, p < 10-4; Sifumanise ukuba i-CP yayinxulunyaniswa kakhulu ne-PI (Umfanekiso 7d; i-geometric mean regression; i-great monkey CP N: r = 0.23, p < 0.01; i-bistable monkey CP N r = 0.21, p = 0.013; i-great monkey CP S: r = 0.013) 0.30, p < 10-4; бистабильный CP обезьяны S: r = 0,29, p < 10-3), что указывает на то, что клетки, классифицированные как PDS, прояолунцьклишу активность, чем клетки, классифицированные как CDS и неклассифицированные. inkawu S bistable CP: r = 0.29, p < 10-3), nto leyo ebonisa ukuba iiseli ezihlelwe njengePDS zibonise umsebenzi omkhulu wokukhetha kuneeseli ezihlelwe njengeCDS kwaye ezingahlelwanga.Ngenxa yokuba zombini i-PI kunye novakalelo lwe-neuron zinxulumene ne-CP, senze uhlalutyo oluninzi lokubuyela umva (nge-PI kunye novakalelo lwe-neuron njengeenguqu ezizimeleyo kunye ne-CP enkulu njengenguqu exhomekeke kuyo) ukuze sisuse ulwalamano phakathi kwala manyathelo mabini akhokelela ekubeni kubekho isiphumo. Zombini ii-coefficients zolwalamano olungaphelelanga zazibalulekile (i-monkey N: threshold vs. CP: r = −0.13, p = 0.04, PI vs. CP: r = 0.23, p < 0.01; inkawu S: threshold vs. CP: r = −0.16, p = 0.03, PI vs CP: 0.29, p < 10−3), nto leyo ebonisa ukuba i-CP iyanda ngovakalelo kwaye ngendlela ezimeleyo iyanda nge-PI. Zombini ii-coefficients zolwalamano olungaphelelanga zazibalulekile (i-monkey N: threshold vs. CP: r = −0.13, p = 0.04, PI vs. CP: r = 0.23, p < 0.01; inkawu S: threshold vs. CP: r = −0.16, p = 0.03, PI vs CP: 0.29, p < 10−3), nto leyo ebonisa ukuba i-CP iyanda ngovakalelo kwaye ngendlela ezimeleyo iyanda nge-PI. Inkcazelo ngokuthe gabalala CP: r = -0,13, p = 0,04, PI против CP: 3, r = 0, 2, p = 0, 2 порог против CP: r = -0,16, p = 0,03, PI vs CP: 0,29, p < 10-3), предполагая, что CP увеличивается с чувствительностью и неваомбисть PI. Zombini ii-coefficients zolwalamano olungaphelelanga zazibalulekile (i-monkey N: threshold vs. CP: r=-0.13, p=0.04, PI vs. CP: r=0.23, p<0.01; inkawu S: threshold vs. CP: r = -0.16, p = 0.03, PI vs CP: 0.29, p <10-3), nto leyo ebonisa ukuba i-CP iyanda ngovakalelo kwaye iyanda 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 = 阈倎倎 = 阈 CP: S = 6; 0.03,PI vs CP:0.29,p <10-3），表明CP 随灵敏度增加而增加，并且以独立方式随PI 增加。两个偏相关系数均显着（猴子N:阈值与CP:r = -0.13,p = 0.04，PI = 0.03，PI vs CP:0.29，p <10-3明， Inkcazelo ngokuthe gabalala CP: r = -0,13, p = 0,04, PI против CP: 3, r = 0, 2, p = 0, 2 порог против CP: r = -0,16, p = 0,03 , PI против CP: 0,29, p < 10-3), что указывает на то, что CP увеличивалась с чувстевительность независимым образом. Zombini ii-coefficients zolwalamano olungaphelelanga zazibalulekile (i-monkey N: threshold vs. CP: r=-0.13, p=0.04, PI vs. CP: r=0.23, p<0.01; inkawu S: threshold vs. CP: r = -0.16, p = 0.03, PI vs CP: 0.29, p <10-3), nto leyo ebonisa ukuba i-CP inyukile ngovakalelo kwaye inyukile nge-PI ngendlela ezimeleyo.
Sirekhode umsebenzi omnye kwindawo ye-MT, kwaye iinkawu zibike indlela ezibona ngayo iipateni ezinokubonakala njengeentshukumo ezihambelanayo okanye ezicacileyo. Uvakalelo lwee-neurons kwiimpawu zokwahlulwahlulwa ezongezwe kwiimbono ezicalucalulo luyahluka kakhulu kwaye lumiselwa, ubuncinane ngokuyinxenye, lulwalamano phakathi kolwalathiso olukhethwayo lweyunithi kunye necala lentshukumo yokukhuthaza. Kubo bonke abantu, uvakalelo lwe-neuronal lwaluphantsi kakhulu kunovakalelo lwe-psychophysical, nangona iiyunithi ezinovakalelo kakhulu zifanise okanye zidlule uvakalelo lokuziphatha kwiimpawu zokwahlulwahlulwa. Ukongeza, kukho ukuhambelana okubalulekileyo phakathi kokuphindaphinda kokudubula kunye nokuqonda, okubonisa ukuba i-MT signaling idlala indima ekuhlukaniseni. Iiseli ezine-orientation ekhethwayo ziphucule uvakalelo lwazo kumahluko kwizibonakaliso zokwahlulwahlulwa kwe-lattice kwaye zithande ukubonisa intshukumo yehlabathi kwi-stimuli nge-orientation ezininzi zasekuhlaleni, zibonisa ulwalamano oluphezulu lokuqonda. Apha siqwalasela ezinye iingxaki ezinokubakho ngaphambi kokuthelekisa ezi ziphumo nomsebenzi wangaphambili.
Ingxaki enkulu ngophando olusebenzisa izinto ezibangela ukuba umntu angakwazi ukuzibamba kwiimodeli zezilwanyana kukuba iimpendulo zokuziphatha azinakusekelwa kubungakanani bomdla. Umzekelo, iinkawu zethu zinokuxela indlela ezibona ngayo indlela ezibona ngayo izinto ngokuzimeleyo ngendlela ezibona ngayo ukuhambelana kwelattice. Iinkalo ezimbini zedatha zibonisa ukuba oku akunjalo. Okokuqala, ngokuhambelana neengxelo zangaphambili, ukutshintsha i-angle yokujonga enxulumeneyo yezinto ezihlukanisayo kutshintshe ngokucwangcisiweyo amathuba okuqonda okuvumelanayo. Okwesibini, ngokomyinge, isiphumo siyafana kwiipateni eziqulathe okanye ezingenazo izibonakaliso zokuthungwa. Xa zizonke, ezi mbono zibonisa ukuba iimpendulo zeenkawu zihlala zibonisa indlela ezibona ngayo ukunxibelelana/ukucaca.
Enye ingxaki enokubakho kukuba asikayilungisi kakuhle iparameters yentshukumo ye-grating kwimeko ethile. Kwimisebenzi emininzi yangaphambili ethelekisa uvakalelo lwe-neuronal kunye nolwe-psychophysical, izikhuthazo zikhethwe ngokwahlukeneyo kwiyunithi nganye ebhalisiweyo [31, 32, 34, 36, 37, 38, 39, 40, 41, 42, 43, 44, 45]. Apha sisebenzise iindlela ezimbini ezifanayo zokuhamba kwepateni ye-lattice nokuba ulungelelwaniso lolwalathiso lweseli nganye lunjani. Olu yilo lusivumele ukuba sifunde indlela uvakalelo olutshintsha ngayo ngokudibana phakathi kwentshukumo ye-lattice kunye nolwalathiso olukhethwayo, nangona kunjalo, aluzange lubonelele ngesiseko sokuqala sokumisela ukuba iiseli zikhetha ii-lattice ezihambelanayo okanye ezicacileyo. Ke ngoko, sixhomekeke kwiikhrayitheriya zovavanyo, sisebenzisa impendulo yeseli nganye kwi-mesh ebhalwe ngendlela ethile, ukunika ukhetho kunye neelebhile ezingezizo kudidi ngalunye lwentshukumo ye-mesh. Nangona kungenakwenzeka, oku kunokujika ngokucwangcisiweyo iziphumo zohlalutyo lwethu lovakalelo kunye nokufunyanwa kwesignali ye-CP, okunokuthi kugqithise nakuphi na umlinganiselo. Nangona kunjalo, iinkalo ezininzi zohlalutyo kunye nedatha exutyushwayo ngezantsi zibonisa ukuba oku akunjalo.
Okokuqala, ukunika amagama akhethwayo (angenanto) kwizikhuthazo ezibangele umsebenzi omninzi (ongaphantsi) akuzange kuchaphazele ukwahlulwa kwale miba yokusabela. Endaweni yoko, kuqinisekisa kuphela ukuba imisebenzi ye-neurometric kunye neye-psychometric inophawu olufanayo, ukuze ithelekiswe ngokuthe ngqo. Okwesibini, iimpendulo ezisetyenziselwa ukubala i-CP (izilingo "ezingalunganga" ze-textured gratings kunye nazo zonke izilingo ze-textured gratings ngaphandle kokungafani kwe-texture) azifakwanga kuhlalutyo lokubuyela umva oluye lwamisela ukuba iseli nganye "ikhetha" imidlalo edibeneyo okanye ecacileyo. Oku kuqinisekisa ukuba iziphumo zokukhetha azikhethi cala kwiindlela ezikhethwayo/ezingasebenziyo, nto leyo ekhokelela kumathuba amakhulu okukhetha.
Izifundo zikaNewsom kunye noogxa bakhe [36, 39, 46, 47] zezokuqala ukumisela indima ye-MT ekuqikeleleni okuqikelelweyo kwendlela yokuhamba. Iingxelo ezilandelayo ziqokelele idatha malunga nokubandakanyeka kwe-MT kubunzulu34,44,48,49,50,51 kunye nesantya32,52, ulwalathiso oluhle33 kunye nokuqonda ulwakhiwo lwe-3D ukusuka kwintshukumo31,53,54 (amahlathi azinzileyo e-3D). ukuya kulawula). Siyandisa ezi ziphumo ngeendlela ezimbini ezibalulekileyo. Okokuqala, sinika ubungqina bokuba iimpendulo ze-MT zinegalelo ekuhlukaniseni imiqondiso ye-visuomotor. Okwesibini, sibonile ubudlelwane phakathi kokukhetha ukujongwa kwendlela ye-MT kunye nalo mqondiso wokukhetha.
Ngokwengqiqo, iziphumo zangoku zifana kakhulu nomsebenzi kwi-3-D SFM, njengoko zombini ziyimbono enzima ye-bistable ebandakanya intshukumo kunye nokulandelelana kobunzulu. UDodd et al.31 bafumene ithuba elikhulu lokukhetha (0.67) kumsebenzi wenkawu bexela ukujongwa kokujikeleza kwesilinda ye-bistable 3D SFM. Sifumene isiphumo sokukhetha esincinci kakhulu kwi-bistable grid stimuli (malunga ne-0.55 kwiinkawu zombini). Ekubeni uvavanyo lwe-CP luxhomekeke kwi-coefficient yokukhetha, kunzima ukutolika i-CP efunyenwe phantsi kweemeko ezahlukeneyo kwimisebenzi eyahlukeneyo. Nangona kunjalo, ubukhulu besiphumo sokukhetha esisibonileyo babufana kwi-zero kunye ne-low texture contrast gratings, kwaye naxa sidibanise i-low/no texture contrast stimuli yokwandisa amandla. Ke ngoko, lo mahluko kwi-CP akunakwenzeka ukuba ubangelwe kumahluko kumazinga okukhetha phakathi kweeseti zedatha.
Utshintsho oluncinci kwizinga lokudubula le-MT oluhamba nombono kwimeko yokugqibela lubonakala luyingxaki xa kuthelekiswa neemeko zokuqonda ezinzulu nezingafaniyo ngokwemigangatho ezibangelwa kukukhuthazwa kwe-3-D SFM kunye nezakhiwo zegridi eziguquguqukayo. Enye into enokwenzeka kukuba siwuthathe kancinci umphumo wokukhetha ngokubala izinga lokudubula kulo lonke ixesha lokukhuthazwa. Ngokwahlukileyo kwimeko ye-31 3-D SFM, apho umahluko kwimisebenzi ye-MT uphuhliswe malunga ne-250 ms kwizilingo waza wanda ngokuthe chu kuzo zonke iimvavanyo, uhlalutyo lwethu lwe-temporal dynamics yesignali zokukhetha (jonga i-500 ms emva kokuqala kwe-stimulus kwiinkawu zombini. Ukongeza, emva kokunyuka okukhulu ngeli xesha, sibone ukuguquguquka kwi-CP ngexesha eliseleyo lovavanyo. UHupe noRubin55 baxela ukuba imbono yabantu ngee-bistable rectangular arrays zihlala zitshintsha ngexesha lovavanyo olude. Nangona i-stimuli zethu zaziswa imizuzwana eyi-1.5 kuphela, imbono yeenkawu zethu inokwahluka ukusuka ekuhambelaneni ukuya ekukhanyeni ngexesha lovavanyo (iimpendulo zabo zabonisa imbono yabo yokugqibela ekukhetheni i-cue.) Ke ngoko, inguqulelo yexesha lokuphendula yomsebenzi wethu, okanye isicwangciso apho iinkawu zinokubika khona imbono yazo ngokuqhubekayo, kulindeleke ukuba zibe nefuthe elikhulu lokukhetha. Inokwenzeka yokugqibela kukuba isibonakaliso se-MT sifundwa ngokwahlukileyo kwimisebenzi emibini. Nangona kudala kucingwa ukuba imiqondiso ye-CPU ivela kwi-sensory decoding kunye nengxolo edibeneyo,56 UGu kunye noogxa bakhe57 bafumanise ukuba kwiimodeli zokubala, amaqhinga ahlukeneyo okudibanisa, endaweni yamanqanaba e-correlated ukuguquguquka, kunokuchaza ngcono i-CPU kwi-dorsal media-superior temporal neurons. umsebenzi wokuqaphela utshintsho lweshiti (MSTd). Isiphumo sokukhetha esincinci esisibonileyo kwi-MT mhlawumbi sibonisa ukuhlanganiswa okubanzi kwee-neurons ezininzi ezinolwazi oluphantsi ukudala imbono yokuhambelana okanye ukucaca. Nokuba yeyiphi na imeko apho ii-local motion cues zaza kuhlanganiswa zibe yinto enye okanye ezimbini (ii-bistable gratings) okanye iindawo ezahlukeneyo zezinto eziqhelekileyo (3-D SFM), ubungqina obuzimeleyo bokuba iimpendulo ze-MT zazinxulunyaniswa kakhulu nokugweba okubonakalayo, bekukho iimpendulo ze-MT ezinamandla. Kucetyiswa ukuba kudlale indima ekuhlukaniseni imifanekiso enzima ibe yimiboniso yezinto ezininzi kusetyenziswa ulwazi lwentshukumo ebonakalayo.
Njengoko kukhankanyiwe apha ngasentla, besingabaqalayo ukuxela unxulumano phakathi komsebenzi weseli yeMT pattern kunye nokuqonda. Njengoko kwaqulunqwa kwimodeli yokuqala yezigaba ezibini nguMovshon kunye noogxa bakhe, iyunithi yemowudi yinqanaba lokukhupha leMT. Nangona kunjalo, umsebenzi wakutshanje ubonise ukuba iiseli zemowudi kunye neenxalenye zimele iziphelo ezahlukeneyo ze-continuum kwaye umahluko we-parametric kwisakhiwo sentsimi yokwamkela uxanduva lwe-tuning spectrum yeenxalenye zemowudi. Ke ngoko, sifumene ulwalamano olubalulekileyo phakathi kwe-CP kunye ne-PI, olufana nolwalamano phakathi kwe-binocular mismatch adjustment symmetry kunye ne-CP kumsebenzi wokuqaphela ubunzulu okanye ulungiselelo lokuseta ulwalathiso kumsebenzi wokwahlulahlula ulwalathiso olucokisekileyo. Ubudlelwane phakathi kwamaxwebhu kunye ne-CP 33. UWang kunye ne-Movshon62 bahlalutye inani elikhulu leeseli ezine-MT orientation selectivity kwaye bafumanisa ukuba, ngokomndilili, i-mode index yayinxulunyaniswa neepropati ezininzi zokulungisa, nto leyo ebonisa ukuba i-mode selectivity ikhona kwezinye iintlobo ezininzi zeempawu ezinokufundwa kubemi be-MT. Ngoko ke, kwizifundo ezizayo zobudlelwane phakathi komsebenzi we-MT kunye nokuqonda okuxhomekeke kwingqondo yomntu, kuya kuba yimfuneko ukumisela ukuba isalathisi sepateni sihambelana ngokufanayo nezinye iimpawu zokukhetha umsebenzi kunye ne-stimulus, okanye ukuba olu budlelwane buxhomekeke kwimeko yokwahlulwahlulwa kwembono.
Ngokufanayo, uNienborg kunye noCumming 42 bafumanise ukuba nangona iiseli ezikufutshane nezikude ezikhetha ukungafani kwe-binocular kwi-V2 zazinobuntununtunu obufanayo kumsebenzi wocalucalulo olunzulu, kuphela uluntu lweeseli oluthanda kakhulu olubonakalise i-CP ebalulekileyo. Nangona kunjalo, ukuqeqesha kwakhona iinkawu ukuba zilinganise umahluko okhethekileyo okude kubangele ii-CP ezibalulekileyo kwiikheyiji ezithandwa kakhulu. Ezinye izifundo zikwaxele ukuba imbali yoqeqesho ixhomekeke kulwalamano lokuqonda34,40,63 okanye ubudlelwane obubangela phakathi komsebenzi we-MT kunye nocalucalulo oluhlukileyo48. Ubudlelwane esibubonileyo phakathi kwe-CP kunye nokukhetha indlela yerejimeni kusenokwenzeka ukuba bubonakalisa icebo elithile elisetyenziswe ziinkawu ukusombulula ingxaki yethu, hayi indima ethile yeempawu zokukhetha imo ekuqondeni kwe-visual-motor. Kumsebenzi wexesha elizayo, kuya kuba yimfuneko ukumisela ukuba imbali yokufunda inempembelelo ebalulekileyo ekumiseleni ukuba zeziphi iimpawu ze-MT ezilinganiswa ngokukhethekileyo kwaye ziguquguquke ukuze zenze izigqibo zokwahlulahlula.
UStoner kunye noogxa bakhe14,23 ngabokuqala ukuxela ukuba ukutshintsha ukukhanya kweendawo zegridi ezidityanisiweyo kwachaphazela ukuhambelana kunye nokucaca kweengxelo zabantu abajongayo kunye nohlengahlengiso lwendlela kwii-neurons ze-MT ezinkulu. Ababhali bafumanise ukuba xa ukukhanya kweendawo ezidityanisiweyo kuhambelana nokucaca ngokwasemzimbeni, abajongi baxele ukuqonda okucacileyo ngakumbi, ngelixa ii-neurons ze-MT zibonisa intshukumo yezinto ze-raster. Ngokwahlukileyo koko, xa ukukhanya okudityanisiweyo kunye nokugqagqana okucacileyo kungahambelani ngokwasemzimbeni, umjongi ubona intshukumo ehambelanayo, kwaye ii-neurons ze-MT zibonisa intshukumo yehlabathi yepateni. Ke ngoko, ezi zifundo zibonisa ukuba utshintsho olubonakalayo kwi-stimuli ebonakalayo oluchaphazela ngokuthembekileyo iingxelo zokwahlulwahlulwa kwe-MT lukwabangela utshintsho oluqikelelweyo kwi-MT arousal. Umsebenzi wakutshanje kule ndawo uphonononge ukuba zeziphi iimpawu ze-MT ezilandelela ukubonakala kokubona kwezinto eziyinkimbinkimbi18,24,64. Umzekelo, i-subset yee-neurons ze-MT ibonakalisiwe ukuba ibonisa ukulungiswa kwe-bimodal kwimephu yentshukumo yenqaku elingacwangciswanga (RDK) eneendlela ezimbini ezikude kancinci kune-RDK engeyiyo. I-bandwidth yokulungiswa kweselula 19, 25. Abaqapheli bahlala bebona ipateni yokuqala njengentshukumo ecacileyo, nangona uninzi lwee-neurons ze-MT zibonisa uhlengahlengiso lwe-unimodal ekuphenduleni ezi stimuli, kwaye umyinge olula wazo zonke iiseli ze-MT unika impendulo ye-unimodal population. Ngoko ke, iqela elincinci leeseli ezibonisa ukulungiswa kwe-bimodal linokwenza i-neural substrate yolu luvo. Okunomdla kukuba, kwii-marmosets, eli nani lihambelana neeseli ze-PDS xa kuvavanywa kusetyenziswa i-grid eqhelekileyo kunye ne-grid stimuli.
Iziphumo zethu zihamba inyathelo elinye ngaphezu kwezi zingasentla, ezibalulekileyo ekusekeni indima ye-MT ekuhlukaniseni ukuqonda. Enyanisweni, ukuhlukana yinto eyenzeka ngokwembono. Iimbonakalo ezininzi ezibonakalayo eziguquguqukayo zibonisa amandla enkqubo yokubona ukuhlela nokutolika izinto eziqhubekayo ngeendlela ezingaphezu kwesinye. Ukuqokelela ngaxeshanye iimpendulo ze-neural kunye neengxelo zokuqonda kwisifundo sethu kusivumele ukuba sihlolisise i-covariance phakathi kwesantya sokudubula kwe-MT kunye nokuchazwa kokuqonda kokukhuthaza okuqhubekayo. Emva kokuba sibonise olu lwalamano, siyavuma ukuba ulwalathiso lwesizathu alukasekwa, oko kukuthi, kufuneka iimvavanyo ezongezelelweyo ukuze kuqinisekiswe ukuba uphawu lokuhlukana kokuqonda olubonwa sithi, njengoko abanye bephikisana [65, 66, 67], luyazenzekela na. Le nkqubo iphinda imele imiqondiso ehlayo ebuyela kwi-sensory cortex evela kwiindawo eziphezulu 68, 69, 70 (Umzobo 8). Iingxelo zenxalenye enkulu yeeseli ezikhetha iipateni kwi-MSTd71, enye yezona njongo ziphambili ze-cortical ze-MT, zibonisa ukuba ukwandisa olu vavanyo ukuze lubandakanye ukurekhodwa ngaxeshanye kwe-MT kunye ne-MSTd kuya kuba linyathelo lokuqala elihle lokuqonda ngakumbi iindlela ze-neural zokuqonda.
Imodeli enamanqanaba amabini yokukhetha icandelo kunye nolwalathiso lwendlela kunye nefuthe elinokubakho lempendulo evela phezulu ukuya ezantsi kwimisebenzi enxulumene nokukhetha ekuguquleleni komatshini. Apha, ukukhetha isikhokelo sendlela (PDS – “P”) kwinyathelo le-MT kudalwe (i) isampulu enkulu yedatha yokufaka ekhethiweyo yendlela ehambelana nesantya esithile sendlela, kunye (ii) noxinzelelo oluqinileyo lokulungisa. Icandelo elikhethayo kwicala (CDS) lesigaba se-MT (“C”) linoluhlu oluncinci lwesampulu kwicala lokufaka kwaye alinaxinzelelo olukhulu lokulungisa. Uthintelo olungalungiswanga lunika ulawulo kuzo zombini iindawo. Iintolo ezinemibala zibonisa ulwalathiso lwesixhobo olukhethwayo. Ukuze kucace, kuphela icandelo elincinci lonxibelelwano lwe-V1-MT kunye nebhokisi yokukhetha icandelo elinye kunye ne-orientation. Kwimeko yokutolika iziphumo zethu ze-feed-forward (FF), useto olubanzi lokufaka kunye nothintelo oluqinileyo lokulungisa (oluboniswe ngombala obomvu) kwiiseli ze-PDS lubangele umahluko omkhulu kumsebenzi ekuphenduleni iipatheni ezininzi zokuhamba. Kwingxaki yethu yokwahlulahlula, eli qela liqhuba imixokelelwane yesigqibo kwaye liphazamisa ukuqonda. Ngokwahlukileyo koko, kwimeko yempendulo (FB), izigqibo zokuqonda zenziwa kwiisekethe eziphezulu ngedatha yeemvakalelo kunye nocalucalulo lwengqondo, kwaye isiphumo esikhulu se-FB esezantsi kwiiseli ze-PDS (imigca etyebileyo) ivelisa imiqondiso yokukhetha. b Ukubonakaliswa kwesicwangciso seemodeli ezizezinye zezixhobo ze-CDS kunye ne-PDS. Apha imiqondiso ye-PDS kwi-MT ayiveliswanga kuphela ngokufaka ngqo kwe-V1, kodwa nangokufaka ngokungathanga ngqo kwendlela ye-V1-V2-MT. Iindlela ezingathanga ngqo zemodeli zihlengahlengiswa ukuze zinike ukukhetha kwimida yokuthungwa (iindawo ezigqubanayo zegridi). Imodyuli ye-CDS yomaleko we-MT yenza isixa esilinganiselweyo sezinto ezifakwa ngqo nezingathanga ngqo kwaye ithumela imveliso kwimodyuli ye-PDS. I-PDS ilawulwa kukuthintelwa kokhuphiswano. Kwakhona, kuphela ezo nxibelelwano ziboniswayo ezifunekayo ukuzoba uyilo olusisiseko lwemodeli. Apha, indlela eyahlukileyo ye-FF kunaleyo icetywayo kwi-a inokukhokelela kutshintsho olukhulu kwimpendulo ye-lattice yeselula kwi-PDS, kwakhona ikhokelele ekucalucaluleni kwiipateni zesigqibo. Ngaphandle koko, i-CP enkulu kwiiseli ze-PDS isenokuba sisiphumo sokucalucalula kumandla okanye ukusebenza kakuhle kokuncamathiselwa kwe-FB kwiiseli ze-PDS. Ubungqina buxhasa iimodeli ze-MT PDS ezinamanqanaba amabini nantathu kunye neengcaciso ze-CP FF kunye ne-FB.
Iimacaque ezimbini ezindala (i-macaca mulatta), enye inkunzi nenye imazi (iminyaka esi-7 kunye ne-5 ubudala ngokulandelelana), ezinobunzima obuqala kwi-4.5 ukuya kwi-9.0 kg, zasetyenziswa njengezinto zokufunda. Ngaphambi kokuba kwenziwe uvavanyo lotyando olungenazintsholongwane, izilwanyana zafakwa igumbi lokurekhoda elenziwe ngokwezifiso zee-electrode ezithe nkqo ezisondela kwindawo ye-MT, isitendi sentloko yentsimbi engagqwaliyo (iCrist Instruments, iHagerstown, MD), kunye nendawo yamehlo ene-coil yokukhangela ye-scleral elinganisiweyo. (iCooner Wire, eSan Diego, eCalifornia). Zonke iiprotokholi ziyahambelana nemigaqo yeSebe lezoLimo lase-United States (i-USDA) kunye neMigaqo yeSizwe yezeMpilo (i-NIH) yoKhathalelo loLuntu kunye nokusetyenziswa kwezilwanyana zaseLebhu kwaye zivunyiwe yiKomiti yoKhathalelo lweZilwanyana zaseYunivesithi yaseChicago (IAUKC).
Zonke izinto ezibonakalayo ziboniswe kwindawo evulekileyo ejikelezileyo ngokuchasene nemvelaphi emnyama okanye engwevu. Ngexesha lokurekhoda, indawo kunye nobubanzi balo mngxuma zilungiswe ngokuhambelana nentsimi yokwamkela ye-neurons kwincam ye-electrode. Sisebenzise iindidi ezimbini ezibanzi zezinto ezibonakalayo: izinto zengqondo kunye nezinto ezibonakalayo.
I-psychometric stimulus yipatheni ye-grating (20 cd/m2, 50% contrast, 50% duty cycle, 5 degrees/sec) eyenziwe ngokubeka ii-rectangular gratings ezimbini ezijikelezayo kwicala elithe nkqo kwicala lazo (Umzobo 1b). Kuye kwaboniswa ngaphambili ukuba abantu ababukeleyo babona ezi patheni zegridi njengezinto ezikhuthazayo eziguquguqukayo, ngamanye amaxesha njengepatheni enye ehamba kwicala elinye (intshukumo ehambelanayo) kwaye ngamanye amaxesha njengeendawo ezimbini ezahlukeneyo ezihamba kwicala ezahlukeneyo (intshukumo ebonakalayo). Izinto zepatheni ye-lattice, ezijolise ngokulinganayo - i-engile ephakathi kwee-lattice isuka kwi-95° ukuya kwi-130° (ethathwe kwiseti: 95°, 100°, 105°, 115°, 120°, 125°, 130° °, kuyo yonke iseshoni. Ii-Isolation angle neurons kwi-115° azizange zigcinwe, kodwa sibandakanya idatha ye-psychophysical apha) - malunga ne-90° okanye i-270° (ulwalathiso lwepatheni). Kwiseshoni nganye, kusetyenziswe ikona enye kuphela yelatisi ye-interlattice; ngexesha leseshoni nganye, indlela yokujonga ipateni yovavanyo ngalunye ikhethwe ngokungacwangciswanga kwiindlela ezimbini ezinokwenzeka.
Ukuze sihlukanise indlela ebonwa ngayo igridi size sinike isiseko esibonakalayo somvuzo wesenzo, singenisa ii-random point textures kwinyathelo le-light bar 72 le-grid component nganye. Oku kufezekiswa ngokunyusa okanye ukunciphisa (ngenani elimiselweyo) ukukhanya kwe-subset ekhethiweyo ngokungacwangciswanga yee-pixels (Umzobo 1c). Icala lentshukumo ye-texture linika isignali enamandla etshintsha indlela obona ngayo umjongi ukuya kwi-coherent okanye transparent movement (Umzobo 1c). Phantsi kweemeko ezihambelanayo, zonke ii-textures, nokuba zeziphi ii-partition ze-texture lattice covers, ziguqulelwa kwicala le-pattern (Umzobo 1c, coherent). Kwimeko ecacileyo, i-texture ihamba ngokuthe nkqo kwicala le-grating eyigubungelayo (Umzobo 1c, transparent) (I-Supplementary Movie 1). Ukulawula ubunzima bomsebenzi, kwiiseshoni ezininzi i-Michelson contrast (Lmax-Lmin/Lmax+Lmin) yale texture mark yahluka kwi-set ye-(-80, -40, -20, -10, -5, 0, 5). , 10, 20, 40, 80). Umahluko uchazwa njengokukhanya okuqhelekileyo kwe-raster (ngoko ke ixabiso lomahluko le-80% liya kubangela ubume be-36 okanye i-6 cd/m2). Kwiiseshoni ezi-6 kwi-monkey N kunye neeseshoni ezi-5 kwi-monkey S, sisebenzise uluhlu oluncinci lomahluko wombhalo (-30, -20, -15, -10, -5, 0, 5, 10, 15, 20, 30), apho iimpawu zengqondo zilandela iphethini efanayo nomahluko opheleleyo, kodwa ngaphandle kokwanda.
Izinto ezivuselelayo zi-sinusoidal grids (umahluko we-50%, umjikelo/idigri e-1, iidigri ezi-5/isekhondi) ezihamba kwelinye lamacala ali-16 ahlukeneyo ngokulinganayo, okanye ii-sinusoidal grids ezihamba kwezi ndlela (eziqulathe ii-angles ezimbini ezichaseneyo ze-135° superimposed sinusoidal gratings phezu kwenye), kwicala elifanayo lepateni.