Alamomin jijiyoyi na rarrabuwar fahimta a cikin cortex na lokaci na macaques

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Carousel yana nuna zamiya guda uku a lokaci guda. Yi amfani da maɓallan da suka gabata da na gaba don motsawa ta zamiya guda uku a lokaci guda, ko kuma yi amfani da maɓallan zamiya a ƙarshen don motsawa ta zamiya guda uku a lokaci guda.
Ganin gani mai ƙuduri mai girma yana buƙatar samfurin retina mai kyau da haɗin kai don sake gina halayen abu. Yana da mahimmanci a lura cewa lokacin haɗa samfuran gida daga abubuwa daban-daban, daidaito yana ɓacewa. Saboda haka, rarrabawa, haɗa wuraren hoto don sarrafawa daban, yana da mahimmanci ga fahimta. A cikin aikin da ya gabata, an yi amfani da tsarin lattice na bistable, wanda za'a iya ɗauka azaman ɗaya ko fiye da saman motsi, don nazarin wannan tsari. A nan, muna ba da rahoton alaƙar da ke tsakanin aiki da hukunce-hukuncen rarrabuwa a yankuna na tsakiya na hanyar gani ta asali. Musamman, mun gano cewa karkatar da hankali tsakanin ƙwayoyin jijiyoyi na ɗan lokaci yana da alaƙa da alamun rubutu da aka yi amfani da su don karkatar da fahimtar bistable gratings kuma ya nuna muhimmiyar alaƙa tsakanin gwaji da fahimtar ra'ayi na tunani na abubuwan da ke ci gaba. Wannan alaƙar ta fi girma a cikin raka'a waɗanda ke nuna motsi na duniya a cikin alamu tare da alƙawura da yawa na gida. Don haka, mun kammala da cewa yankin lokaci na tsaka-tsaki ya ƙunshi sigina da ake amfani da su don raba al'amuran rikitarwa zuwa abubuwa da saman.
Gani ba wai kawai ya dogara ne akan bambancin halayen hoto na asali kamar yanayin gefen da saurin ba, amma mafi mahimmanci akan haɗakar waɗannan halaye daidai don ƙididdige halayen muhalli kamar siffar abu da hanyar da ke tafiya1. Duk da haka, matsaloli suna tasowa lokacin da hotunan retinal ke tallafawa ƙungiyoyin fasali da dama masu yuwuwa iri ɗaya 2, 3, 4 (Hoto na 1a). Misali, lokacin da saitin siginar gudu guda biyu suka yi kusa sosai, ana iya fassara wannan a matsayin abu ɗaya mai motsi ko abubuwa da yawa masu motsi (Hoto na 1b). Wannan yana nuna yanayin rarrabuwar kai, watau ba ƙayyadadden sigar ba ce, amma tsari ne na fassara. Duk da mahimmancin fahimta ta al'ada, fahimtarmu game da tushen jijiyoyi na rarrabuwar fahimta har yanzu ba ta cika ba.
Zane mai zane na matsalar rarrabuwar fahimta. Fahimtar mai kallo game da zurfin da ke cikin kube na Necker (hagu) yana canzawa tsakanin bayanai guda biyu masu yiwuwa (dama). Wannan saboda babu sigina a cikin hoton da ke ba wa kwakwalwa damar tantance yanayin 3D na siffar ta musamman (wanda siginar rufewa ta monocular a dama ta bayar). b Lokacin da aka gabatar da siginar motsi da yawa a kusancin sarari, tsarin hangen nesa dole ne ya tantance ko samfuran gida sun fito ne daga abu ɗaya ko fiye. Rashin tabbas na siginar motsi na gida, watau jerin motsin abu na iya samar da motsi iri ɗaya na gida, wanda ke haifar da fassarori da yawa masu dacewa na shigarwar gani, watau filayen vector a nan na iya tasowa daga motsi mai daidaituwa na saman ɗaya ko motsi mai haske na saman da ke haɗuwa. c (hagu) Misali na motsin grid ɗinmu mai laushi. Gratings masu kusurwa huɗu suna shawagi a tsaye zuwa ga alkiblarsu ("alkiblar sashi" - kibiyoyi fari) suna haɗuwa da juna don samar da tsarin grating. Ana iya fahimtar raga a matsayin motsi ɗaya, na yau da kullun, mai haɗawa na al'ada (kibiyoyi ja) ko motsi mai haske na alkiblar mahaɗi. Fahimtar layin raga ta karkace ta hanyar ƙara alamun rubutu na bazuwar. (Tsakiya) Yankin da aka haskaka da rawaya an faɗaɗa shi kuma ana nuna shi azaman jerin firam don sigina masu haɗawa da bayyana, bi da bi. Motsin digo a kowane yanayi ana wakilta shi da kibiyoyi kore da ja. (Dama) Zane na matsayi (x, y) na wurin zaɓi idan aka kwatanta da adadin firam. A cikin yanayin da aka haɗa, duk laushi suna tafiya a hanya ɗaya. Idan aka haɗa haske, yanayin yana motsawa a alkiblar ɓangaren. d Misalin zane mai ban dariya na aikin raba motsi. Birai sun fara kowace gwaji ta hanyar gyara ƙaramin digo. Bayan ɗan lokaci kaɗan, wani nau'in tsarin grating (daidaituwa/bayyananne) da girman siginar rubutu (misali bambanci) sun bayyana a wurin MT RF. A lokacin kowane gwaji, grating na iya zamewa a ɗaya daga cikin hanyoyi biyu masu yiwuwa na tsarin. Bayan janyewar motsawa, maƙasudin zaɓi sun bayyana a sama da ƙasa da MT RF. Birai dole ne su nuna fahimtarsu game da grid a cikin saccades zuwa ga maƙasudin zaɓi da ya dace.
Tsarin sarrafa motsin gani yana da kyau kuma don haka yana ba da kyakkyawan tsari don nazarin da'irorin jijiyoyi na rarrabuwar fahimta. Nazarce-nazarce da yawa na lissafi sun lura da amfanin samfuran sarrafa motsi masu matakai biyu inda ake bin ƙididdigar farko mai ƙuduri mai girma tare da haɗa samfuran gida don cire hayaniya da dawo da saurin abu7,8. Yana da mahimmanci a lura cewa tsarin gani dole ne ya yi taka tsantsan don iyakance wannan tarin ga waɗancan samfuran gida daga abubuwa na yau da kullun. Nazarin ilimin halayyar ɗan adam ya bayyana abubuwan jiki waɗanda ke tasiri ga yadda ake raba siginar motsi na gida, amma siffar hanyoyin jiki da lambobin jijiyoyi sun kasance tambayoyi a buɗe. Rahotanni da yawa sun nuna cewa ƙwayoyin da ke zaɓar alkibla a yankin lokaci (MT) na ƙwayar halittar primate suna da 'yan takara don substrates na jijiyoyi.
Abu mafi mahimmanci, a cikin waɗannan gwaje-gwajen da suka gabata, canje-canje a cikin ayyukan jijiyoyi suna da alaƙa da canje-canje na zahiri a cikin abubuwan da ke motsa gani. Duk da haka, kamar yadda aka ambata a sama, rarrabuwar abu a zahiri tsari ne na fahimta. Saboda haka, nazarin substrate na jijiyoyi yana buƙatar haɗa canje-canje a cikin ayyukan jijiyoyi tare da canje-canje a cikin fahimtar abubuwan da ke motsa jiki. Saboda haka, mun horar da birai biyu don bayar da rahoton ko tsarin grating da aka gani wanda aka samar ta hanyar gratings masu jujjuyawa masu juyawa shine saman ɗaya ko saman biyu masu zaman kansu. Don nazarin dangantakar da ke tsakanin aikin jijiyoyi da hukunce-hukuncen rarrabuwa, mun rubuta aiki ɗaya a cikin MT lokacin da birai suka yi wannan aikin.
Mun sami wata muhimmiyar alaƙa tsakanin nazarin ayyukan MT da fahimta. Wannan alaƙar ta wanzu ko abubuwan da ke haifar da ita sun ƙunshi alamun rarrabuwa a bayyane ko a'a. Bugu da ƙari, ƙarfin wannan tasirin yana da alaƙa da jin daɗin siginar rarrabuwa, da kuma ma'aunin tsari. Na ƙarshen yana ƙididdige matakin da na'urar ke haskakawa a duniya maimakon motsi na gida a cikin tsare-tsare masu rikitarwa. Kodayake zaɓin alkiblar salo an daɗe ana gane shi a matsayin siffa mai ma'ana ta MT, kuma ƙwayoyin da ke zaɓar salo suna nuna daidaitawa ga abubuwan da ke haifar da rikitarwa daidai da fahimtar ɗan adam game da waɗannan abubuwan, gwargwadon iliminmu, wannan shine shaida ta farko don alaƙa tsakanin tsare-tsare. ma'auni da rarrabuwar fahimta.
Mun horar da birai biyu don nuna fahimtarsu game da motsin grid mai zamewa (motsi mai daidaituwa ko bayyananne). Masu lura da mutane galibi suna ɗaukar waɗannan abubuwan ƙarfafawa a matsayin motsi mai daidaituwa ko bayyananne na kusan mita ɗaya. Domin bayar da amsar da ta dace a cikin wannan gwaji da kuma kafa tushen ladan aiki, mun ƙirƙiri siginar rarrabuwa ta hanyar zana raster na ɓangaren da ke samar da layin (Hoto na 1c, d). A ƙarƙashin yanayi mai daidaituwa, duk laushi suna motsawa tare da alkiblar tsarin (Hoto na 1c, "daidai"). A cikin yanayin haske, laushi yana motsawa daidai da alkiblar grating ɗin da aka ɗora shi a kai (Hoto na 1c, "mai haske"). Muna sarrafa wahalar aikin ta hanyar canza bambancin wannan lakabin rubutu. A cikin gwaje-gwajen da aka ɗauka, an ba birai lada don amsoshi da suka dace da alamun rubutu, kuma an ba da lada bazuwar (50/50 rashin daidaito) a cikin gwaje-gwajen da ke ɗauke da alamu ba tare da alamun rubutu ba (yanayin bambancin rubutu sifili).
An nuna bayanan ɗabi'a daga gwaje-gwaje guda biyu masu wakilci a Hoto na 2a, kuma an tsara amsoshin a matsayin rabo na hukunce-hukuncen daidaito da bambancin siginar rubutu (ana ɗaukar bambancin bayyana a matsayin mara kyau ta ma'anar) don alamu waɗanda ke canzawa sama ko ƙasa, bi da bi. Gabaɗaya, fahimtar birai game da daidaito/bayyananne ya shafi tasirin alamar (mai haske, mai jituwa) da ƙarfi (bambanci) na alamar laushi (ANOVA; biri N: alkibla – F = 0.58, p = 0.45, alamar – F = 1248, p < 10−10, bambanci – F = 22.63, p < 10;−10 biri S: alkibla – F = 0.41, p = 0.52, alama – F = 2876.7, p < 10−10, bambanci – F = 36.5, p < 10−10). Gabaɗaya, fahimtar birai game da daidaito/bayyananne ya shafi tasirin alamar (mai haske, mai jituwa) da ƙarfi (bambanci) na alamar laushi (ANOVA; biri N: alkibla – F = 0.58, p = 0.45, alamar – F = 1248, p < 10−10, bambanci – F = 22.63, p < 10; −10 biri S: alkibla – F = 0.41, p = 0.52, alama – F = 2876.7, p < 10−10, bambanci – F = 36.5, p < 10−10). В целом на в восприятие (контрастность) текстурного признака (ANOVA; обезьяна N: направление — F = 0,58, p = 0,45, знак — F = 1248, p < 10−10, контраст – F = 22,63, p < 10; -10 обезьяна S: направление – F = 0,41, p = 0,52, признак – F = 2876,7, p <10-10, kontrast - F = 36,5, р <10-10). Gabaɗaya, fahimtar daidaito/bayyanannen abu ta birai ya shafi fahimtar daidaito/bayyanannen abu ta hanyar alamar (bayyananne, haɗuwa) da ƙarfi (bambanci) na siffar rubutu (ANOVA; biri N: jagora — F = 0.58, p = 0.45, alama — F = 1248, p < 10−10, bambanci – F = 22.63, p < 10; -10 biri S: jagora – F = 0.41, p = 0.52, alama – F = 2876.7, p < 10 −10, bambanci – 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 = 0.52, 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).Gabaɗaya, fahimtar daidaiton biri/bayyananne ya shafi alamar (bayyananne, daidaito) da kuma ƙarfi (bambanci) na siginar rubutu (ANOVA);обезьяна N: ориентация - F = 0,58, p = 0,45, знак - F = 1248, p <10−10, Контрастность - F = 22,63, p <10; biri N: daidaitawa – F = 0.58, p = 0.45, alama – F = 1248, p < 10−10, Bambanci – 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 Biri S: Gabatarwa – F = 0.41, p = 0.52, Alamar – F = 2876.7, p < 10-10, Bambanci – F = 36.5, p < 10-10).An haɗa ayyukan tarin Gaussian zuwa ga bayanai daga kowane zaman don kwatanta halayen ilimin halayyar birai. A hoto na 2b yana nuna yadda aka rarraba waɗannan samfuran a duk zaman birai biyu. Gabaɗaya, birai sun kammala aikin daidai kuma akai-akai, kuma mun ƙi ƙasa da kashi 13% na zaman birai biyu saboda rashin dacewa da tsarin Gaussian mai tarin yawa.
Misalan Halayyar Birai a Zaman Wakilta (n ≥ 20 gwaje-gwaje ga kowane yanayi na ƙarfafawa). A cikin bangarorin hagu (dama), bayanai daga zaman biri ɗaya na N(S) an tsara su azaman maki masu daidaituwa (daidaitawa) idan aka kwatanta da bambancin alamun siginar rubutu (abscissa). A nan ana ɗauka cewa laushi masu haske (masu daidaituwa) suna da ƙimar korau (masu kyau). An gina amsoshi daban-daban bisa ga alkiblar motsi na tsarin (sama (90°) ko ƙasa (270°)) a cikin gwajin. Ga dabbobi biyu, aiki, ko an raba martanin da bambanci 50/50 (PSE - kibiyoyi masu ƙarfi) ko adadin bambancin rubutu da ake buƙata don tallafawa wani matakin aiki (matsakaicin - kibiyoyi buɗe), yana cikin waɗannan kwatancen karkacewa. b An daidaita histogram na ƙimar R2 na aikin tarin Gaussian. Bayanan Monkey S(N) an nuna su a hagu (dama). c (Sama) An auna PSE don grid ɗin da aka canza ƙasa (ordinate) idan aka kwatanta da PSE da aka canza sama da grid ɗin (abscissa) da aka tsara, tare da gefuna suna wakiltar rarrabawar PSE don kowane yanayi da kibiyoyi suna nuna matsakaicin kowane yanayi. Bayanan duk zaman biri na N(S) an bayar da su a ginshiƙin hagu (dama). (Ƙasa) Daidai da na bayanan PSE, amma don iyakokin fasali da aka daidaita. Babu wani bambanci mai mahimmanci a cikin iyakokin PSE ko yanayin salo (duba rubutu). d An tsara PSE da gangare (ordinate) dangane da yanayin raster da aka daidaita na ɓangaren rabuwar kusurwa ("kusurwar grating mai haɗawa" - abscissa). Da'irori masu buɗewa sune hanyoyin, layin mai ƙarfi shine mafi kyawun samfurin komawa baya, kuma layin da aka nuna shine tazara ta amincewa 95% don samfurin komawa baya. Akwai muhimmiyar alaƙa tsakanin PSE da kusurwar haɗin kai da aka daidaita, amma ba gangare da kusurwar haɗin kai da aka daidaita ba, yana nuna cewa aikin psychometric yana canzawa yayin da kusurwar ke raba layukan kayan, amma ba kaifi ko lanƙwasa ba. (Biri N, n = zaman 32; Biri S, n = zaman 43). A cikin dukkan bangarori, sandunan kuskure suna wakiltar kuskuren daidaito na matsakaicin. Haha. Daidaito, maki daidaito na PSE, ƙa'ida. daidaito.
Kamar yadda aka ambata a sama, bambancin alamomin rubutu da kuma alkiblar motsi na tsarin ya bambanta a cikin gwaje-gwaje, tare da abubuwan da ke motsa jiki suna zagayawa sama ko ƙasa a cikin wani gwaji da aka bayar. Ana yin wannan ne don rage tasirin daidaitawa na psychophysical11 da neuronal28. Tsarin daidaitawa da son zuciya (ma'aunin daidaito na mutum ko PSE) (Gwajin jimlar matsayi na Wilcoxon; biri N: z = 0.25, p = 0.8; biri S: z = 0.86, p = 0.39) ko kuma matakin aiki da aka daidaita (jimillar matsayin Wilcoxon; biri N: z = 0.14, p = 0.89, biri S: z = 0.49, p = 0.62) (Hoto na 2c). Bugu da ƙari, babu wani muhimmin bambanci tsakanin birai a matakin bambancin rubutu da ake buƙata don kiyaye matakan matakin aiki (N biri = 24.5% ± 3.9%, S biri = 18.9% ± 1.9%; jimlar matsayi na Wilcoxon, z = 1.01, p = 0.31).
A kowane zaman, mun canza kusurwar da ke tsakanin layukan sassan da ke raba alkiblar sassan. Nazarin ilimin halayyar ɗan adam ya nuna cewa mutane sun fi fahimtar tantanin halitta 10 a matsayin wanda aka haɗa lokacin da wannan kusurwar ta ƙanƙanta. Idan birai suna ba da rahoton fahimtarsu game da daidaito/bayyananne, to bisa ga waɗannan binciken, za a yi tsammanin PSE, bambancin rubutu wanda ya dace da raba daidai tsakanin daidaito da zaɓin bayyananne, zai ƙaru yayin hulɗa. kusurwar layi. Wannan gaskiya ne (Hoto na 2d; rugujewa ta hanyar kwatancen tsari, Kruskal–Wallis; biri N: χ2 = 23.06, p < 10−3; biri S: χ2 = 22.22, p < 10−3; alaƙa tsakanin kusurwar da aka daidaita tsakanin grating da PSE - biri N: r = 0.67, p < 10−9; biri S: r = 0.76, p < 10−13). Wannan gaskiya ne (Hoto na 2d; rugujewa ta hanyar kwatancen tsari, Kruskal–Wallis; biri N: χ2 = 23.06, p < 10−3; biri S: χ2 = 22.22, p < 10−3; alaƙa tsakanin kusurwar da aka daidaita tsakanin juna da PSE – biri N: r = 0.67, p < 10−9; biri S: r = 0.76, p < 10−13). Это действительно имело место (rис. 2d; колапс поперек направления патерна, Крускал-Уоллис; p:2, 6 N. 10–3; обезьяна S: χ2 = 22,22, p < 10–3; обезьяна S: r = 0,76, p <10-13). Wannan ya faru da gaske (Hoto na 2d; rugujewa a kan alkiblar tsarin, Kruskal-Wallis; biri N: χ2 = 23.06, p < 10–3; biri S: χ2 = 22.22, p < 10–3; alaƙa tsakanin kusurwar layi mai daidaitawa da PSE - biri N: r = 0.67, p < 10-9, biri S: 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; 2.2 10-3;情况 确实 如此 (图 图 2D ; 方向 折叠 , kruskal-wallis ; n 22.22 10-9;猴子S:r = 0.76,p <10-13) Это действительно имело место (rис. 2d; kratность по оси моды, Крускал-Уолис; обезяна N: χ2 = 23, 06 χ2 = 22,22, p <10-3; Wannan gaskiya ne (Hoto na 2d; ninka a kan ma'aunin yanayin, Kruskal-Wallis; biri N: χ2 = 23.06, p < 10-3; biri S: χ2 = 22.22, p < 10-3; kusurwar interlattice mai tsari). PSE-обезьяна N: r = 0,67, p <10-9, обезьяна S: r = 0,76, p <10-13). Birin PSE N: r = 0.67, p < 10–9, biri S: r = 0.76, p < 10–13).Sabanin haka, canza kusurwar interlattice ba ta da wani tasiri mai mahimmanci a kan gangaren aikin psychometric (Hoto na 2d; ninka yanayin daidaitawa, Kruskal-Wallis; biri N: χ2 = 8.09, p = 0.23; biri S χ2 = 3.18, p = 0.67, alaƙa tsakanin kusurwar interlattice da aka daidaita da gangare - biri N: r = -0.4, p = 0.2, biri S: r = 0.03, p = 0.76). Don haka, daidai da bayanan psychophysical na mutum, matsakaicin tasirin canza kusurwar tsakanin gratings shine sauyawa a wuraren ƙaura, ba ƙari ko raguwa a cikin amsawa ga siginar rarrabuwa ba.
A ƙarshe, ana ba da lada bazuwar tare da yuwuwar 0.5 a cikin gwaje-gwajen da ba su da bambancin rubutu sifili. Da dukkan birai sun san wannan bambancin rubutu na musamman kuma sun sami damar bambance tsakanin bambancin rubutu sifili da abubuwan da ke motsa alamun, za su iya haɓaka dabarun daban-daban don nau'ikan gwaje-gwajen guda biyu. Abubuwan lura guda biyu sun nuna cewa ba haka lamarin yake ba. Na farko, canza kusurwar grating yana da tasiri iri ɗaya akan maki bambancin rubutu sifili da sifili (Hoto na 2d da Ƙarin Hoto na 1). Na biyu, ga birai biyu, zaɓin gwajin bistable ba zai zama maimaita zaɓin lada na baya ba (na baya) (gwajin binomial, birai N: 0.52, z = 0.74, p = 0.22; birai S: 0.51, r = 0.9, p = 0.18).
A ƙarshe, halayen birai a cikin aikin raba su yana ƙarƙashin kyakkyawan iko na ƙarfafawa. Dogaro da hukunce-hukuncen fahimta akan alamar da girman alamun rubutu, da kuma canje-canje a cikin PSE tare da kusurwar grating, yana nuna cewa birai sun ba da rahoton fahimtarsu ta zahiri game da daidaito/bayyana motsi. A ƙarshe, martanin birai a cikin gwaje-gwajen bambancin rubutu sifili bai shafi tarihin lada na gwaje-gwajen da suka gabata ba kuma canje-canjen kusurwa tsakanin raster sun shafi shi sosai. Wannan yana nuna cewa birai suna ci gaba da ba da rahoton fahimtarsu ta zahiri game da tsarin saman lattice a ƙarƙashin wannan muhimmin yanayi.
Kamar yadda aka ambata a sama, sauyin bambancin rubutu daga mummunan zuwa mai kyau daidai yake da sauyin fahimta na abubuwan da ke motsa jiki daga haske zuwa mai jituwa. Gabaɗaya, ga wani tantanin halitta, martanin MT yana ƙaruwa ko raguwa yayin da bambancin rubutu ke canzawa daga mummunan zuwa mai kyau, kuma alkiblar wannan tasirin yawanci ya dogara ne da alkiblar motsi na tsari/ɓangaren. Misali, an nuna lanƙwasa na daidaita alkibla na ƙwayoyin MT guda biyu masu wakiltar a Hoto na 3 tare da amsoshin waɗannan ƙwayoyin zuwa ga gratings waɗanda ke ɗauke da ƙananan ko babban siginar rubutu mai jituwa ko mai haske. Mun yi ƙoƙari ta wata hanya don ƙididdige waɗannan amsoshin grid mafi kyau, wanda zai iya danganta da aikin psychophysiological na birai namu.
Tsarin polar na lanƙwasa mai daidaitawa na kwatancen wata kwayar halitta ta biri MT S wacce ke wakiltar S a matsayin martani ga jerin sinusoidal guda ɗaya. Kusurwar tana nuna alkiblar motsi na grating, girman yana nuna fitar da iska, kuma alkiblar tantanin halitta da aka fi so ta haɗu da kusan 90° (sama) tare da alkiblar ɗaya daga cikin abubuwan da ke cikin alkiblar tsarin grating. b Histogram na lokacin motsa jiki na mako-mako (PSTH) na grid ɗin amsawa, an canza shi zuwa alkiblar samfuri da 90° (wanda aka nuna a cikin tsari a hagu) don tantanin da aka nuna a cikin a. Ana rarraba amsoshin ta hanyar nau'in alamar rubutu (haɗe-haɗe/mai haske - tsakiya/dama panel bi da bi) da kuma bambancin Michelson (alamar launi ta PSTH). Ana nuna yunƙurin da ya dace kawai ga kowane nau'in siginar rubutu mai ƙarancin bambanci da babban bambanci. Kwayoyin halitta sun fi mayar da martani ga tsarin lattice mai juyawa sama tare da alamun rubutu mai haske, kuma martani ga waɗannan alamu ya ƙaru tare da ƙaruwar bambancin rubutu. c, d iri ɗaya ne kamar a da b, amma ga ƙwayoyin MT banda biri S, yanayin da suka fi so ya kusan haɗuwa da grid mai juyawa ƙasa. Naúrar ta fi son ramuka masu juyawa ƙasa tare da alamun rubutu masu daidaituwa, kuma martanin waɗannan alamu yana ƙaruwa tare da ƙaruwar bambancin rubutu. A cikin dukkan bangarori, yankin da aka yi wa inuwa yana wakiltar kuskuren daidaitattun matsakaici. Kafafu. Ƙafafu, daƙiƙa. na biyu.
Don bincika alaƙar da ke tsakanin tsarin saman lattice (mai daidaituwa ko bayyananne) kamar yadda siginar rubutu da ayyukan MT suka nuna, da farko mun koma ga alaƙar da ke tsakanin ƙwayoyin halitta don motsi mai daidaituwa (zube mai kyau) ko motsi mai haske (zube mara kyau) ta hanyar komawa baya. wanda aka ba wa rarraba ƙwayoyin ta hanyar ƙimar amsawar alama idan aka kwatanta da bambanci (daban ga kowane alkiblar yanayi). Misalan waɗannan lanƙwasa masu daidaita lattice daga tantanin halitta iri ɗaya a cikin Hoto na 3 an nuna su a Hoto na 4a. Bayan rarrabuwa, mun yi amfani da nazarin aikin mai karɓa (ROC) don ƙididdige ƙarfin kowace tantanin halitta ga daidaitawar siginar rubutu (duba Hanyoyi). Ayyukan neurometric da aka samu ta wannan hanyar za a iya kwatanta su kai tsaye da halayen psychophysical na birai a cikin zaman ɗaya don kwatanta kai tsaye yanayin psychophysical na neurons zuwa yanayin lattice. Mun yi nazarin gano sigina guda biyu don duk raka'a a cikin samfurin, muna ƙididdige fasalulluka daban-daban na neurometric don kowane alkibla na tsarin (kuma, sama ko ƙasa). Yana da mahimmanci a lura cewa, don wannan binciken, mun haɗa da gwaje-gwaje kawai waɗanda (i) abubuwan da ke haifar da motsin jiki sun ƙunshi alamar rubutu kuma (ii) birai sun amsa daidai da wannan alamar (watau, gwaje-gwajen "daidai").
An tsara ƙimar wuta bisa ga bambancin alamun rubutu, bi da bi, don gratings waɗanda ke canzawa sama (hagu) ko ƙasa (dama), layin mai ƙarfi yana wakiltar mafi kyawun daidaitawar layi, kuma an ɗauko bayanan da ke cikin layi na sama (ƙasa) daga waɗanda aka nuna a Hoto na . Shinkafa. 3a Cell, b (Hoto na 3c, d). An yi amfani da fasalulluka na gangaren komawa baya don sanya alamun rubutu da aka fi so (daidaitacce/mai haske) ga kowane haɗin daidaitawar tantanin halitta/lattice (n ≥ gwaje-gwaje 20 a kowane yanayin ƙarfafawa). Sandunan kuskure suna wakiltar daidaitaccen karkacewar matsakaicin. Ayyukan neurometric na raka'a da aka nuna a cikin ba an bayyana su tare da ayyukan psychometric da aka tattara a lokacin zaman ɗaya. Yanzu, ga kowane fasali, mun tsara zaɓin shawarar kayan aiki da aka fi so (daidaitawa) (duba rubutu) a matsayin kashi na bambancin alamar rubutu (abscissa). An canza bambancin rubutu don haka shawarwarin kayan aiki da aka fi so suna da kyau kuma shawarwarin kayan aiki marasa komai suna da korau. An nuna bayanai daga grids masu juyawa sama (ƙasa) a cikin bangarorin hagu (dama), a cikin layuka na sama (ƙasa) - bayanai daga ƙwayoyin da aka nuna a Hoto na 3a, b (Hoto na 3c, d). An nuna rabon matakin neurometric da psychometric (N/P) a kowane bangare. Kafafu. Ƙafafu, daƙiƙa. sec, jagorar. alkibla, fifikon lardin, psi. Psychometry, Jijiyoyin Jijiyoyi.
An nuna lanƙwasa masu daidaita lattice da ayyukan neurometric na ƙwayoyin MT guda biyu masu wakiltar juna da ayyukan psychometric ɗinsu masu alaƙa, waɗanda aka haɗa tare da waɗannan amsoshin, a saman da ƙasan allon Hoto na 4a, b, bi da bi. Waɗannan ƙwayoyin suna nuna ƙaruwa ko raguwa mai kama da juna yayin da alamar rubutu ke tafiya daga haske zuwa mai jituwa. Bugu da ƙari, alkibla da ƙarfin wannan haɗin sun dogara ne akan alkiblar motsin lattice. A ƙarshe, ayyukan neurometric da aka ƙididdige daga amsoshin waɗannan ƙwayoyin sun kusanci kawai (amma har yanzu ba su yi daidai da) halayen psychophysical na motsi na grid mai jagora ɗaya ba. An taƙaita ayyukan neurometric da psychometric da ƙofofi, watau daidai da kusan 84% na bambancin da aka zaɓa daidai (wanda ya yi daidai da matsakaicin + 1 sd na aikin Gaussian mai tarin yawa da aka daidaita). A duk faɗin samfurin, rabon N/P, rabon iyakar jijiyoyi zuwa na psychometric, ya kai matsakaicin 12.4 ± 1.2 a cikin biri N da 15.9 ± 1.8 a cikin biri S, kuma don layin ya motsa aƙalla alkibla ɗaya, kawai A ~16% (18.%) raka'a daga biri N (biri S) (Hoto na 5a). Daga misalin tantanin halitta da aka nuna a cikin hoton. Kamar yadda aka gani a cikin Hoto na 3 da 4, yanayin jijiyoyi na iya shafar alaƙar da ke tsakanin yanayin da aka fi so na tantanin halitta da kuma alkiblar motsi na layin da aka yi amfani da shi a cikin gwaje-gwajen. Musamman ma, lanƙwasa daidaitawar daidaitawa a cikin Hoto na 3a, c yana nuna alaƙar da ke tsakanin saitin yanayin jijiyoyi na jerin sinusoidal guda ɗaya da kuma yanayinsa ga motsi mai haske/mai jituwa a cikin jerin rubutunmu. Wannan shi ne lamarin ga birai biyu (ANOVA; umarnin da aka fi so waɗanda aka sanya su a cikin kwandon shara tare da ƙudurin 10°; biri N: F = 2.12, p < 0.01; biri S: F = 2.01, p < 0.01). Wannan shi ne lamarin ga birai biyu (ANOVA; umarnin da aka fi so waɗanda aka sanya su a cikin kwandon shara tare da ƙudurin 10°; biri N: F = 2.12, p < 0.01; biri S: F = 2.01, p < 0.01). Это имело место для обеих обезьян (ANOVA; обезьяна N: F = 2,12, p <0,01; обезьяна S: F = 2,01, p <0,01). Wannan shi ne lamarin ga birai biyu (ANOVA; an haɗa umarnin da aka fi so a matakin ƙuduri na 10°; biri N: F=2.12, p<0.01; biri S: F=2.01, p<0.01).两只猴子都是这种情况(方差分析;差分析;以10° 0.01;猴子S:F = 2.01, p <0.01)两 只 猴子 都 是 这 种 (方差 分析 以 以 10 ° 分辨率 合并 的 相对 方向 ; 2 f. , p <0.01 ; : f = 2.01 , p <0.01….. Это имело место для обеих обезьян (ANOVA; F = 2,12, p <0,01; обезьяна S: F = 2,01, p <0,01). Wannan shi ne lamarin ga birai biyu (ANOVA; an haɗa yanayin da aka fi so a ƙudurin 10°; biri N: F=2.12, p<0.01; biri S: F=2.01, p<0.01).Ganin yawan bambancin da ke tsakanin jijiyoyi (Hoto na 5a), domin mu ga yadda jijiyoyi ke dogara da yanayin da aka fi so, da farko mun daidaita yanayin da kowace tantanin halitta ta fi so zuwa ga yanayin "mafi kyau" don motsi na tsarin grid (watau alkibla). wanda tsarin grid ya samar da mafi ƙarancin kusurwa tsakanin yanayin da aka fi so da kuma yanayin tsarin grating). Mun gano cewa iyakokin jijiyoyi (matsakaicin matakin "mafi muni" na lattice/matsakaicin matakin "mafi kyau" na lattice) sun bambanta da wannan yanayin da aka fi so, tare da kololuwa a cikin wannan rabon matakin da ke faruwa a kusa da alamu ko yanayin sassan (Hoto na 5b). )). Ba za a iya bayyana wannan tasirin ta hanyar nuna son kai a cikin rarrabawar alkiblar da aka fi so a cikin raka'o'in a cikin kowane samfurin zuwa ga ɗaya daga cikin tsarin plaid ko alkiblar sassan ba (Hoto na 5c; gwajin Rayleigh; biri N: z = 8.33, p < 10−3, matsakaicin zagaye = 190.13 deg ± 9.83 deg; biri S: z = 0.79, p = 0.45) kuma ya kasance daidai a tsakanin kusurwoyin plaid tsakanin grating (Hoto na Ƙari na 2). Ba za a iya bayyana wannan tasirin ta hanyar nuna son kai a cikin rarrabawar alkiblar da aka fi so a cikin raka'o'in a cikin kowane samfurin zuwa ga ɗaya daga cikin tsarin plaid ko alkiblar sassan ba (Hoto na 5c; gwajin Rayleigh; biri N: z = 8.33, p < 10−3, matsakaicin zagaye = 190.13 deg ± 9.83 deg; biri S: z = 0.79, p = 0.45) kuma ya kasance daidai a tsakanin kusurwoyin plaid tsakanin grating (Hoto na Ƙari na 2). Wannan shi ne karo na farko da za a gudanar da gasar cin kofin duniya a karo na biyu. сторону одного из клетчатых направлений или направлений компонентов (рис. 5в; критерий Рэлея; обезьяна N: 3 - 3 = 8, 3 z). Ba za a iya bayyana wannan tasirin ta hanyar sauyawa a cikin rarrabawar umarnin da aka fi so a cikin raka'a a cikin kowane samfurin zuwa ɗaya daga cikin kwatancen da aka duba ko umarnin sassan ba (Hoto na 5c; gwajin Rayleigh; biri N: z = 8.33, p < 10–3)., matsakaicin zagaye = digiri 190.13 ± digiri 9.83; biri S: z = 0.79, p = 0.45) kuma iri ɗaya ne ga dukkan kusurwoyin grid ɗin plaid (Hoto na Ƙari na 2).这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来解释(图5c;瑞利测试;猴子N:z = 8.33, p <10-3 ,圆形平均值= 190.13 度± 9.83 度;猴子S 0.45)并且在格子间光栅角上是一致的(补充图2)。不 效应 不 能 通过 每来 解释 (图 图 图 瑞利 测试 ; 猴子 n : z = 8.33 , p <10-3圆形 圆形 圆形 圆形 圆形 圆形 圆形 Wannan shi ne karo na farko da za a yi amfani da su a cikin dakin gwaje-gwajen. смещено либо в сторону структуры решетки, либо в сторону одной из ориентаций компонентов (рис. 5е; обезьяна N: z = 8,33, p <10-3). Ba za a iya bayyana wannan tasirin ta hanyar gaskiyar cewa rarrabawar abubuwan da aka fi so a cikin ƙwayoyin halitta a cikin kowane samfurin yana nuna karkacewa ko dai ga tsarin layi ko zuwa ga ɗaya daga cikin abubuwan da aka haɗa ba (Hoto na 5c; gwajin Rayleigh; biri N: z = 8.33, p < 10–3)., matsakaicin zagaye) = digiri 190.13 ± digiri 9.83; biri S: z = 0.79, p = 0.45) kuma sun yi daidai a kusurwoyin layi tsakanin grids (Hoto na Ƙari na 2).Saboda haka, saurin ji na jijiyoyi ga layukan rubutu ya dogara, aƙalla a wani ɓangare, akan mahimman halayen daidaita MT.
Bangaren hagu yana nuna rarrabawar rabon N/P (ƙasa ta jijiyoyi/hanyar tunani); kowace tantanin halitta tana ba da maki biyu na bayanai, ɗaya ga kowace alkibla inda tsarin ke motsawa. Bangaren dama yana nuna iyakokin psychophysical (ordinate) idan aka kwatanta da iyakokin neuronal (abscissa) ga dukkan raka'a a cikin samfurin. Bayanan da ke cikin layi na sama (ƙasa) sun fito ne daga biri N (S). b An tsara rabon iyakokin da aka daidaita bisa girman bambanci tsakanin mafi kyawun yanayin lattice da yanayin tantanin halitta da aka fi so. Alkiblar "mafi kyau" an ayyana ita ce alkiblar tsarin grating (an auna ta da grating guda ɗaya na sinusoidal) mafi kusa da alkiblar tantanin halitta da aka fi so. An fara tattara bayanai ta hanyar alkiblar da aka fi so (bankunan 10°), sannan aka daidaita rabon iyakokin zuwa matsakaicin ƙima kuma aka daidaita su a cikin kowane akwati. Kwayoyin da ke da alkiblar da aka fi so sun ɗan fi girma ko ƙanƙanta fiye da alkiblar sassan lattice suna da babban bambanci a cikin ji da yanayin lattice. c Histogram mai launin ruwan hoda na rarrabawar yanayin da aka fi so na duk raka'o'in MT da aka rubuta a cikin kowane biri.
A ƙarshe, martanin MT yana daidaitawa ta hanyar alkiblar motsin grating da cikakkun bayanai na siginar raba mu (launi). Kwatanta yanayin jijiyoyi da na psychophysical ya nuna cewa, gabaɗaya, raka'o'in MT ba su da saurin amsawa ga siginar rubutu masu bambanci fiye da birai. Duk da haka, yanayin jijiyoyi ya canza dangane da bambanci tsakanin yanayin da aka fi so na naúrar da alkiblar motsi. Kwayoyin da suka fi saurin amsawa sun fi saurin amsawa waɗanda suka kusan rufe tsarin lattice ko ɗaya daga cikin yanayin da aka haɗa, kuma ƙaramin rukuni na samfuranmu sun fi saurin amsawa ko kuma sun fi saurin amsawa fiye da yadda birai ke fahimtar bambance-bambancen bambanci. Don tantance ko siginar daga waɗannan raka'o'in da suka fi saurin amsawa suna da alaƙa da fahimta a cikin birai, mun bincika alaƙar da ke tsakanin fahimta da martanin jijiyoyi.
Wani muhimmin mataki wajen kafa alaƙa tsakanin ayyukan jijiyoyi da ɗabi'a shine a kafa alaƙa tsakanin jijiyoyi da martanin ɗabi'a ga abubuwan da ke motsa jiki akai-akai. Domin a haɗa martanin jijiyoyi da hukunce-hukuncen rarrabuwa, yana da mahimmanci a ƙirƙiri wani abin ƙarfafawa wanda, duk da cewa iri ɗaya ne, ana ganinsa daban-daban a cikin gwaje-gwaje daban-daban. A cikin wannan binciken, wannan an wakilta shi a sarari ta hanyar grating na sifili. Kodayake mun jaddada cewa, bisa ga ayyukan tunani na dabbobi, gratings masu ƙarancin (ƙasa da ~20%) bambancin rubutu galibi ana ɗaukar su a matsayin masu haɗin kai ko bayyananne.
Domin auna yadda amsoshin MT suka yi daidai da rahotannin fahimta, mun yi nazarin yiwuwar zaɓi (CP) na bayanan grid ɗinmu (duba 3). A takaice, CP ma'auni ne mara mizani, wanda ba na yau da kullun ba wanda ke auna alaƙar da ke tsakanin amsoshin ƙaruwa da hukunce-hukuncen fahimta30. Taƙaita nazarin zuwa gwaje-gwaje ta amfani da grid tare da sifili bambancin rubutu da zaman da birai suka yi aƙalla zaɓi biyar ga kowane nau'in waɗannan gwaje-gwajen, mun ƙididdige SR daban-daban don kowane alkibla na motsi na grid. A cikin birai, mun lura da matsakaicin ƙimar CP mafi girma fiye da yadda muke tsammani ta hanyar kwatsam (Hoto na 6a, d; biri N: matsakaicin CP: 0.54, 95% CI: (0.53, 0.56), gwajin t-gefe biyu akan null na CP = 0.5, t = 6.7, p < 10−9; biri S: matsakaicin CP: 0.55, 95% CI: (0.54, 0.57), gwajin t-gefe biyu, t = 9.4, p < 10−13). A cikin birai, mun lura da matsakaicin ƙimar CP mafi girma fiye da yadda muke tsammani ta hanyar kwatsam (Hoto na 6a, d; biri N: matsakaicin CP: 0.54, 95% CI: (0.53, 0.56), gwajin t mai gefe biyu akan null na CP = 0.5, t = 6.7, p < 10−9; biri S: matsakaicin CP: 0.55, 95% CI: (0.54, 0.57), gwajin t mai gefe biyu, t = 9.4, p < 10−13).A cikin birai, mun lura da matsakaicin CP ya fi girma fiye da yadda aka zata bazuwar (Hoto na 6a, d; biri N: ma'anar CP: 0.54, 95% CI: (0.53, 0.56), gwajin t-mai wutsiya biyu da ƙimar mara kyau).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) . biri S: matsakaicin CP: 0.55, 95% CI: (0.54, 0.57), gwajin t mai wutsiya biyu, t = 9.4, p < 10–13).在猴子中。 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,: 95.5双边t 检验, t = 9.4, p <10−13).在 猴子 中 , 我们 观察 平均 平均 值 显着 大于0.54, 95% Ci: 0.53 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 обезьяна N: среднее CP: 0,54, 95% ДИ: (0,53, 0,56), двусторонний t- тест CP против нуля = 0,5, t = 6,7-p, p < 10; CP: 0,55, 95% ДИ: (0,54, 0,57), двусторонний t-критерий, t = 9,4, p <10- 13) . A cikin birai, mun lura da matsakaicin ƙimar CP sama da abin da za mu iya tsammani ta hanyar kwatsam (Hoto na 6a, d; biri N: ma'anar CP: 0.54, 95% CI: (0.53, 0.56), gwajin t-test mai wutsiya biyu vs. sifili = 0.5, t = 6.7, p < 10-9, biri S: ma'anar CP: 0.55, 95% CI: (0.54, 0.57), ma'aunin t-wutsiya biyu, t = 9.4, p < 10-13).Saboda haka, ƙwayoyin halittar MT suna ƙara yin ƙarfi ko da babu wata alama ta rarrabuwa a fili, lokacin da fahimtar dabba game da motsin layin ya dace da fifikon ƙwayar halitta.
Rarraba yiwuwar zaɓi don grids ba tare da siginar rubutu ba don samfuran da aka rubuta daga biri N. Kowace tantanin halitta na iya ba da gudummawa har zuwa maki biyu na bayanai (ɗaya ga kowane alkibla na motsi na grid). Matsakaicin ƙimar CP sama da kibiyoyi fari bazuwar (farin kibiyoyi) yana nuna cewa gabaɗaya akwai muhimmiyar alaƙa tsakanin aikin MT da fahimta. b Don bincika tasirin duk wani zaɓi mai yuwuwar nuna bambanci, mun ƙididdige CP daban don duk wani abin ƙarfafawa wanda birai suka yi aƙalla kuskure ɗaya. An tsara yiwuwar zaɓi azaman aikin rabon zaɓi (pref/null) don duk abubuwan ƙarfafawa (hagu) da ƙimar cikakkiyar bambancin alamar rubutu (dama, bayanai daga ƙwayoyin halitta 120). Layin mai ƙarfi da yankin inuwa a cikin allon hagu suna wakiltar matsakaicin ± sem na matsakaicin motsi na maki 20. Yiwuwar zaɓi da aka ƙididdige don abubuwan ƙarfafawa tare da rabon zaɓi mara daidaito, kamar grids tare da babban bambancin sigina, sun bambanta sosai kuma an haɗa su a kusa da yuwuwar. Yankin da aka yi wa inuwa launin toka a cikin allon dama yana jaddada bambancin fasalulluka da aka haɗa a cikin lissafin babban yuwuwar zaɓi. c An zana yiwuwar babban zaɓi (ordinate) a kan iyakar jijiyoyi (abscissa). Yiwuwar zaɓi ya kasance yana da alaƙa da maƙasudi mara kyau. Tsarin df iri ɗaya ne da ac amma yana aiki ga bayanai guda 157 daga biri S sai dai idan an lura da wani abu daban. g Mafi girman yuwuwar zaɓi (ordinate) an zana shi akan alkiblar da aka fi so (abscissa) ga kowane birai biyu. Kowane tantanin halitta na MT ya ba da gudummawar maki biyu na bayanai (ɗaya ga kowane alkibla na tsarin lattice). h Manyan zane-zanen akwatin yuwuwar zaɓi ga kowane kusurwar inter-raster. Layin mai ƙarfi yana nuna matsakaici, ƙananan gefuna na akwatin suna wakiltar kashi 25 da 75 bi da bi, an faɗaɗa gashin baki zuwa sau 1.5 na kewayon interquartile, kuma an lura da abubuwan da suka wuce wannan iyaka. Bayanan da ke cikin bangarorin hagu (dama) sun fito ne daga ƙwayoyin biri N(S) guda 120 (157). i Mafi girman yuwuwar zaɓi (ordinate) an zana shi akan lokacin fara motsawa (abscissa). An ƙididdige babban CP a cikin murabba'ai masu zamiya (faɗin 100 ms, mataki na 10 ms) a cikin gwajin sannan aka auna shi a kan raka'a.
Wasu bincike da aka yi a baya sun ba da rahoton cewa CP ya dogara ne da adadin gwaje-gwajen da aka yi a cikin rarrabawar ƙimar basal, ma'ana wannan ma'aunin ba shi da inganci ga abubuwan da ke haifar da manyan bambance-bambance a cikin rabon kowane zaɓi. Don gwada wannan tasirin a cikin bayananmu, mun ƙididdige CP daban-daban don duk abubuwan da ke haifar da sakamako, ba tare da la'akari da bambancin yanayin alamun ba, kuma birai sun yi aƙalla gwajin ƙarya guda ɗaya. An tsara CP akan rabon zaɓi (pref/null) ga kowane dabba a Hoto na 6b da e (a ɓangaren hagu), bi da bi. Idan aka duba matsakaicin motsi, a bayyane yake cewa CP ya kasance sama da yuwuwar a kan nau'ikan zaɓuɓɓuka masu yawa, yana raguwa ne kawai lokacin da damar ta faɗi (ƙaruwa) a ƙasa (sama) 0.2 (0.8). Dangane da halayen psychometric na dabbobi, za mu yi tsammanin ma'aunin zaɓi na wannan girman zai shafi abubuwan da ke haifar da sakamako masu bambanci (masu jituwa ko masu haske) (duba misalan fasalulluka na psychometric a Hoto na 2a, b). Domin tantance ko haka lamarin yake kuma ko wani babban PC ya ci gaba har ma da abubuwan ƙarfafawa tare da siginar rarrabuwa bayyanannu, mun bincika tasirin ƙimar bambancin rubutu cikakke akan PC (Hoto na 6b, e-dama). Kamar yadda aka zata, CP ya fi girma fiye da yuwuwar abubuwan ƙarfafawa waɗanda ke ɗauke da alamun rarrabuwa har zuwa matsakaici (~20% bambanci ko ƙasa da haka).
A cikin ayyukan gane yanayin, saurin gudu, da rashin daidaito, MT CP ya fi yawa a cikin ƙwayoyin jijiyoyi masu saurin fahimta, wataƙila saboda waɗannan ƙwayoyin jijiyoyi suna ɗauke da sigina mafi yawan bayanai30,32,33,34.Daidai da waɗannan binciken mun lura da wata alaƙa mai sauƙi amma mai mahimmanci tsakanin babban CP, wanda aka ƙididdige daga ƙimar harbi da aka zira a cikin bambancin yanayin rubutu wanda aka nuna a cikin sashin dama na Hoto.6b, e, da kuma iyakar jijiyoyi (Hoto na 6c, f; matsakaicin juyi na geometric; biri N: r = −0.12, p = 0.07 biri S: r = −0.18, p < 10−3). 6b, e, da kuma iyakar jijiyoyi (Hoto na 6c, f; matsakaicin juyi na geometric; biri N: r = −0.12, p = 0.07 biri S: r = −0.18, p < 10−3).Daidai da waɗannan binciken, mun lura da ƙaramin alaƙa mai mahimmanci tsakanin babban CP da aka ƙididdige daga ƙimar z-mita na motsawa daga bambancin siginar rubutu da aka nuna a cikin sashin da ke kusa da dama na Hoto na 6b, e, da kuma iyakar jijiyoyi (Hoto na 6c, f; na lissafi). matsakaicin juzu'i na geometric;обезьяна N: r = -0,12, p = 0,07 обезьяна S: r = -0,18, p <10-3). biri N: r = -0.12, p = 0.07 biri S: r = -0.18, p < 10-3).与这些发现一致,我们观察到大CP 之间存在适度但显着的相关性,这是根据图6、和神经元阈值(图6c、f;几何平均回归;猴子N:r = -0.12,p = 0.07 猴子S:r , -0.18与 些 发现 一致 , 我们 到 大 大 之间 存在 适度 但和 元 阈值 (图 图 6c 、 f ; 回归 ; 猴子 n ; 猴子 n : r = -0.12 , p = 0.07 猴子 Daidai da waɗannan binciken, mun lura da ƙaramin alaƙa amma mai mahimmanci tsakanin manyan CVs kamar yadda aka nuna a Hoto na 6b,e da iyakokin neuron (Hoto na 6c,f; matsakaicin juyi na geometric; biri N: r = -0.12, p = 0.07).Обезьяна S: г = -0,18, р <10-3). Biri S: r = -0.18, p < 10-3).Saboda haka, alamu daga sassa mafi ilimi sun fi nuna bambanci sosai da hukunce-hukuncen rarrabuwar kai a cikin birai, wanda yake da mahimmanci ba tare da la'akari da duk wani alamun rubutu da aka ƙara wa son zuciya ba.
Ganin cewa mun riga mun kafa dangantaka tsakanin jin daɗin siginar rubutu da kuma yanayin jijiyoyi da aka fi so, mun yi mamakin ko akwai irin wannan dangantaka tsakanin CP da yanayin da aka fi so (Hoto na 6g). Wannan alaƙar ta ɗan yi tasiri kaɗan a cikin biri S (ANOVA; biri N: 1.03, p=0.46; biri S: F=1.73, p=0.04). Ba mu ga wani bambanci a cikin CP ba don kusurwoyin layi tsakanin layi a cikin kowace dabba (Hoto na 6h; ANOVA; biri N: F = 1.8, p = 0.11; biri S: F = 0.32, p = 0.9).
A ƙarshe, aikin da ya gabata ya nuna cewa CP yana canzawa a duk lokacin gwajin. Wasu bincike sun ba da rahoton ƙaruwa mai kaifi tare da tasirin zaɓi mai laushi,30 yayin da wasu suka ba da rahoton ƙaruwa mai ɗorewa a cikin siginar zaɓi a cikin gwajin31. Ga kowane biri, mun ƙididdige CP na kowane raka'a a cikin gwaje-gwaje tare da sifili bambancin rubutu (bi da bi, bisa ga yanayin tsari) a cikin ƙwayoyin ms 100 waɗanda ke tafiya kowane ms 20 daga farkon farawa zuwa matsakaicin matsakaicin ƙaddamarwar motsawa. An nuna matsakaicin yanayin CP na birai biyu a cikin Hoto na 6i. A cikin duka halayen, CP ya kasance a matakin bazuwar ko kusa da shi har zuwa kusan ms 500 bayan farawar motsawar, bayan haka CP ya ƙaru sosai.
Baya ga canjin yanayin ji, an nuna cewa CP yana da wasu halaye na halayen daidaita ƙwayoyin halitta. Misali, Uka da DeAngelis34 sun gano cewa CP a cikin aikin gane rashin daidaiton binocular ya dogara ne akan daidaiton lanƙwasa daidaita rashin daidaiton binocular na na'urar. A wannan yanayin, tambaya mai alaƙa ita ce ko ƙwayoyin zaɓin alkiblar tsari (PDS) sun fi ƙarfin alkiblar sashi (CDS). Kwayoyin PDS suna ɓoye alkiblar gabaɗaya na alamu waɗanda ke ɗauke da alkiblar gida da yawa, yayin da ƙwayoyin CDS ke amsawa ga motsi na sassan tsarin alkibla (Hoto na 7a).
Wakiltar tsari na gyaran yanayin motsa jiki da kuma daidaita lanƙwasa na hasashe (hagu) da kuma daidaita lanƙwasa na daidaita lanƙwasa (dama) (duba Kayan Aiki da Hanyoyi). A takaice, idan tantanin halitta ya haɗu a cikin sassan grid don motsa lanƙwasa na sigina, mutum zai yi tsammanin lanƙwasa iri ɗaya don daidaita lanƙwasa na grid da abubuwan ƙarfafawa na grid (shafi na ƙarshe, lanƙwasa mai ƙarfi). Akasin haka, idan tantanin halitta bai haɗa alkiblar abubuwan haɗin cikin motsi na siginar ba, mutum zai yi tsammanin lanƙwasa na daidaitawa biyu tare da kololuwa a kowane alkibla na motsi na grating wanda ke fassara sashi ɗaya zuwa alkiblar da tantanin halitta ta fi so (shafi na ƙarshe, lanƙwasa mai lanƙwasa). . lanƙwasa b (hagu) don daidaita yanayin jerin sinusoidal don ƙwayoyin da aka nuna a cikin Hotuna na 1 da 2. 3 da 4 (layi na sama - ƙwayoyin a cikin Hotuna na 3a, b da 4a, b (sama); ɓangaren ƙasa - ƙwayoyin Hoto na 3c, d da 4a, b (ƙasa)). (Tsakiya) Tsarin da hasashen sassan da aka ƙididdige daga bayanan gyaran lattice. (Dama) Daidaita grid na waɗannan ƙwayoyin. An rarraba ƙwayoyin saman (ƙasa) a matsayin ƙwayoyin samfuri (bangaren). Lura cewa babu wata alaƙa tsakanin rarrabuwar sassan tsarin da kuma fifikon motsi na ƙwayoyin da ke da alaƙa/bayyananne (duba amsoshin layin rubutu don waɗannan ƙwayoyin a Hoto na 4a). c Ma'aunin haɗin ɓangaren yanayin z-score (ordinate) wanda aka zana akan ma'aunin haɗin ɓangaren ɓangaren z-score (abscissa) don duk ƙwayoyin da aka rubuta a cikin birai N (hagu) da S (dama). Layuka masu kauri suna nuna ma'aunin mahimmanci da ake amfani da su don rarraba ƙwayoyin. d Tsarin babban yuwuwar zaɓi (ordinate) idan aka kwatanta da ma'aunin yanayi (Zp - Zc) (abscissa). Bayanai a cikin bangarorin hagu (dama) suna nufin biri N(S). Da'irori baƙi suna nuna bayanai a cikin raka'o'i kimanin. A cikin dabbobi biyu, akwai muhimmiyar alaƙa tsakanin babban yuwuwar zaɓi da ma'aunin tsari, yana nuna babban haɗin fahimta ga ƙwayoyin da ke da ma'aunin sigina a cikin abubuwan da ke haifar da yanayin da ke da ma'anar abubuwa da yawa.
Saboda haka, a cikin wani gwaji daban, mun auna martanin da aka bayar ga grids da grids na sinusoidal don rarraba neurons a cikin samfuranmu a matsayin PDS ko CDS (duba Hanyoyi). Lanƙwasa na daidaita lattice, hasashen sassan samfuri da aka gina daga wannan bayanan daidaitawa, da lanƙwasa na daidaita lattice don ƙwayoyin da aka nuna a cikin Figures 1 da 3. Hotuna 3 da 4 da Ƙarin Hoto na 3 an nuna su a cikin Hoto na 7b. An nuna rarraba tsari da zaɓin sassan, da kuma yanayin da aka fi so a cikin kowane rukuni, ga kowane biri a cikin Hoto na 7c da ƙarin Hoto na 4 bi da bi.
Domin tantance dogaro da CP akan gyaran sassan tsarin, mun fara ƙididdige ma'aunin tsarin 35 (PI), manyan ƙimar (ƙananan) waɗanda ke nuna babban ɗabi'ar PDS (CDS). Ganin yadda aka nuna a sama cewa: (i) ƙarfin jijiyoyi ya bambanta da bambanci tsakanin yanayin da aka fi so na ƙwayar halitta da kuma alkiblar motsin motsa jiki, kuma (ii) akwai muhimmiyar alaƙa tsakanin ƙarfin jijiyoyi da yuwuwar zaɓi a cikin samfurinmu, mun sami alaƙa tsakanin PI da jimlar CP da aka yi nazari a kai don "mafi kyawun" alkiblar motsi na kowace ƙwayar halitta (duba sama). Mun gano cewa CP yana da alaƙa sosai da PI (Hoto na 7d; matsakaicin juyi na geometric; babban biri na CP N: r = 0.23, p < 0.01; bi-stable CP biri N r = 0.21, p = 0.013; babban biri na CP S: r = 0.30, p < 10−4; bi-stable CP biri S: r = 0.29, p < 10−3), yana nuna cewa ƙwayoyin da aka rarraba a matsayin PDS sun nuna mafi girman aiki da ya shafi zaɓi fiye da CDS da ƙwayoyin da ba a rarraba ba. Mun gano cewa CP yana da alaƙa sosai da PI (Hoto na 7d; matsakaicin juyi na geometric; babban biri na CP N: r = 0.23, p < 0.01; bi-stable CP biri N r = 0.21, p = 0.013; babban biri na CP S: r = 0.30, p < 10−4; bi-stable CP biri S: r = 0.29, p < 10−3), yana nuna cewa ƙwayoyin da aka rarraba a matsayin PDS sun nuna mafi girman aiki da ya shafi zaɓi fiye da CDS da ƙwayoyin da ba a rarraba ba. Мы обнаружили, что CP значительно корррелирует с PI (рис. 7d; 0,23, p <0,01; бистабильная обезьяна CP N r = 0,21, p = 0,013; большая обезьяна CP S: r = 0,30, p <10-4; = 0,29, p <10-3), что указывает на то, что клетки, классифицированные как PDS, проявляли большую активность, связанную с выborом, чем CDS kletky. Mun gano cewa CP yana da alaƙa sosai da PI (Hoto na 7d; matsakaicin juyi na geometric; babban biri CP N: r = 0.23, p < 0.01; bistable biri CP N r = 0.21, p = 0.013; babban biri CP S: r = 0.30, p < 10-4; biri S bistable CP: r = 0.29, p < 10-3), yana nuna cewa ƙwayoyin da aka rarraba a matsayin PDS sun nuna ƙarin aiki, waɗanda ke da alaƙa da zaɓi fiye da CDS da ƙwayoyin da ba a rarraba ba.我们发现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;双焳0.21,p = 0.013;大CP 猴S: r = 0.30,p <10-4 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 0.21, p3; 0.0 Мы обнаружили, что CP был значительно связан с PI (рис. 7d; 0,23, p <0,01; бистабильная обезьяна CP N r = 0,21, p = 0,013; большая обезьяна CP S: r = 0,013) 0,30, p <10-10. Mun gano cewa CP yana da alaƙa mai mahimmanci da PI (Hoto na 7d; matsakaicin juzu'i na geometric; babban biri CP N: r = 0.23, p < 0.01; bistable biri CP N r = 0.21, p = 0.013; babban biri 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, класссифипицированные большую селекционную активность, чем клетки, класссифицированные как CDS da неклассифицированые. biri S bistable CP: r = 0.29, p < 10-3), yana nuna cewa ƙwayoyin da aka rarraba a matsayin PDS sun nuna ayyukan zaɓi mafi girma fiye da ƙwayoyin da aka rarraba a matsayin CDS kuma ba a rarraba su ba.Saboda duka PI da neuron suna da alaƙa da CP, mun yi nazarin komawa baya da yawa (tare da PI da neuron masu canzawa a matsayin masu canzawa masu zaman kansu da kuma babban CP a matsayin masu canzawa masu dogaro) don kawar da alaƙa tsakanin ma'aunin guda biyu wanda ke haifar da yuwuwar tasiri. Dukansu ma'aunin haɗin kai na ɓangare sun kasance masu mahimmanci (biri N: ma'auni vs. CP: r = −0.13, p = 0.04, PI vs. CP: r = 0.23, p < 0.01; biri S: ma'auni vs. CP: r = −0.16, p = 0.03, PI vs CP: 0.29, p < 10−3), yana nuna cewa CP yana ƙaruwa tare da jin daɗi kuma a cikin yanayi mai zaman kansa yana ƙaruwa tare da PI. Dukansu ma'aunin haɗin kai na ɓangare sun kasance masu mahimmanci (biri N: ma'auni vs. CP: r = −0.13, p = 0.04, PI vs. CP: r = 0.23, p < 0.01; biri S: ma'auni vs. CP: r = −0.16, p = 0.03, PI vs CP: 0.29, p < 10−3), yana nuna cewa CP yana ƙaruwa tare da jin daɗi kuma a cikin yanayi mai zaman kansa yana ƙaruwa tare da PI. Оба частныh коэффициента kорреляции были значимыmy ( N: порог против CP: r = -0,13, p = 0,04, PI 0,04, PI 0,04, PI 0,04, PI 0,04, PI 0,04, PI. <0,01; обезьяна S: порог против CP: r = -0,16, p = 0,03, PI vs CP: 0,29, p <10-3), da CP увеличи чувствительностью и независимым образом увеличивается с PI. Dukansu ma'aunin haɗin kai na ɓangare sun kasance masu mahimmanci (biri N: ma'auni vs. CP: r=-0.13, p=0.04, PI vs. CP: r=0.23, p<0.01; biri S: ma'auni vs. CP: r = -0.16, p = 0.03, PI vs CP: 0.29, p < 10-3), yana nuna cewa CP yana ƙaruwa tare da jin daɗi kuma yana ƙaruwa da kansa tare da 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;猴子S:阈值与CP:r = -0.16, p = 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 <10CP Оба частныh коэффициента kорреляции были значимыmy ( N: порог против CP: r = -0,13, p = 0,04, PI 0,04, PI 0,04, PI 0,04, PI 0,04, PI 0,04, PI. <0,01; обезьяна S: порог против CP: r = -0,16, p = 0,03 чувствительностью и увеличивалась с PI независимым образом. Dukansu ma'aunin haɗin kai na ɓangare sun kasance masu mahimmanci (biri N: ma'auni vs. CP: r=-0.13, p=0.04, PI vs. CP: r=0.23, p<0.01; biri S: ma'auni vs. CP: r = -0.16, p = 0.03, PI vs CP: 0.29, p < 10-3), yana nuna cewa CP ya ƙaru da hankali kuma ya ƙaru da PI ta hanyar da ba ta da tushe.
Mun yi rikodin aiki guda ɗaya a yankin MT, kuma birai sun ba da rahoton fahimtarsu game da alamu waɗanda za su iya bayyana a matsayin motsi masu daidaituwa ko bayyananne. Jin daɗin jijiyoyi ga alamun rarrabuwa da aka ƙara wa fahimtar son zuciya ya bambanta sosai kuma ana tantance shi, aƙalla a wani ɓangare, ta hanyar alaƙar da ke tsakanin yanayin da aka fi so na sashin da alkiblar motsin ƙarfafawa. A cikin dukkan jama'a, jin daɗin jijiyoyi ya yi ƙasa da na psychophysical, kodayake raka'o'in da suka fi dacewa sun yi daidai ko sun wuce yanayin halayyar siginar rarrabuwa. Bugu da ƙari, akwai babban bambanci tsakanin mitar harbi da fahimta, yana nuna cewa siginar MT tana taka rawa a cikin rarrabuwa. Kwayoyin da ke da yanayin da aka fi so sun inganta yanayin su ga bambance-bambance a cikin siginar rarrabuwa ta layi kuma suna nuna alamar motsi na duniya a cikin abubuwan da ke haifar da yanayi na gida da yawa, suna nuna mafi girman alaƙar fahimta. A nan muna la'akari da wasu matsaloli masu yuwuwa kafin kwatanta waɗannan sakamakon da aikin da ya gabata.
Babbar matsala da bincike ke amfani da abubuwan da ke motsa jiki na bistable a cikin samfuran dabbobi ita ce martanin ɗabi'a ba zai dogara da girman abin sha'awa ba. Misali, birai namu za su iya bayar da rahoton fahimtarsu game da yanayin rubutu ba tare da la'akari da fahimtarsu game da daidaiton layi ba. Abubuwa biyu na bayanan sun nuna cewa ba haka lamarin yake ba. Na farko, daidai da rahotannin da suka gabata, canza kusurwar yanayin da ke tsakanin sassan rarrabuwa ta hanyar tsari ya canza yuwuwar fahimtar juna. Na biyu, a matsakaici, tasirin iri ɗaya ne ga alamu waɗanda ke ɗauke da ko ba su ƙunshi siginar rubutu ba. Idan aka haɗa su, waɗannan abubuwan lura sun nuna cewa martanin biri koyaushe yana nuna fahimtarsu game da haɗin kai/bayyananne.
Wata matsala mai yuwuwa ita ce ba mu inganta sigogin motsin grating don takamaiman yanayin ba. A cikin ayyukan da suka gabata da yawa waɗanda suka kwatanta ƙarfin jijiyoyi da na psychophysical, an zaɓi abubuwan ƙarfafawa daban-daban ga kowane rukunin da aka yi rijista [31, 32, 34, 36, 37, 38, 39, 40, 41, 42, 43, 44, 45]. A nan mun yi amfani da hanyoyi guda biyu na motsi na tsarin lattice ba tare da la'akari da daidaitawar yanayin kowace tantanin halitta ba. Wannan ƙira ta ba mu damar yin nazarin yadda ƙarfin ji ya canza tare da haɗuwa tsakanin motsi na lattice da yanayin da aka fi so, duk da haka, bai samar da tushen farko ba don tantance ko ƙwayoyin sun fi son lattice masu haɗin kai ko masu haske. Saboda haka, muna dogara da ƙa'idodin gwaji, ta amfani da martanin kowane tantanin halitta zuwa raga mai laushi, don sanya fifiko da sifili lakabi ga kowane rukuni na motsi na raga. Kodayake ba zai yiwu ba, wannan na iya karkatar da sakamakon nazarin ƙarfin ji da gano siginar CP ta hanyar tsari, yana iya wuce gona da iri kan kowane ma'auni. Duk da haka, fannoni da yawa na bincike da bayanai da aka tattauna a ƙasa sun nuna cewa ba haka lamarin yake ba.
Da farko, sanya sunayen da aka fi so (marasa amfani) ga abubuwan da ke haifar da ƙarin aiki (ƙasa) bai shafi bambance-bambancen waɗannan rarrabawar amsawar ba. Madadin haka, yana tabbatar da cewa ayyukan neurometric da psychometric suna da alama iri ɗaya, don haka za a iya kwatanta su kai tsaye. Na biyu, amsoshin da aka yi amfani da su don ƙididdige CP (gwaji "mara kyau" don gratings masu laushi da duk gwaje-gwaje don gratings ba tare da bambancin rubutu ba) ba a haɗa su a cikin nazarin komawa baya wanda ya ƙayyade ko kowace tantanin halitta "ta fi son" wasanni masu alaƙa ko masu gaskiya. Wannan yana tabbatar da cewa tasirin zaɓi ba ya karkata ga sunayen da aka fi so/marasa inganci, wanda ke haifar da yuwuwar zaɓi mai mahimmanci.
Nazarin Newsom da abokan aikinsa [36, 39, 46, 47] su ne na farko da suka tantance rawar da MT ke takawa a kimantawa na alkiblar motsi. Rahotannin da suka biyo baya sun tattara bayanai kan shiga MT a zurfin34,44,48,49,50,51 da sauri32,52, kyakkyawan tsari33 da fahimtar tsarin 3D daga motsi31,53,54 (dazuzzukan 3D masu dorewa). don mulki). Muna faɗaɗa waɗannan sakamakon ta hanyoyi biyu masu mahimmanci. Na farko, muna ba da shaida cewa martanin MT suna ba da gudummawa ga rarrabuwar siginar visuomotor. Na biyu, mun lura da alaƙa tsakanin zaɓin yanayin MT da wannan siginar zaɓi.
A ra'ayi, sakamakon yanzu ya fi kama da aikin da aka yi akan 3-D SFM, domin duka biyun fahimta ce mai rikitarwa wacce ta shafi tsari da zurfin motsi. Dodd et al.31 sun sami babban yuwuwar zaɓi (0.67) a cikin aikin biri wanda ke ba da rahoton yanayin juyawa na silinda 3D SFM ta bistable. Mun sami ƙaramin tasirin zaɓi don abubuwan ƙarfafa grid na bistable (kimanin 0.55 ga birai biyu). Tunda kimantawar CP ta dogara da ma'aunin zaɓi, yana da wuya a fassara CP da aka samu a ƙarƙashin yanayi daban-daban a cikin ayyuka daban-daban. Duk da haka, girman tasirin zaɓi da muka lura iri ɗaya ne ga sifili da ƙarancin zane-zanen bambanci, kuma lokacin da muka haɗa ƙananan/babu abubuwan ƙarfafawa na rubutu don ƙara ƙarfi. Saboda haka, wannan bambanci a cikin CP ba zai yiwu ba saboda bambance-bambance a cikin ƙimar zaɓi tsakanin bayanai.
Ƙananan canje-canje a cikin ƙimar harbin MT waɗanda ke tare da fahimta a cikin yanayin na ƙarshe sun zama abin mamaki idan aka kwatanta da yanayin fahimta mai ƙarfi da inganci wanda ke haifar da motsawar SFM 3-D da tsarin grid bistable. Wata yuwuwar ita ce mun raina tasirin zaɓi ta hanyar ƙididdige ƙimar harbin a duk tsawon lokacin da aka yi amfani da shi. Sabanin lamarin 31 3-D SFM, inda bambance-bambance a cikin ayyukan MT suka haɓaka kusan 250 ms a cikin gwaje-gwaje sannan suka ƙaru a hankali a duk lokacin gwaje-gwajen, bincikenmu game da yanayin motsa jiki na lokaci-lokaci na siginar zaɓi (duba 500 ms bayan fara motsawa a cikin birai biyu. Bugu da ƙari, bayan ƙaruwa mai ƙarfi a wannan lokacin, mun lura da canje-canje a cikin CP a lokacin sauran gwajin. Hupe da Rubin55 sun ba da rahoton cewa fahimtar ɗan adam game da jerin murabba'i masu sassauƙa sau da yawa yana canzawa a lokacin gwaje-gwaje masu tsawo. Kodayake an gabatar da abubuwan ƙarfafawa namu na daƙiƙa 1.5 kawai, fahimtar birai namu kuma na iya bambanta daga daidaituwa zuwa bayyananne yayin gwajin (amsoshinsu sun nuna fahimtarsu ta ƙarshe a lokacin zaɓin alama.) Saboda haka, ana sa ran sigar lokacin amsawa ta aikinmu, ko shirin da birai za su iya ci gaba da ba da rahoton fahimtarsu, za ta sami babban tasirin zaɓi. Yiwuwar ƙarshe ita ce ana karanta siginar MT daban-daban a cikin ayyukan biyu. Kodayake an daɗe ana tunanin cewa siginar CPU tana haifar da gano yanayin ji da hayaniyar da ke da alaƙa,56 Gu da abokan aiki57 sun gano cewa a cikin samfuran lissafi, dabarun haɗuwa daban-daban, maimakon matakan haɗin gwiwa. Bambanci, zai iya bayyana CPU mafi kyau a cikin ƙwayoyin jijiyoyi na dorsal media-superior temporal neurons. aikin gane yanayin canza takarda (MSTd). Ƙaramin tasirin zaɓi da muka lura a cikin MT wataƙila yana nuna tarin ƙwayoyin jijiyoyi masu ƙarancin bayanai don ƙirƙirar fahimtar daidaito ko bayyana gaskiya. A duk lokacin da za a haɗa alamun motsi na gida zuwa abubuwa ɗaya ko biyu (bistable gratings) ko kuma saman abubuwa daban-daban (3-D SFM), shaida mai zaman kanta cewa amsoshin MT suna da alaƙa sosai da hukunce-hukuncen fahimta, akwai amsoshin MT masu ƙarfi. An ba da shawarar taka rawa wajen rarraba hotuna masu rikitarwa zuwa wuraren abubuwa da yawa ta amfani da bayanan motsin gani.
Kamar yadda aka ambata a sama, mu ne farkon waɗanda suka ba da rahoton alaƙa tsakanin aikin ƙwayoyin halitta na tsarin MT da fahimta. Kamar yadda aka tsara a cikin samfurin mataki biyu na asali ta Movshon da abokan aikinsa, sashin yanayin shine matakin fitarwa na MT. Duk da haka, aikin da aka yi kwanan nan ya nuna cewa ƙwayoyin yanayi da sassan suna wakiltar ƙarshen daban-daban na ci gaba kuma cewa bambance-bambancen parametric a cikin tsarin filin karɓa suna da alhakin daidaita bakan abubuwan da ke cikin yanayin. Saboda haka, mun sami babban alaƙa tsakanin CP da PI, kama da alaƙar da ke tsakanin daidaiton daidaitawar binocular da CP a cikin aikin gane zurfin ko saitin saitin jagora a cikin aikin nuna bambancin shugabanci mai kyau. Alaƙa tsakanin takardu da CP 33. Wang da Movshon62 sun bincika adadi mai yawa na ƙwayoyin halitta tare da zaɓin daidaitawar MT kuma sun gano cewa, a matsakaici, ma'aunin yanayin yana da alaƙa da kaddarorin daidaitawa da yawa, yana nuna cewa zaɓin yanayi yana wanzu a cikin wasu nau'ikan sigina da yawa waɗanda za a iya karantawa daga yawan MT. Saboda haka, don nazarin da za a yi nan gaba game da alaƙar da ke tsakanin ayyukan MT da fahimtar mutum, zai zama da muhimmanci a tantance ko ma'aunin tsarin ya yi daidai da sauran siginar zaɓi na aiki da motsa jiki, ko kuma ko wannan alaƙar ta musamman ce ga yanayin rarrabuwar fahimta.
Hakazalika, Nienborg da Cumming 42 sun gano cewa duk da cewa ƙwayoyin da ke kusa da nesa waɗanda ke zaɓar rashin daidaiton binocular a cikin V2 sun kasance masu saurin kamuwa da cutar a cikin aikin wariyar launin fata, yawan ƙwayoyin da suka fi so ne kawai suka nuna babban CP. Duk da haka, sake horar da birai don bambance-bambancen nisa da suka fi so ya haifar da manyan CPs a cikin keji da aka fi so. Wasu nazarin kuma sun ba da rahoton cewa tarihin horo ya dogara ne akan alaƙar fahimta34,40,63 ko alaƙar dalili tsakanin aikin MT da bambancin wariya48. Alaƙar da muka lura tsakanin CP da zaɓin alkiblar tsarin yana nuna takamaiman dabarun da birai suka yi amfani da su don magance matsalarmu, ba takamaiman rawar da siginar zaɓin yanayi ke takawa a cikin fahimtar gani da motsi ba. A cikin aikin gaba, zai zama mahimmanci a tantance ko tarihin ilmantarwa yana da tasiri mai mahimmanci akan tantance waɗanne siginar MT ne aka fi so kuma a sassauƙa don yin yanke hukunci na rarrabuwa.
Stoner da abokan aikinsa14,23 su ne suka fara bayar da rahoton cewa canza hasken yankunan grid masu haɗuwa ya shafi daidaito da bayyananniya na rahotannin masu lura da ɗan adam da kuma daidaitawar alkibla a cikin ƙwayoyin MT masu kama da juna. Marubutan sun gano cewa lokacin da hasken yankunan da suka haɗu suka yi daidai da gaskiya, masu lura sun ba da rahoton ƙarin fahimta mai haske, yayin da ƙwayoyin MT ke nuna motsi na abubuwan raster. Akasin haka, lokacin da haske mai haɗuwa da haɗuwa mai haske ba su dace da jiki ba, mai lura yana fahimtar motsi mai daidaituwa, kuma ƙwayoyin MT suna nuna motsi na duniya na tsarin. Don haka, waɗannan nazarin sun nuna cewa canje-canje na zahiri a cikin abubuwan da ke haifar da gani waɗanda ke tasiri ga rahotannin rarrabuwa suma suna haifar da canje-canje da ake iya faɗi a cikin motsin MT. Aikin da aka yi kwanan nan a wannan fanni ya bincika waɗanne siginar MT ke bin diddigin bayyanar fahimta na abubuwan da ke haifar da rikitarwa18,24,64. Misali, an nuna wani ɓangare na ƙwayoyin MT suna nuna daidaitawar bimodal zuwa taswirar motsi bazuwar (RDK) tare da hanyoyi biyu waɗanda ba su da tazara fiye da RDK mai jagora ɗaya. Bandwidth na daidaita ƙwayoyin halitta 19, 25. Masu lura koyaushe suna ganin tsarin farko a matsayin motsi mai haske, duk da cewa yawancin ƙwayoyin MT suna nuna daidaitawar unimodal don mayar da martani ga waɗannan abubuwan ƙarfafawa, kuma matsakaicin matsakaici na duk ƙwayoyin MT yana ba da amsawar yawan jama'a mara tsari. Don haka, wani ɓangare na ƙwayoyin da ke nuna daidaitawar bimodal na iya samar da tushen jijiyoyi don wannan fahimta. Abin sha'awa, a cikin marmosets, wannan yawan jama'a ya dace da ƙwayoyin PDS lokacin da aka gwada su ta amfani da grid na al'ada da abubuwan ƙarfafa grid.
Sakamakonmu ya wuce mataki ɗaya fiye da abin da ke sama, waɗanda suke da mahimmanci wajen kafa rawar da MT ke takawa a cikin rarrabuwar fahimta. A zahiri, rarrabuwa wani abu ne na mutum ɗaya. Yawancin nunin gani na polystable suna nuna ikon tsarin gani don tsarawa da fassara abubuwan ƙarfafawa masu ɗorewa ta hanyoyi da yawa. A lokaci guda tattara martanin jijiyoyi da rahotannin fahimta a cikin bincikenmu ya ba mu damar bincika haɗin kai tsakanin ƙimar harbin MT da fassarar fahimta na motsawar akai-akai. Bayan mun nuna wannan alaƙar, mun yarda cewa ba a kafa alkiblar dalili ba, wato, ana buƙatar ƙarin gwaje-gwaje don tantance ko siginar rarrabuwar fahimta da muka lura, kamar yadda wasu ke jayayya [65, 66, 67], ta atomatik ne. Tsarin kuma yana wakiltar siginar saukowa da ke dawowa zuwa ga kwakwalwar jijiyoyi daga manyan yankuna 68, 69, 70 (Hoto na 8). Rahotannin babban rabo na ƙwayoyin zaɓaɓɓun tsari a cikin MSTd71, ɗaya daga cikin manyan manufofin cortical na MT, ya nuna cewa faɗaɗa waɗannan gwaje-gwajen don haɗawa da rikodin MT da MSTd a lokaci guda zai zama kyakkyawan mataki na farko don ƙara fahimtar hanyoyin jijiyoyi na fahimta. rarrabawa.
Tsarin matakai biyu na zaɓin sassa da yanayin da kuma tasirin da zai iya haifar da martani daga sama zuwa ƙasa kan ayyukan da suka shafi zaɓi a cikin fassarar na'ura. A nan, zaɓin alkiblar yanayi (PDS - "P") a cikin matakin MT an ƙirƙira shi ta hanyar (i) babban samfurin bayanan shigarwar alkiblar da suka dace da takamaiman saurin yanayi, da (ii) ƙarfin hana daidaitawa. Sashen zaɓin alkiblar (CDS) na matakin MT ("C") yana da kunkuntar kewayon samfuri a cikin alkiblar shigarwa kuma ba shi da yawan hana daidaitawa. Hanawa mara gyara yana ba da iko akan alkiblar duka biyun. Kibiyoyi masu launi suna nuna yanayin na'urar da aka fi so. Don fayyace, an nuna ƙaramin yanki na haɗin V1-MT da akwatin zaɓin sassa ɗaya da yanayin alkiblar. A cikin mahallin fassara sakamakonmu na gaba-gaba (FF), saitin shigarwa mai faɗi da hana daidaitawa mai ƙarfi (wanda aka haskaka da ja) a cikin ƙwayoyin PDS sun haifar da manyan bambance-bambance a cikin aiki don mayar da martani ga tsarin motsi da yawa. A cikin matsalar rarrabuwar mu, wannan rukunin yana jagorantar sarƙoƙin yanke shawara kuma yana karkatar da fahimta. Sabanin haka, a yanayin martani (FB), ana samar da yanke shawara mai fahimta a cikin da'irori na sama ta hanyar bayanan ji da kuma son zuciya, kuma babban tasirin FB na ƙasa akan ƙwayoyin PDS (layuka masu kauri) yana haifar da siginar zaɓi. b Wakiltar tsari na madadin samfuran na'urorin CDS da PDS. A nan ana samar da siginar PDS a cikin MT ba kawai ta hanyar shigar kai tsaye na V1 ba, har ma ta hanyar shigar kai tsaye na hanyar V1-V2-MT. Ana daidaita hanyoyin da ba kai tsaye na samfurin don ba da zaɓi ga iyakokin rubutu (wuraren da ke haɗuwa da grid). Tsarin CDS na Layer MT yana yin jimlar shigarwa kai tsaye da kai tsaye kuma yana aika fitarwa zuwa tsarin PDS. Ana tsara PDS ta hanyar hana gasa. Kuma, waɗannan haɗin ne kawai aka nuna waɗanda suka zama dole don zana tsarin asali na samfurin. A nan, wata hanyar FF daban fiye da wacce aka gabatar a cikin a na iya haifar da babban canji a cikin martanin layin sel zuwa PDS, wanda kuma ke haifar da son kai a cikin tsarin yanke shawara. A madadin haka, babban CP a cikin ƙwayoyin PDS har yanzu yana iya zama sakamakon son kai a cikin ƙarfi ko ingancin haɗewar FB zuwa ƙwayoyin PDS. Shaidun sun goyi bayan samfuran MT PDS guda biyu da uku da kuma fassarar CP FF da FB.
An yi amfani da manyan macaques guda biyu (macaca mulatta), namiji ɗaya da mace ɗaya (shekaru 7 da 5 bi da bi), waɗanda nauyinsu ya kama daga kilogiram 4.5 zuwa 9.0, a matsayin abubuwan da za a yi nazari a kansu. Kafin duk gwaje-gwajen tiyata marasa tsafta, an dasa dabbobi da ɗakin rikodi na musamman don electrodes masu tsaye kusa da yankin MT, wurin riƙe kai na bakin ƙarfe (Crist Instruments, Hagerstown, MD), da kuma matsayin ido tare da na'urar bincike ta scleral da aka auna. (Cooner Wire, San Diego, California). Duk yarjejeniyoyi sun bi ƙa'idodin Ma'aikatar Aikin Gona ta Amurka (USDA) da Ka'idojin Cibiyoyin Lafiya na Ƙasa (NIH) don Kula da Dan Adam da Amfani da Dabbobin Dakunan Gwaji kuma Kwamitin Kula da Dabbobi na Jami'ar Chicago (IAUKC) ya amince da su.
An gabatar da dukkan abubuwan da ke motsa gani a cikin zagaye mai kusurwa a kan bango mai baƙi ko launin toka. A lokacin rikodi, an daidaita matsayi da diamita na wannan ramin daidai da filin karɓa na gargajiya na ƙwayoyin jijiyoyi a ƙarshen lantarki. Mun yi amfani da manyan nau'ikan abubuwan da ke motsa gani guda biyu: abubuwan da ke motsa hankali da kuma abubuwan da ke motsa tunani.
Ƙarfafawar psychometric wani tsari ne na grating (20 cd/m2, bambanci 50%, zagayowar aiki 50%, digiri 5/sec) wanda aka ƙirƙira ta hanyar ɗora gratings guda biyu masu kusurwa huɗu waɗanda ke shawagi a cikin alkibla daidai da alkiblarsu (Hoto na 1b). An nuna a baya cewa masu lura da ɗan adam suna ɗaukar waɗannan tsarin grid a matsayin abubuwan ƙarfafawa masu ƙarfi, wani lokacin a matsayin tsari ɗaya da ke motsawa a cikin alkibla ɗaya (motsi mai daidaituwa) kuma wani lokacin a matsayin saman guda biyu daban-daban suna motsawa a cikin alkibla daban-daban (motsi mai haske). Abubuwan da ke cikin tsarin lattice, waɗanda aka daidaita daidai - kusurwar da ke tsakanin lattices ɗin daga 95° zuwa 130° (wanda aka zana daga saitin: 95°, 100°, 105°, 115°, 120°, 125°, 130°, a duk tsawon zaman ba a kiyaye ƙwayoyin kwakwalwan kusurwa na keɓewa a 115° ba, amma mun haɗa da bayanan psychophysical a nan) - kusan 90° ko 270° (daidaitaccen tsari). A kowane zaman, an yi amfani da kusurwa ɗaya kawai na layin interlattice; a lokacin kowane zaman, an zaɓi yanayin tsarin don kowane gwaji bazuwar daga hanyoyi biyu.
Domin mu fayyace fahimtar grid ɗin da kuma samar da tushen gwaji don ladan aiki, mun gabatar da zane-zanen maki bazuwar a cikin sandar haske mataki na 72 na kowane ɓangaren grid. Ana samun wannan ta hanyar ƙara ko ragewa (ta hanyar adadi mai ƙayyadadden) hasken wani ƙaramin pixel da aka zaɓa bazuwar (Hoto na 1c). Alkiblar motsin rubutu yana ba da sigina mai ƙarfi wanda ke canza fahimtar mai kallo zuwa motsi mai haɗin kai ko mai haske (Hoto na 1c). A ƙarƙashin yanayi mai haɗin kai, duk zane-zane, ba tare da la'akari da wane ɓangare na murfin layin rubutu ba, ana fassara su zuwa ga alkiblar tsarin (Hoto na 1c, mai haɗin kai). A cikin yanayin haske, yanayin yana motsawa daidai da alkiblar grating ɗin da yake rufewa (Hoto na 1c, mai haske) (Fim ɗin Ƙarin 1). Don sarrafa sarkakiyar aikin, a yawancin zaman, bambancin Michelson (Lmax-Lmin/Lmax+Lmin) don wannan alamar rubutu ya bambanta daga saitin (-80, -40, -20, -10, -5, 0, 5). , 10, 20, 40, 80). Ana bayyana bambanci a matsayin haske na ɗan lokaci na raster (don haka ƙimar bambanci ta 80% zai haifar da yanayin 36 ko 6 cd/m2). Don zaman 6 a cikin biri N da zaman 5 a cikin biri S, mun yi amfani da kewayon bambancin rubutu mai kunkuntar (-30, -20, -15, -10, -5, 0, 5, 10, 15, 20, 30), inda halayen psychophysical ke bin tsari iri ɗaya kamar bambancin cikakken zango, amma ba tare da cikawa ba.
Abubuwan da ke motsa sitiriyo sune grids na sinusoidal (bambanci 50%, zagaye 1/digiri, digiri 5/sec) suna motsawa a cikin ɗaya daga cikin kwatance 16 masu tazara daidai gwargwado, ko grids na sinusoidal suna motsawa a cikin waɗannan kwatance (wanda ya ƙunshi kusurwoyi biyu masu gaba da juna 135° gratings a saman juna). a cikin alkibla ɗaya ta tsarin.


Lokacin Saƙo: Nuwamba-13-2022