Umusaruro w'umuceri urimo kugabanuka bitewe n'imihindagurikire y'ikirere n'ihindagurika ry'ikirere muri Kolombiya.Abashinzwe kugenzura imikurire y'ibimerabyakoreshejwe nk'ingamba zo kugabanya ubushyuhe mu bihingwa bitandukanye. Kubwibyo, intego y'ubu bushakashatsi kwari ugusuzuma ingaruka z'umubiri (gukora umuceri mu nda, gukora umuceri mu nda, ingano ya chlorophyll yose, igipimo cya Fv/Fm cy'ubwoko bubiri bw'umuceri bucuruzwa bwakorewe ku bushyuhe buvanze (ubushyuhe bwinshi bwa manywa na nijoro), ubushyuhe bwo mu ihema n'ingano y'amazi) hamwe n'ibintu bihindura imiterere y'ibinyabuzima (malondialdehyde (MDA) na aside prolinic). Igerageza rya mbere n'irya kabiri byakozwe hakoreshejwe ibimera by'ubwoko bubiri bw'umuceri Federrose 67 ("F67") na Federrose 2000 ("F2000"), uko bikurikirana. Igerageza ryombi ryasesenguwe hamwe nk'urukurikirane rw'igerageza. Uburyo bwashyizweho ni ubu bukurikira: kugenzura burundu (AC) (ibimera by'umuceri byahinzwe ku bushyuhe bwiza (ubushyuhe bwa manywa na nijoro 30/25°C)), kugenzura ubushyuhe (SC) [ibimera by'umuceri byakorewe ku bushyuhe buvanze gusa (40/25°C). 30°C)], kandi ibihingwa by'umuceri byashyizwemo imbaraga nyinshi kandi bisukwaho imiti igenzura imikurire y'ibimera (stress+AUX, stress+BR, stress+CK cyangwa stress+GA) kabiri (iminsi 5 mbere na nyuma y'imikurire y'ubushyuhe). Gutera SA byongereye ingano ya chlorophyll y'ubwoko bwombi (uburemere bushya bw'ibihingwa by'umuceri "F67″ na "F2000″ bwari 3.25 na 3.65 mg/g, uko bikurikirana) ugereranije n'ibihingwa bya SC (uburemere bushya bw'ibihingwa "F67″ bwari 2.36 na 2.56 mg). g-1)" n'umuceri "F2000″, gukoresha CK ku bimera byabyo muri rusange byazamuye uburyo ibimera "F2000" bitwara mu nda (499.25 ugereranyije na 150.60 mmol m-2 s) ugereranije n'uburyo ubushyuhe butwara. Imikurire y'ubushyuhe, ubushyuhe bw'ikamba ry'ibimera bugabanukaho 2–3 °C, kandi ingano ya MDA mu bimera iragabanuka. Igipimo cy’ubuhanga mu guhangana n’imboga z’umuceri kigaragaza ko gukoresha CK (97.69%) na BR (60.73%) bishobora gufasha kugabanya ikibazo cy’ubushyuhe buvanze cyane cyane mu bihingwa by’umuceri bya F2000. Muri make, gutera BR cyangwa CK mu bihingwa bishobora gufatwa nk’ingamba z’ubuhinzi kugira ngo bigabanye ingaruka mbi z’ubushyuhe buvanze ku myitwarire y’ibihingwa by’umuceri.
Umuceri (Oryza sativa) uri mu muryango wa Poaceae kandi ni kimwe mu binyampeke bihingwa cyane ku isi hamwe n'ibigori n'ingano (Bajaj na Mohanty, 2005). Ubuso bw'umuceri buhingwaho ni hegitari 617,934, kandi umusaruro w'igihugu mu 2020 wari toni 2,937,840 hamwe n'umusaruro mpuzandengo wa toni 5.02 kuri hegitari (Federarroz (Federación Nacional de Arroceros), 2021).
Ubushyuhe bw'isi burimo kugira ingaruka ku bihingwa by'umuceri, bigatera ubwoko butandukanye bw'imihangayiko idashingiye ku binyabuzima nko ubushyuhe bwinshi n'ibihe by'amapfa. Imihindagurikire y'ikirere irimo gutuma ubushyuhe bw'isi bwiyongera; Biteganyijwe ko ubushyuhe buziyongeraho 1.0–3.7°C mu kinyejana cya 21, ibi bikaba bishobora kongera ubushyuhe n'ubukana bw'umuvuduko w'ubushyuhe. Ubwiyongere bw'ubushyuhe ku bidukikije bwagize ingaruka ku muceri, bituma umusaruro w'ibihingwa ugabanukaho 6–7%. Ku rundi ruhande, imihindagurikire y'ikirere inatera ibidukikije bibi ku bihingwa, nk'ibihe by'amapfa akomeye cyangwa ubushyuhe bwinshi mu turere dushyuha n'utwo mu turere dushyuha. Byongeye kandi, ibintu bitandukanye nka El Niño bishobora gutera ubushyuhe bwinshi no kwangiza imyaka mu turere tumwe na tumwe dushyuha. Muri Kolombiya, ubushyuhe mu turere duhinga umuceri buteganijwe kwiyongeraho 2–2.5°C mu 2050, bigabanyije umusaruro w'umuceri kandi bikagira ingaruka ku isoko no ku miyoboro y'ibicuruzwa.
Ibyinshi mu bihingwa by'umuceri bihingwa mu turere aho ubushyuhe buri hafi y'urugero rwiza rwo gukura kw'ibihingwa (Shah et al., 2011). Byaragaragaye ko ubushyuhe bwiza bwo ku manywa na nijoro kuriiterambere n'iterambere ry'umuceriMuri rusange ni 28°C na 22°C, uko bikurikirana (Kilasi et al., 2018; Calderón-Páez et al., 2021). Ubushyuhe buri hejuru y’iyi ngano bushobora gutera ibihe by’ubushyuhe buri hagati kugeza ku bunini mu bihe bikomeye byo gukura k’umuceri (guhinga, gutera indabo, kurabya, no kuzuza ibinyampeke), bityo bigira ingaruka mbi ku musaruro w’ibinyampeke. Uku kugabanuka k’umusaruro biterwa ahanini n’igihe kirekire cy’ubushyuhe, bigira ingaruka ku miterere y’ibimera. Bitewe n’imikoranire y’ibintu bitandukanye, nko kumara igihe cy’ubushyuhe n’ubushyuhe ntarengwa bugerwaho, ubushyuhe bushobora gutera ingaruka mbi ku mikorere y’ibimera no ku mikurire yabyo.
Ubushyuhe bugira ingaruka ku mikorere itandukanye y’ibinyabuzima n’ibinyabutabire mu bimera. Filime y’amababi ni imwe mu mikorere ishobora kwibasirwa cyane n’ubushyuhe mu bimera by’umuceri, kuko igipimo cya fotositeyine kigabanukaho 50% iyo ubushyuhe bwa buri munsi burenze 35°C. Uburyo ibimera by’umuceri bikora buratandukanye bitewe n’ubwoko bw’ubushyuhe. Urugero, igipimo cya fotositeyine n’uburyo umwijima ugenda utera mu nda birakumirwa iyo ibimera bihuye n’ubushyuhe bwinshi bwa manywa (33–40°C) cyangwa ubushyuhe bwinshi bwa manywa na nijoro (35–40°C ku manywa, 28–30°C). C bivuze nijoro) (Lü et al., 2013; Fahad et al., 2016; Chaturvedi et al., 2017). Ubushyuhe bwinshi bwa nijoro (30°C) butera fotositeyine kugabanya ubushyuhe ariko byongera guhumeka nijoro (Fahad et al., 2016; Alvarado-Sanabria et al., 2017). Uko igihe cy’ihungabana cyaba kiri kose, ubushyuhe bukabije bugira ingaruka ku ngano ya chlorophyll mu mababi, igipimo cy’urumuri ruhindagurika rwa chlorophyll ugereranyije n’urumuri runini rwa chlorophyll (Fv/Fm), hamwe n’imikorere ya Rubisco mu bimera by’umuceri (Cao et al. 2009; Yin et al. 2010). ) Sanchez Reynoso et al., 2014).
Impinduka mu binyabuzima ni ikindi kintu kijyanye no guhangana n’ubushyuhe bw’ibimera (Wahid et al., 2007). Ingano ya proline yakoreshejwe nk'ikimenyetso cy’ubushyuhe bw’ibimera (Ahmed na Hassan 2011). Proline igira uruhare runini mu mikorere y’ibimera kuko ikora nk'isoko ya karuboni cyangwa azote ndetse ikaba n'iy’ubushyuhe bw’ibimera mu gihe cy’ubushyuhe bwinshi (Sánchez-Reinoso et al., 2014). Ubushyuhe bwinshi nabwo bigira ingaruka ku buziranenge bw’ibimera binyuze mu gukwirakwiza lipide, bigatera ikorwa rya malondialdehyde (MDA) (Wahid et al., 2007). Kubwibyo, ingano ya MDA nayo yakoreshejwe mu gusobanukirwa imiterere y’uturemangingo tw’uturemangingo mu gihe cy’ubushyuhe (Cao et al., 2009; Chavez-Arias et al., 2018). Amaherezo, ubushyuhe buvanze [37/30°C (amanywa/ijoro)] bwongereye ijanisha ry’amazi ya electrolyte n’ingano ya malondialdehyde mu muceri (Liu et al., 2013).
Ikoreshwa ry’ibikoresho bigenzura imikurire y’ibimera (GRs) ryasuzumwe kugira ngo bigabanye ingaruka mbi z’ubushyuhe, kuko ibi bintu bigira uruhare runini mu gusubiza ibimera cyangwa mu buryo bwo kwirinda iyo mikurire (Peleg na Blumald, 2011; Yin et al. et al., 2011; Ahmed et al., 2015). Gukoresha umutungo kamere hanze byagize ingaruka nziza ku kwihanganira ubushyuhe mu bihingwa bitandukanye. Ubushakashatsi bwagaragaje ko hormone nka gibberellins (GA), cytokinins (CK), auxins (AUX) cyangwa brassinosteroids (BR) bituma habaho kwiyongera kw’ibintu bitandukanye bigize umubiri n’ibikomoka kuri biokemike (Peleg na Blumald, 2011; Yin et al. Ren, 2011; Mitler et al., 2012; Zhou et al., 2014). Muri Kolombiya, ikoreshwa ry’umutungo kamere hanze n’ingaruka zawo ku bihingwa by’umuceri ntirirasobanuka neza kandi ntiryigwa. Ariko, ubushakashatsi bwakozwe mbere bwagaragaje ko gutera BR ku mababi y'umuceri bishobora kongera ubushobozi bwo kwihanganira umuceri binyuze mu kunoza imiterere y'ihindagurika ry'imyuka, chlorophyll cyangwa ingano ya proline mu mababi y'imbuto z'umuceri (Quintero-Calderón et al., 2021).
Cytokinins zifasha ibimera guhangana n’ibibazo byo mu mubiri, harimo n’ubushyuhe (Ha et al., 2012). Byongeye kandi, byatangajwe ko gukoresha CK hanze bishobora kugabanya kwangirika k’ubushyuhe. Urugero, gukoresha zeatin hanze byongera umuvuduko wa photosynthesis, ingano ya chlorophyll a na b, ndetse no gutwara neza electron mu bimera bikurura bentgrass (Agrotis estolonifera) mu gihe cy’ubushyuhe (Xu na Huang, 2009; Jespersen na Huang, 2015). Gukoresha zeatin hanze bishobora no kunoza imikorere ya antioxydants, kongera imikorere ya poroteyine zitandukanye, kugabanya kwangirika kwa ogisijeni (ROS) no gukora malondialdehyde (MDA) mu ngingo z’ibimera (Chernyadyev, 2009; Yang et al., 2009). , 2016; Kumar et al., 2020).
Gukoresha aside gibberellic nabyo byagaragaje ko bitanga igisubizo cyiza ku bushyuhe. Ubushakashatsi bwagaragaje ko GA ikora mu buryo butandukanye kandi yongera ubushobozi bwo kwihanganira ubushyuhe bwinshi (Alonso-Ramirez et al. 2009; Khan et al. 2020). Abdel-Nabi et al. (2020) basanze gutera GA hanze y’ibimera (25 cyangwa 50 mg*L) bishobora kongera umuvuduko wa photosynthesis n’imikorere ya antioxydants mu bimera by’amacunga bifite ubushyuhe bwinshi ugereranije n’ibimera bigenzura. Byagaragaye kandi ko gukoresha HA hanze byongera ubushuhe, chlorophyll na carotenoid kandi bigabanya lipide peroxidation mu bimera bya tende (Phoenix dactylifera) biri mu bushyuhe bwinshi (Khan et al., 2020). Auxin kandi igira uruhare runini mu kugenzura uburyo bwo guhangana n’ubushyuhe bwinshi (Sun et al., 2012; Wang et al., 2016). Iyi regulator ikora nk'ikimenyetso cya biochemical mu mikorere itandukanye nko gukora proline cyangwa kwangirika kw'ibimera bitewe n'igabanuka ry'ibimera (Ali et al. 2007). Byongeye kandi, AUX yongera imikorere ya antioxidant, bigatuma MDA igabanuka mu bimera bitewe no kugabanuka kwa lipide peroxidation (Bielach et al., 2017). Sergeev et al. (2018) babonye ko mu bimera by'amashaza (Pisum sativum) biri mu bushyuhe, ingano ya proline - dimethylaminoethoxycarbonylmethyl) naphthylchloromethyl ether (TA-14) yiyongera. Muri ubwo bushakashatsi, babonye kandi urugero rwa MDA ruri hasi mu bimera byavuwe ugereranije n'ibimera bitavuwe na AUX.
Brassinosteroids ni ikindi cyiciro cy’imikurire ikoreshwa mu kugabanya ingaruka z’ubushyuhe. Ogweno et al. (2008) bavuze ko spray ya BR yo hanze yongera umuvuduko wa fotosynthetic, imikorere y’igifu n’umuvuduko ntarengwa wa Rubisco carboxylation y’ibimera by’inyanya (Solanum lycopersicum) biri mu bushyuhe mu gihe cy’iminsi 8. Gutera epibrassinosteroids ku bimera bya concombre (Cucumis sativus) biri mu bushyuhe mu gihe cy’iminsi 8. Byongeye kandi, gukoresha BR hanze bitinda kwangirika kwa chlorophyll kandi byongera imikorere myiza y’amazi n’umusaruro ntarengwa wa fotochimie ya PSII mu bimera biri mu bushyuhe mu gihe cy’ubushyuhe mu gihe cy’iminsi 8 (Holá et al., 2010; Toussagunpanit et al., 2015).
Kubera imihindagurikire y’ikirere n’imihindagurikire, ibihingwa byumuceri bihura nibihe byubushyuhe buri munsi (Lesk et al., 2016; Garcés, 2020; Federarroz (Federación Nacional de Arroceros), 2021). Muri fenotipi y’ibimera, ikoreshwa rya phytonutrients cyangwa biostimulants ryizwe nkingamba zo kugabanya ubukana bw’ubushyuhe mu turere duhinga umuceri (Alvarado-Sanabria et al., 2017; Calderón-Páez et al., 2021; Quintero-Calderón et al., 2021). Byongeye kandi, ikoreshwa ry’ibintu bihindura imiterere y’ibinyabuzima n’imiterere y’umubiri (ubushyuhe bw’amababi, uburyo bwo kuyobora imiterere y’ibimera mu nda, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, n’imiterere y’ibimera mu kirere) n’ingaruka z’uburyo bwo gushyira imiterere y’ibimera mu kirere mu buryo bw’ibimera bine bigenzura imikurire y’ibimera (AUX, CK, GA na BR). (Amabara y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, uburyo bwo gushyira imiterere y’ibimera mu kirere, n’uburyo bwo gushyira imiterere y’ibimera mu kirere) Impinduka mu bwoko bubiri bw'umuceri bucuruzwa zahuye n'ubushyuhe bwinshi (ubushyuhe bwinshi ku manywa na nijoro).
Muri ubu bushakashatsi, hakozwe igerageza ribiri ryigenga. Ubwoko bwa genotypes Federrose 67 (F67: ubwoko bwa genotype bwavutse mu bushyuhe bwinshi mu myaka icumi ishize) na Federrose 2000 (F2000: ubwoko bwa genotype bwavutse mu myaka icumi ishize y'ikinyejana cya 20 bugaragaza ubudahangarwa bwa virusi y'amababi yera) bwakoreshejwe ku nshuro ya mbere. Ubwoko bwa genotypes zombi zihingwa cyane n'abahinzi bo muri Kolombiya. Imbuto zatewe mu masahani ya litiro 10 (uburebure bwa cm 39.6, ubugari bwa cm 28.8, uburebure bwa cm 16.8) zirimo ubutaka bw'umucanga bufite 2% by'ibinyabuzima. Imbuto eshanu zabanje kumera zatewe muri buri sahani. Amasahani yashyizwe mu busitani bw'ishami ry'ubumenyi bw'ubuhinzi rya Kaminuza Nkuru ya Kolombiya, ishami rya Bogotá (43°50′56″ N, 74°04′051″ W), ku butumburuke bwa metero 2556 hejuru y'inyanja (asl). m.) kandi byakozwe kuva mu Ukwakira kugeza Ukuboza 2019. Igerageza rimwe (Federroz 67) n'igerageza rya kabiri (Federroz 2000) muri icyo gihembwe cya 2020.
Imiterere y'ibidukikije mu busitani bw'ubutaka muri buri gihembwe cy'ihinga ni iyi ikurikira: ubushyuhe bw'amanywa na nijoro 30/25°C, ubushyuhe 60~80%, igihe cy'ifoto karemano amasaha 12 (imirasire ikora ikoresheje fotositike 1500 µmol (fotoni) m-2 s-). 1 saa sita z'amanywa). Ibimera byafumbirijwe hakurikijwe ibikubiye muri buri kintu nyuma y'iminsi 20 imbuto zivutse (DAE), nk'uko Sánchez-Reinoso et al. (2019) abivuga: 670 mg azote kuri buri kimera, 110 mg fosifore kuri buri kimera, 350 mg potasiyumu kuri buri kimera, 68 mg kalisiyumu kuri buri kimera, 20 mg manyeziyumu kuri buri kimera, 20 mg sulfur kuri buri kimera, 17 mg silikoni kuri buri kimera. Ibimera birimo 10 mg boron kuri buri kimera, 17 mg umuringa kuri buri kimera, na 44 mg zinc kuri buri kimera. Ibimera by'umuceri byagumanye DAE kugeza kuri 47 muri buri gerageza ubwo byageraga ku cyiciro cya V5 muri iki gihe. Ubushakashatsi bwakozwe mbere bwagaragaje ko iki cyiciro cya phenological ari igihe gikwiye cyo gukora inyigo ku bushyuhe mu muceri (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2017).
Muri buri gerageza, hakozwe uburyo bubiri butandukanye bwo kugenzura imikurire y'amababi. Itsinda rya mbere rya spray ya phytohormone yakoreshejwe iminsi 5 mbere y'uko ubushyuhe butangira (42 DAE) kugira ngo bitegure ibimera ku mikurire y'ibidukikije. Hashyizweho ubundi spray ya phytohormone nyuma y'iminsi 5 ibimera byahuye n'imikurire y'ihungabana (52 DAE). Hakoreshejwe phytohormone enye kandi imiterere ya buri kintu gikora cyasutswe muri ubu bushakashatsi igaragara mu mbonerahamwe y'inyongera ya 1. Ingano y'ibipimo by'imikurire y'amababi byakoreshejwe ni iyi ikurikira: (i) Auxin (aside 1-naphthylacetic: NAA) ku gipimo cya 5 × 10−5 M (ii) 5 × 10–5 M gibberellin (aside gibberelllic: NAA); GA3); (iii) Cytokinin (trans-zeatin) 1 × 10-5 M (iv) Brassinosteroids [Spirostan-6-one, 3,5-dihydroxy-, (3b,5a,25R)] 5 × 10-5; M. Ibi bipimo byatoranijwe kuko bitera ibisubizo byiza kandi byongera ubushobozi bw'ibimera bwo kurwanya ubushyuhe (Zahir et al., 2001; Wen et al., 2010; El-Bassiony et al., 2012; Salehifar et al., 2017). Ibimera by'umuceri bidafite imiti igabanya ubukana bw'ibimera byavuwe n'amazi meza gusa. Ibimera byose by'umuceri byasutsweho umuti utera intoki. Shyira mililitiro 20 za H2O ku kimera kugira ngo utose ubuso bwo hejuru no hasi bw'amababi. Imiti yose itera imboga yakoreshejwe mu buhinzi (Agrotin, Bayer CropScience, Kolombiya) kuri 0.1% (v/v). Intera iri hagati y'inkono n'umuti utera imboga ni cm 30.
Uburyo bwo kuvura ubushyuhe bwakoreshejwe iminsi 5 nyuma yo gutera ubwa mbere (47 DAE) muri buri gerageza. Ibimera by'umuceri byimuwe mu cyumba cy'ubushyuhe bijyanwa mu cyumba cy'ubushyuhe cya litiro 294 (MLR-351H, Sanyo, IL, USA) kugira ngo hamenyekane ubushyuhe cyangwa hakomeze kubaho ibidukikije bimwe (47 DAE). Uburyo bwo kuvura ubushyuhe bwakoreshejwe hamwe bwakozwe hashyirwa icyumba ku bushyuhe bukurikira ku manywa/ijoro: ubushyuhe bwinshi ku manywa [40°C mu masaha 5 (kuva saa 11:00 kugeza 16:00)] n'igihe cyo nijoro [30°C mu masaha 5]. Iminsi 8 yikurikiranya (kuva saa 19:00 kugeza 24:00). Ubushyuhe bw'ubushyuhe n'igihe cyo kugaragara byatoranijwe hashingiwe ku bushakashatsi bwabanje (Sánchez-Reynoso et al. 2014; Alvarado-Sanabía et al. 2017). Ku rundi ruhande, itsinda ry'ibimera byimuriwe mu cyumba cy'imikurire byashyizwe mu busitani ku bushyuhe bumwe (30°C ku manywa/25°C nijoro) mu minsi 8 yikurikiranya.
Mu mpera z'igerageza, habonetse amatsinda akurikira yo kuvura: (i) imiterere y'ubushyuhe bw'ikura + ikoreshwa ry'amazi yaciwe [Absolute control (AC)], (ii) imiterere y'ubushyuhe + ikoreshwa ry'amazi yaciwe [Heat stress control (SC)], (iii) imiterere y'imiterere y'ubushyuhe + auxin application (AUX), (iv) imiterere y'ubushyuhe + gibberellin application (GA), (v) imiterere y'ubushyuhe + cytokinin application (CK), na (vi) imiterere y'ubushyuhe + brassinosteroid (BR) Umugereka. Aya matsinda yo kuvura yakoreshejwe ku bwoko bubiri bwa genotypes (F67 na F2000). Ubuvuzi bwose bwakozwe mu buryo bwa buri gihe hamwe n'ibipimo bitanu, buri kimwe kigizwe n'igihingwa kimwe. Buri gihingwa cyakoreshejwe mu gusoma ibihinduka byagenwe mu mpera y'igerageza. Igerageza ryamaze 55 DAE.
Uburyo bwo kohereza amazi mu nda (gs) bwapimwe hakoreshejwe porosometer igendanwa (SC-1, METER Group Inc., USA) iri hagati ya 0 na 1000 mmol m-2 s-1, ifite umwanya w'icyitegererezo wa mm 6.35. Ibipimo bifatwa hakoreshejwe icyuma gipima stomameter ku kibabi gikuze hamwe n'umushibu w'ingenzi w'ikimera wagutse cyane. Kuri buri buryo bwo kuvura, ingano ya gs yafashwe ku mababi atatu ya buri kimera hagati ya saa tanu na saa kumi n'ebyiri z'umugoroba kandi ikoreshwa mu mpuzandengo.
RWC yagenwe hakurikijwe uburyo bwasobanuwe na Ghoulam et al. (2002). Urupapuro rwagutse rwose rwakoreshejwe mu gupima g narwo rwakoreshejwe mu gupima RWC. Uburemere bushya (FW) bwagenwe ako kanya nyuma yo gusarurwa hakoreshejwe igipimo cya digitale. Amababi yashyizwe mu gikoresho cya pulasitiki cyuzuyemo amazi maze asigara mu mwijima ku bushyuhe bw'icyumba (22°C) mu gihe cy'amasaha 48. Hanyuma upime ku gipimo cya digitale hanyuma wandike uburemere bwagutse (TW). Amababi yabyimbye yumishijwe mu ifuru kuri 75°C mu gihe cy'amasaha 48 hanyuma uburemere bwayo bwumye (DW) bwanditswe.
Ingano ya chlorophyll yapimwe hakoreshejwe icyuma gipima chlorophyll (atLeafmeter, FT Green LLC, USA) kandi igaragazwa mu byiciro bya atLeaf (Dey et al., 2016). Ibipimo bya PSII bya quantum efficiency efficiency (Fv/Fm ratio) byafashwe hakoreshejwe fluorimeter ya chlorophyll ikomeza gushyuha (Handy PEA, Hansatech Instruments, UK). Amababi yahinduwe yijimye hakoreshejwe clamps z'amababi mu minota 20 mbere yo gupima Fv/Fm (Restrepo-Diaz na Garces-Varon, 2013). Nyuma y'uko amababi yijimye, hapimwe basal (F0) na max fluorescence (Fm). Muri aya makuru, fluorescence ihinduka (Fv = Fm – F0), igipimo cy’urumuri ruhinduka kugeza ku ruziga runini (Fv/Fm), umusaruro ntarengwa wa quantum wa photochemistry ya PSII (Fv/F0) n’igipimo cya Fm/F0 (Baker, 2008; Lee et al. ., 2017). Ingano ya chlorophyll na chlorophyll fluorescence yafashwe ku mababi amwe yakoreshejwe mu gupima gs.
Hafi mg 800 z'uburemere bw'amababi mashya byakusanyijwe nk'ibintu bihinduka mu buryo bwa biochemical. Ingero z'amababi zashyizwe mu buryo bumwe muri azote y'amazi hanyuma zibikwa kugira ngo zikorerwe isesengura rirambuye. Uburyo bwa spectrometric bwakoreshejwe mu gupima ingano ya chlorophyll a, b na carotenoid bushingiye ku buryo n'ibigereranyo byasobanuwe na Wellburn (1994). Ingero z'amababi (30 mg) zakusanyijwe kandi zishyirwa mu buryo bumwe muri mililitiro 3 za acetone ya 80%. Ingero zashyizwe mu buryo bwa centrifuge (model 420101, Becton Dickinson Primary Care Diagnostics, USA) ku muvuduko wa 5000 rpm mu gihe cy'iminota 10 kugira ngo haveho uduce duto. Supernatant yagabanijwe kugeza kuri mililitiro 6 hongewemo 80% ya acetone (Sims na Gamon, 2002). Ingano ya chlorophyll yapimwe kuri 663 (chlorophyll a) na 646 (chlorophyll b) nm, naho carotenoids kuri 470 nm hakoreshejwe spectrophotometer (Spectronic BioMate 3 UV-vis, Thermo, USA).
Uburyo bwa aside thiobarbituric (TBA) bwasobanuwe na Hodges et al. (1999) bwakoreshejwe mu gusuzuma peroxidation ya lipide mu membrane (MDA). Hafi garama 0.3 z'amababi nazo zahinduwe muri azote y'amazi. Ingero zashyizwe muri centrifuge kuri 5000 rpm kandi absorbance yapimwe kuri spectrophotometer kuri 440, 532 na 600 nm. Amaherezo, igipimo cya MDA cyabazwe hakoreshejwe coefficient yo kuzimira (157 M mL−1).
Ingano ya proline mu miti yose yapimwe hakoreshejwe uburyo bwasobanuwe na Bates et al. (1973). Ongeraho mililitiro 10 z'umuti w'amazi wa 3% wa aside sulfosalicylic ku gipimo cyabitswe hanyuma uyungurure unyuze ku rupapuro rwa Whatman filter (No. 2). Hanyuma mililitiro 2 z'iyi filtrate zasubiwemo mililitiro 2 za aside ninhydric na mililitiro 2 za aside glacial acetic. Uruvange rwashyizwe mu bwogero bw'amazi kuri 90°C mu gihe cy'isaha 1. Hagarika igisubizo ushyira ku rubura. Kangura umuyoboro cyane ukoresheje vortex shaker hanyuma ushongeshe umuti wavuyemo muri mililitiro 4 za toluene. Ingano y'umuti yapimwe kuri 520 nm ukoresheje spectrophotometer imwe yakoreshejwe mu gupima pigments za photosynthetic (Spectronic BioMate 3 UV-Vis, Thermo, Madison, WI, USA).
Uburyo bwasobanuwe na Gerhards et al. (2016) bwo kubara ubushyuhe bw'inyuma na CSI. Amafoto y'ubushyuhe yafashwe hifashishijwe kamera ya FLIR 2 (FLIR Systems Inc., Boston, MA, USA) ifite ubuziranenge bwa ± 2°C mu mpera y'igihe cy'ihungabana. Shyira ubuso bwera inyuma y'uruganda kugira ngo rufotore. Nanone, inganda ebyiri zafashwe nk'icyitegererezo. Ibimera byashyizwe ku buso bwera; bumwe bwari butwikiriwe n'igikoresho cy'ubuhinzi (Agrotin, Bayer CropScience, Bogotá, Kolombiya) kugira ngo bigane uburyo stomata zose zifunguka [wet mode (Twet)], naho ubundi bwari ikibabi kidakoreshejwe [Dry mode (Tdry)] (Castro -Duque et al., 2020). Intera iri hagati ya kamera n'inkono mu gihe cyo gufata amashusho yari metero 1.
Igipimo cy’ubwihangane bw’ibimera cyabazwe mu buryo butaziguye hakoreshejwe uburyo bwo kuyobora ibimera byavuwe (gs) ugereranije n’ibimera bigenzura (ibimera bidafite imiti igabanya ubukana kandi hakoreshejwe uburyo bwo kugenzura imikurire) kugira ngo hamenyekane ubushobozi bwo kwihanganira ubwoko bw’ibinyabuzima byavuwe bwasuzumwe muri ubu bushakashatsi. RTI yabonetse hakoreshejwe ikigereranyo cyakuwe muri Chávez-Arias et al. (2020).
Muri buri gerageza, ibintu byose bihinduka mu miterere y’umubiri byavuzwe haruguru byagenwe kandi byandikiwe kuri 55 DAE hakoreshejwe amababi yagutse yose yakusanyijwe mu gisenge cyo hejuru. Byongeye kandi, ibipimo byakorewe mu cyumba cy’ikuramo kugira ngo hirindwe guhindura imiterere y’ibidukikije ibimera bikuriramo.
Amakuru yavuye mu igerageza rya mbere n’irya kabiri yasesenguwe hamwe nk'urukurikirane rw'igerageza. Buri tsinda ry'igerageza ryari rigizwe n'ibimera 5, kandi buri kimera cyari kigize igice cy'igerageza. Isesengura ry'impinduka (ANOVA) ryakozwe (P ≤ 0.05). Iyo hagaragaye itandukaniro rikomeye, ikizamini cyo kugereranya cya Tukey cyakoreshejwe kuri P ≤ 0.05. Koresha imikorere ya arcsine kugira ngo uhindure agaciro k'ijanisha. Amakuru yasesenguwe hakoreshejwe porogaramu ya Statistix v 9.0 (Analytical Software, Tallahassee, FL, USA) kandi yashushanyijwe hakoreshejwe SigmaPlot (version 10.0; Systat Software, San Jose, CA, USA). Isesengura ry'ibice by'ingenzi ryakozwe hakoreshejwe porogaramu ya InfoStat 2016 (Analysis Software, Kaminuza Nkuru ya Cordoba, Arijantine) kugira ngo hamenyekane abagenzuzi beza b'imikurire y'ibimera bari kwigwaho.
Imbonerahamwe ya 1 igaragaza muri make ANOVA igaragaza igerageza, uburyo butandukanye bwo kuvura, n'imikoranire yabyo n'amabara y'ibibabi bya photosynthetic (chlorophyll a, b, total, na carotenoids), malondialdehyde (MDA) na proline, hamwe n'uburyo bwo kuyobora stomatal. Ingaruka za gs, ingano y'amazi. (RWC), ingano ya chlorophyll, imiterere ya chlorophyll alpha fluorescence, ubushyuhe bw'ikamba (PCT) (°C), ingano y'umuvuduko w'ibihingwa (CSI) n'ingano y'umusaruro w'ibimera by'umuceri kuri 55 DAE.
Imbonerahamwe ya 1. Incamake y'amakuru ya ANOVA ku bihinduka by'umubiri w'umuceri n'ibihinduka bya biokemikoro hagati y'igerageza (genotypes) n'uburyo bwo kuvura ubushyuhe.
Itandukaniro (P≤0.01) mu mikoranire y’amabara ya fotosite y’amababi, ingano ya chlorophyll (ibipimo bya Atleaf), n’ibipimo bya fluorescence bya alpha-chlorophyll hagati y’igerageza n’imiti bigaragazwa mu mbonerahamwe ya 2. Ubushyuhe bwinshi bwo ku manywa na nijoro bwongereye ingano ya chlorophyll na carotenoid yose. Imbuto z’umuceri zidafite imyuka ya phytohormones (2.36 mg g-1 kuri "F67″ na 2.56 mg g-1 kuri "F2000″)) ugereranije n’ibimera byahinzwe mu gihe cy’ubushyuhe bwiza (2.67 mg g -1)) byagaragaje ingano nto ya chlorophyll yose. Muri izo gerageza zombi, "F67" yari 2.80 mg g-1 na "F2000" yari 2.80 mg g-1. Byongeye kandi, ingemwe z'umuceri zavuwe hakoreshejwe uburyo bwa AUX na GA sprays mu gihe cy'ubushyuhe, zagaragaje ko chlorophyll yagabanutse mu bwoko bwa genotypes zombi (AUX = 1.96 mg g-1 na GA = 1.45 mg g-1 kuri "F67"; AUX = 1.96 mg g-1 na GA = 1.45 mg g-1 kuri "F67″; AUX = 2.24 mg) g-1 na GA = 1.43 mg g-1 (kuri "F2000″) mu gihe cy'ubushyuhe. Mu gihe cy'ubushyuhe, BR treatment y'ibimera byavuwe n'ibimera byazamutseho gato muri ubwo bwoko bwa genotypes zombi. Amaherezo, CK foliar spray yagaragaje agaciro ka fotosynthetic pigment mu buryo bwose bwo kuvura (AUX, GA, BR, SC na AC treatments) mu bwoko bwa genotypes F67 (3.24 mg g-1) na F2000 (3.65 mg g-1). Ingano ya chlorophyll (Atleaf unit) nayo yagabanutse bitewe n’ubushyuhe buvanze. Agaciro gakomeye cyane kagaragaye no mu bimera byatewe CC muri genotypes zombi (41.66 kuri "F67" na 49.30 kuri "F2000"). Igipimo cya Fv na Fv/Fm cyagaragaje itandukaniro rinini hagati y’uburyo bwo kuvura n’ubwoko bw’imbuto (Imbonerahamwe ya 2). Muri rusange, muri ibi bipimo, ubwoko bw’imbuto F67 ntibwashoboraga kwibasirwa n’ubushyuhe bwinshi kurusha ubwoko bw’imbuto F2000. Igipimo cya Fv na Fv/Fm cyagize ingaruka mbi cyane mu igerageza rya kabiri. Imbuto za 'F2000′ zari zifite ubushyuhe bwinshi zitateweho phytohormones zari zifite igipimo cya Fv kiri hasi cyane (2120.15) na Fv/Fm (0.59), ariko gutera imbuto za CK byafashije kugarura ibi gipimo (Fv: 2591, 89, igipimo cya Fv/Fm: 0.73). , kubona ibipimo bisa n'ibyanditswe ku bimera bya "F2000" byahinzwe mu gihe cy'ubushyuhe bwiza (Fv: 2955.35, igipimo cya Fv/Fm: 0.73: 0.72). Nta tandukaniro rigaragara mu miterere y'urumuri rw'ibanze (F0), urumuri rw'ikirenga (Fm), umusaruro w'ingufu za quantum za PSII (Fv/F0) na Fm/F0. Amaherezo, BR yagaragaje icyerekezo nk'icyo cyagaragaye kuri CK (Fv 2545.06, igipimo cya Fv/Fm 0.73).
Imbonerahamwe ya 2. Ingaruka z'ubushyuhe buvanze (40°/30°C ku manywa/ijoro) ku mabara y'amababi akoze muri photosynthesis [chlorophyll yose (Chl Total), chlorophyll a (Chl a), chlorophyll b (Chl b) na carotenoids Cx+c] ingaruka ], ingano ya chlorophyll (Atliff unit), ibipimo bya chlorophyll fluorescence (fluorescence y'ibanze (F0), fluorescence ntarengwa (Fm), fluorescence ihinduka (Fv), imikorere ntarengwa ya PSII (Fv/Fm), umusaruro ntarengwa wa quantum wa fotochemical wa PSII (Fv/F0 ) na Fm/F0 mu bimera by'ubwoko bubiri bw'umuceri [Federrose 67 (F67) na Federrose 2000 (F2000)] iminsi 55 nyuma yo kumera (DAE)).
Ingano y'amazi (RWC) y'ibimera by'umuceri byavuwe mu buryo butandukanye yagaragaje itandukaniro (P ≤ 0.05) mu mibanire hagati y'imiti ikoreshwa mu bushakashatsi n'imiti ikoreshwa mu bushakashatsi (Ishusho ya 1A). Iyo bivuwe na SA, agaciro ko hasi cyane kanditswe kuri genotype zombi (74.01% kuri F67 na 76.6% kuri F2000). Mu gihe cy'ubushyuhe bwinshi, RWC y'ibimera by'umuceri bya genotype zombi byavuwe hakoreshejwe phytohormones zitandukanye yariyongereye cyane. Muri rusange, ikoreshwa rya CK, GA, AUX, cyangwa BR ku bimera byongereye RWC ku gaciro kangana n'ak'ibimera byahinzwe mu bihe byiza mu gihe cy'igerageza. Gugenzura burundu no gutera imiti ku bimera byo mu bwoko bwa foil byagaragaje agaciro ka 83% kuri genotype zombi. Ku rundi ruhande, gs na zo zagaragaje itandukaniro rikomeye (P ≤ 0.01) mu mibanire ikoreshwa mu bushakashatsi no mu mibanire (Ishusho ya 1B). Uruganda rugenzura burundu (AC) narwo rwanditse agaciro kanini kuri buri bwoko bw’ibinyabuzima (440.65 mmol m-2s-1 kuri F67 na 511.02 mmol m-2s-1 kuri F2000). Ibihingwa by’umuceri byahuye n’ubushyuhe bwinshi byonyine byagaragaje agaciro ka gs kari hasi cyane kuri ubwoko bw’ibinyabuzima byombi (150.60 mmol m-2s-1 kuri F67 na 171.32 mmol m-2s-1 kuri F2000). Gutera ibimera hakoreshejwe uburyo bwose bwo kugenzura imikurire y’ibimera na byo byiyongereye. Ku bimera by’umuceri bya F2000 byasizwe CC, ingaruka zo gutera ibimera hifashishijwe phytohormones zari zigaragara cyane. Iri tsinda ry’ibimera nta tandukaniro ryagaragaje ugereranije n’ibimera bigenzura burundu (AC 511.02 na CC 499.25 mmol m-2s-1).
Ishusho ya 1. Ingaruka z'ubushyuhe buvanze (40°/30°C ku manywa/ijoro) ku gipimo cy'amazi (RWC) (A), imikorere y'inyama (gs) (B), umusaruro wa malondialdehyde (MDA) (C), n'ingano ya proline. (D) mu bimera by'ubwoko bubiri bw'umuceri (F67 na F2000) mu minsi 55 nyuma yo kugaragara (DAE). Ubuvuzi bwapimwe kuri buri bwoko bw'ibinyabuzima burimo: kugenzura burundu (AC), kugenzura ubushyuhe (SC), guhangayika mu bushyuhe + auxin (AUX), guhangayika mu bushyuhe + gibberellin (GA), guhangayika mu bushyuhe + mitogen y'uturemangingo (CK), no guhangayika mu bushyuhe + brassinosteroid. (BR). Buri nkingi igaragaza ikosa ry'ikigereranyo ± risanzwe ry'amakuru atanu (n = 5). Inkingi zikurikiwe n'inyuguti zitandukanye zigaragaza itandukaniro rikomeye mu mibare hakurikijwe ikizamini cya Tukey (P ≤ 0.05). Inyuguti zifite ikimenyetso kingana zigaragaza ko impuzandengo idasobanutse mu mibare (≤ 0.05).
Ibikubiye muri MDA (P ≤ 0.01) na proline (P ≤ 0.01) nabyo byagaragaje itandukaniro rikomeye mu mikoranire iri hagati y’igerageza n’imiti ya phytohormone (Ishusho ya 1C, D). Kwiyongera kwa lipid peroxidation byagaragaye mu miti ya SC muri genotypes zombi (Ishusho ya 1C), ariko ibimera byavuwe na spray igenzura ikura ry’amababi byagaragaje ko lipid peroxidation yagabanutse muri genotypes zombi; Muri rusange, ikoreshwa rya phytohormones (CA, AUC, BR cyangwa GA) rituma lipid peroxidation igabanuka (MDA). Nta tandukaniro ryabonetse hagati y’ibimera bya AC by’ubwoko bubiri bw’uturemangingo tw’umubiri n’ibimera biri munsi y’ubushyuhe kandi byateweho imisemburo ya phytohormones (byagaragaye ko agaciro ka FW mu bimera bya “F67” kari hagati ya 4.38–6.77 µmol g-1, naho mu bimera bya “F2000” bya “F2000” “byagaragaye ko agaciro ka 2.84 kugeza kuri 9.18 µmol g-1 (ibimera). Ku rundi ruhande, ikorwa rya proline mu bimera bya “F67″ ryari hasi ugereranyije n’ibimera bya “F2000″ biri munsi y’ubushyuhe buvanze, byatumye umusaruro wa proline wiyongera. mu bimera by’umuceri biri munsi y’ubushyuhe, mu bushakashatsi bwombi, byagaragaye ko gutanga iyi misemburo byongereye cyane aside amine mu bimera bya F2000 (AUX na BR byari 30.44 na 18.34 µmol g-1) uko bikurikirana (Ishusho ya 1G).
Ingaruka zo gutera imiti igabanya ubushyuhe ku bimera n’ibipimo by’ubushyuhe ku bimera bipima ubushyuhe n’ibipimo by’ubushyuhe (RTI) bigaragara ku ishusho ya 2A na B. Ku bimera byombi bipima ubushyuhe, ubushyuhe bw’ibimera bipima ubushyuhe bwari hafi ya 27°C, naho ubw’ibimera bipima ubushyuhe bwari hafi ya 28°C. WITH. Byagaragaye kandi ko gukoresha imiti ipima ubushyuhe bwa CK na BR byatumye ubushyuhe bw’ibimera bigabanukaho 2–3°C ugereranije n’ibimera bipima ubushyuhe (Ishusho ya 2A). RTI yagaragaje imyitwarire isa n’izindi mpinduka mu mikorere y’ibinyabuzima, igaragaza itandukaniro rikomeye (P ≤ 0.01) mu mibanire hagati y’igerageza n’imiti (Ishusho ya 2B). Ibimera bipima ubushyuhe byagaragaje ko ibimera bihangana bike mu bimera bipima ubushyuhe (34.18% na 33.52% ku bimera bipima umuceri bya "F67" na "F2000", uko bikurikirana). Gutanga hormones zo mu bimera bipima ubushyuhe ku bimera bipima ubushyuhe bwinshi bituma RTI irushaho kuba nziza mu bimera bihura n’ubushyuhe bwinshi. Iyi ngaruka yagaragaye cyane mu bimera bya “F2000” byasutsweho CC, aho RTI yari 97.69. Ku rundi ruhande, itandukaniro rikomeye ryagaragaye gusa mu gipimo cy’umusaruro (CSI) cy’ibimera by’umuceri munsi y’imiterere y’umusaruro wa foliar factor (P ≤ 0.01) (Ishusho ya 2B). Ibimera by’umuceri gusa byasutsweho umuvuduko w’ubushyuhe bwinshi ni byo byagaragaje agaciro kanini k’umuvuduko (0.816). Ubwo ibimera by’umuceri byasukwagaho imisemburo itandukanye ya phytohormones, igipimo cy’umuvuduko cyari hasi (agaciro kari kuva kuri 0.6 kugeza kuri 0.67). Amaherezo, igihingwa cy’umuceri cyahinzwe mu bihe byiza cyari gifite agaciro ka 0.138.
Ishusho ya 2. Ingaruka z'ubushyuhe buvanze (40°/30°C ku manywa/ijoro) ku bushyuhe bw'izuba (A), igipimo cyo kwihanganirana (RTI) (B), n'igipimo cyo guhangana n'ibihingwa (CSI) (C) by'amoko abiri y'ibimera. Ubwoko bw'ibimera by'umuceri (F67 na F2000) byakorewe ubushyuhe butandukanye. Ubuvuzi bwapimwe kuri buri bwoko bw'ibimera burimo: kugenzura burundu (AC), kugenzura guhangana n'ubushyuhe (SC), guhangana n'ubushyuhe + auxin (AUX), guhangana n'ubushyuhe + gibberellin (GA), guhangana n'ubushyuhe + mitogen y'uturemangingo (CK), n'ikibazo cyo guhangana n'ubushyuhe + brassinosteroid. (BR). Ubushyuhe buvanze burimo gushyira ibimera by'umuceri ku bushyuhe bwinshi bw'amanywa/ijoro (40°/30°C ku manywa/ijoro). Buri nkingi igaragaza ikosa ry'ikigereranyo ± ry'amakuru atanu (n = 5). Inkingi zikurikiwe n'inyuguti zitandukanye zigaragaza itandukaniro rikomeye ry'imibare hakurikijwe ikizamini cya Tukey (P ≤ 0.05). Inyuguti zifite ikimenyetso kingana zigaragaza ko impuzandengo idafatika mu mibare (≤ 0.05).
Isesengura ry’ibice by’ingenzi (PCA) ryagaragaje ko ibipimo byapimwe kuri 55 DAE byasobanuye 66.1% by’ibisubizo by’umubiri n’ibinyabutabire by’ibimera by’umuceri byafashwe n’ubushyuhe bivurwa hakoreshejwe spray igenzura ikura (Ishusho ya 3). Ibipimo bigaragaza ibipimo naho utudomo tugaragaza ibipimo bigaragaza ikura ry’ibimera (GRs). Ibipimo bya gs, chlorophyll, ubushobozi ntarengwa bwa PSII (Fv/Fm) n’ibipimo bya biochemical (TChl, MDA na proline) biri hafi y’aho bikomoka, bigaragaza isano iri hagati y’imyitwarire y’ibimera n’ibipimo byabyo. Itsinda rimwe (V) ryari rigizwe n’imbuto z’umuceri zahinzwe ku bushyuhe bwiza (AT) n’ibimera bya F2000 byavuwe na CK na BA. Muri icyo gihe, ibimera byinshi byavuwe na GR byagize itsinda ritandukanye (IV), naho kuvurwa na GA muri F2000 byagize itsinda ritandukanye (II). Mu buryo bunyuranye, ingemwe z'umuceri zishyushye cyane (itsinda rya I n'irya III) zidafite imisemburo ya phytohormones (ubwoko bwombi bw'uturemangingo twa genotypes bwari SC) zashyizwe mu gace kari imbere y'itsinda rya V, bigaragaza ingaruka z'ubushyuhe ku miterere y'ibimera.
Ishusho ya 3. Isesengura ry'ibisekuruza ku ngaruka z'ubushyuhe buvanze (40°/30°C ku manywa/ijoro) ku bimera by'ubwoko bubiri bw'umuceri (F67 na F2000) mu minsi 55 nyuma yo kumera (DAE). Incamake: AC F67, kugenzura burundu F67; SC F67, kugenzura ubushyuhe F67; AUX F67, kugenzura ubushyuhe + auxin F67; GA F67, kugenzura ubushyuhe + gibberellin F67; CK F67, kugenzura ubushyuhe + igabanywa ry'uturemangingo BR F67, kugenzura ubushyuhe + brassinosteroid. F67; AC F2000, kugenzura burundu F2000; SC F2000, kugenzura ubushyuhe F2000; AUX F2000, kugenzura ubushyuhe + auxin F2000; GA F2000, kugenzura ubushyuhe + gibberellin F2000; CK F2000, umuvuduko w'ubushyuhe + cytokinin, BR F2000, umuvuduko w'ubushyuhe + steroid y'umuringa; F2000.
Ibintu bitandukanye nka chlorophyll, stomatal conductance, Fv/Fm ratio, CSI, MDA, RTI na proline ingano bishobora gufasha gusobanukirwa uburyo imiterere y’uturemangingo tw’umuceri ihinduka no gusuzuma ingaruka z’ingamba z’ubuhinzi mu gihe cy’ubushyuhe (Sarsu et al., 2018; Quintero-Calderon et al., 2021). Intego y’ubu bushakashatsi kwari ukureba ingaruka z’ikoreshwa ry’ibikoresho bine bigenzura imikurire ku bipimo by’umubiri n’ibinyabutabire by’imbuto z’umuceri mu gihe cy’ubushyuhe bugoye. Gupima imbuto ni uburyo bworoshye kandi bwihuse bwo gusuzuma icyarimwe ibimera by’umuceri bitewe n’ingano cyangwa imiterere y’ibikorwa remezo bihari (Sarsu et al. 2018). Ibisubizo by’ubu bushakashatsi byagaragaje ko ubushyuhe buvanze butera ibisubizo bitandukanye by’umubiri n’ibinyabutabire mu turemangingo tubiri tw’umuceri, bigaragaza inzira yo guhangana n’ingaruka. Ibi bisubizo kandi bigaragaza ko imiti igabanya ubukana bw'ibimera (cyane cyane cytokinins na brassinosteroids) ifasha umuceri kumenyera ubushyuhe bukabije kuko ubukire bugira ingaruka cyane cyane ku bimera, RWC, Fv/Fm ratio, ibara rya photosynthesis n'ingano ya proline.
Gukoresha uburyo bwo kugenzura imikurire bifasha kunoza imiterere y'amazi y'ibimera by'umuceri biri mu bushyuhe, bishobora kuba bifitanye isano n'umuvuduko mwinshi no kugabanuka k'ubushyuhe bw'ibimera. Ubu bushakashatsi bwerekanye ko mu bimera bya "F2000" (bishobora kwangirika mu buryo bw'uturemangingo), ibimera by'umuceri byavuwe ahanini na CK cyangwa BR byari bifite agaciro ka gs kandi bifite agaciro ka PCT kari hasi ugereranyije n'ibimera byavuwe na SC. Ubushakashatsi bwabanje bwagaragaje kandi ko gs na PCT ari ibimenyetso nyabyo by'umubiri bishobora kugena uburyo ibimera by'umuceri bihangana n'ingaruka z'ingamba z'ubuhinzi ku bushyuhe (Restrepo-Diaz na Garces-Varon, 2013; Sarsu et al., 2018; Quintero). -Carr DeLong et al., 2021). CK y'ikibabi cyangwa BR yongera g iri mu bushyuhe kuko iyi misemburo y'ibimera ishobora gutuma stomatal ifunguka binyuze mu gukorana n'izindi molekile z'ibimenyetso nka ABA (promoter of stomatal closure under abiotic stress) (Macková et al., 2013; Zhou et al., 2013). 2013). , 2014). Gufungura igifu bituma amababi akonja kandi bigafasha kugabanya ubushyuhe bw'igifu (Sonjaroon et al., 2018; Quintero-Calderón et al., 2021). Kubera izi mpamvu, ubushyuhe bw'igifu ku bimera by'umuceri byasizwe CK cyangwa BR bushobora kuba buke iyo ubushyuhe buvanze.
Ubushyuhe bwinshi bushobora kugabanya ingano y’amabara y’ibibabi mu bushyuhe kandi nta miti ikoreshwa mu kugenzura imikurire y’ibimera, ingano y’ibibabi mu bushyuhe yagabanukaga mu bwoko bwa genotypes zombi (Imbonerahamwe ya 2). Feng et al. (2013) nabo bavuze ko ingano ya chlorophyll yagabanutse cyane mu mababi y’ubwoko bubiri bw’ingano bwagaragayeho ubushyuhe bwinshi. Guhura n’ubushyuhe bwinshi akenshi bituma ingano ya chlorophyll igabanuka, bishobora guterwa no kugabanuka kwa chlorophyll mu bushyuhe, kwangirika kw’amabara, cyangwa ingaruka zabyo zose hamwe mu bushyuhe bwinshi (Fahad et al., 2017). Ariko, ibimera by’umuceri byavuwe ahanini na CK na BA byongereye ingano y’ibibabi mu bushyuhe bwinshi. Ibisubizo nk’ibyo byatangajwe na Jespersen na Huang (2015) na Suchsagunpanit et al. (2015), wabonye ubwiyongere bw'ingano ya chlorophyll mu mababi nyuma yo gukoresha imisemburo ya zeatin na epibrassinosteroid mu bushyuhe bukabije bwa bentgrass n'umuceri, uko bikurikirana. Impamvu ifatika ituma CK na BR bitera kwiyongera kw'ingano ya chlorophyll mu mababi mu gihe cy'ubushyuhe buvanze ni uko CK ishobora kongera gutangira uburyo bwo gukomeza gutera imbere kw'ibitera imbaraga mu kugaragaza (nk'ibitera imbaraga mu gusaza (SAG12) cyangwa HSP18) no kugabanya ibura rya chlorophyll mu mababi. , bigatinda gusaza kw'amababi no kongera ubushobozi bw'ibimera bwo kurwanya ubushyuhe (Liu et al., 2020). BR ishobora kurinda chlorophyll mu mababi no kongera umubare wa chlorophyll mu mababi binyuze mu gukora cyangwa gutera imbaraga mu gukora enzymes zigize uruhare mu gukora chlorophyll mu gihe cy'ubushyuhe (Sharma et al., 2017; Siddiqui et al., 2018). Amaherezo, imisemburo ibiri ya phytohormones (CK na BR) nayo itera imbere igaragara rya poroteyine zishyuha cyane kandi ikanoza inzira zitandukanye zo guhindura imikorere y’umubiri, nko kongera ubwiyongere bwa chlorophyll biosynthesis (Sharma et al., 2017; Liu et al., 2020).
Ibipimo bya Chlorophyll a fluorescence bitanga uburyo bwihuse kandi butangiza bushobora gusuzuma ubushobozi bw'ibimera bwo kwihanganira cyangwa guhangana n'imimerere y'imihangayiko ya abiotic (Chaerle et al. 2007; Kalaji et al. 2017). Ibipimo nka Fv/Fm ratio byakoreshejwe nk'ibipimo by'uko ibimera bihangana n'imimerere y'imihangayiko (Alvarado-Sanabria et al. 2017; Chavez-Arias et al. 2020). Muri ubu bushakashatsi, ibimera bya SC byagaragaje agaciro kari hasi cyane k'iki gipimo, cyane cyane ibimera by'umuceri bya "F2000". Yin et al. (2010) nabo basanze igipimo cya Fv/Fm cy'amababi y'umuceri yo guhinga menshi cyaragabanutse cyane ku bushyuhe buri hejuru ya 35°C. Nk'uko Feng et al. (2013 abivuga, igipimo gito cya Fv/Fm munsi y'imihangayiko y'ubushyuhe kigaragaza ko igipimo cyo gufata no guhindura ingufu z'ubushyuhe n'ikigo cy'imyitozo cya PSII cyagabanutse, bigaragaza ko ikigo cy'imyitozo cya PSII gisenyuka bitewe n'imihangayiko y'ubushyuhe. Iyi ngingo idufasha kwemeza ko ibibazo biri mu bikoresho bitanga fotosynthesis bigaragara cyane mu bwoko bw’ibimera bishobora kwangirika (Fedearroz 2000) kurusha mu bwoko bw’ibimera birwanya imihindagurikire y’ikirere (Fedearroz 67).
Gukoresha CK cyangwa BR muri rusange byongereye imikorere ya PSII mu bihe bigoye byo gushyuha. Ibisubizo nk'ibyo byabonetse na Suchsagunpanit et al. (2015), babonye ko gukoresha BR byongera imikorere ya PSII mu gihe cy'ubushyuhe mu muceri. Kumar et al. (2020) nabo basanze ibimera by'amashaza byavuwe na CK (6-benzyladine) kandi bigashyirwa mu bushyuhe byongera igipimo cya Fv/Fm, basoza bavuga ko gukoresha CK mu mashaza binyuze mu gukora uruziga rwa zeaxanthin byateraga imbaraga za PSII. Byongeye kandi, spray y'amababi ya BR yakunze fotosite ya PSII mu gihe cy'ubushyuhe buhuriweho, bigaragaza ko gukoresha iyi phytohormone byatumye ingufu zo gushyuha za antene za PSII zigabanuka kandi bikanatera ubwiyongere bwa poroteyine nto zo gushyuha mu ma chloroplast (Ogweno et al. 2008; Kothari na Lachowitz)., 2021).
Ibikubiye muri MDA na proline bikunze kwiyongera iyo ibimera biri munsi y’ubushyuhe bukabije ugereranije n’ibimera byahinzwe mu bihe byiza (Alvarado-Sanabria et al. 2017). Ubushakashatsi bwabanje bwagaragaje kandi ko urugero rwa MDA na proline ari ibipimo bya biochemical bishobora gukoreshwa mu gusobanukirwa inzira yo guhangana n’imihindagurikire y’ibihe cyangwa ingaruka z’ubuhinzi mu muceri munsi y’ubushyuhe bwinshi bw’amanywa cyangwa nijoro (Alvarado-Sanabria et al., 2017; Quintero-Calderón et al., 2021). Ubu bushakashatsi bwagaragaje kandi ko ibikubiye muri MDA na proline byakundaga kuba byinshi mu bimera by’umuceri bihura n’ubushyuhe bwinshi nijoro cyangwa ku manywa, uko bikurikirana. Ariko, gutera CK na BR ku mashanyarazi byagize uruhare mu kugabanuka kwa MDA no kwiyongera kwa proline, cyane cyane mu bwoko bwa genotype bwihanganira (Federroz 67). CK spray ishobora gutuma cytokinin oxidase/dehydrogenase irushaho kwiyongera, bityo yongera ingano y’ibinyabutabire nka betaine na proline (Liu et al., 2020). BR itera imbere mu kongeramo osmoprotectors nka betaine, isukari, na aside amine (harimo na proline yigenga), ikomeza kuringaniza osmotic y'uturemangingo mu gihe hari ibidukikije bibi byinshi (Kothari na Lachowiec, 2021).
Igipimo cy’imihangayiko y’ibihingwa (CSI) n’igipimo cy’imihangayiko y’ibihingwa (RTI) bikoreshwa mu kumenya niba uburyo bwo kuvura burimo gusuzumwa bufasha kugabanya imihangayiko itandukanye (abiotic na biotic) kandi bukagira ingaruka nziza ku miterere y’ibimera (Castro-Duque et al., 2020; Chavez-Arias et al., 2020). Igipimo cya CSI gishobora kuva kuri 0 kugeza kuri 1, kigaragaza imiterere idasaba imihangayiko n’imihangayiko, uko bikurikirana (Lee et al., 2010). Igipimo cya CSI cy’ibimera bifite imihangayiko y’ubushyuhe (SC) cyari kuva kuri 0.8 kugeza kuri 0.9 (Ishusho ya 2B), bigaragaza ko ibimera by’umuceri byagizweho ingaruka mbi n’imihangayiko ihuriweho. Ariko, gutera BC (0.6) cyangwa CK (0.6) ku bimera by’umuceri mu mashanyarazi byatumye iki kimenyetso kigabanuka bitewe n’imihangayiko y’ibimera ugereranije n’ibimera by’umuceri bya SC. Mu bimera bya F2000, RTI yagaragaje ubwiyongere bwinshi ubwo yakoreshaga CA (97.69%) na BC (60.73%) ugereranije na SA (33.52%), bigaragaza ko izi nganda zigenzura imikurire y'ibimera nazo zigira uruhare mu kunoza uburyo umuceri wihanganira imiterere yawo. Gushyuha cyane. Ibi bipimo byatanzwe kugira ngo bicunge imiterere y'imihangayiko mu moko atandukanye. Ubushakashatsi bwakozwe na Lee et al. (2010) bwagaragaje ko CSI y'ubwoko bubiri bw'ipamba buri munsi y'amazi make yari hafi 0.85, mu gihe agaciro ka CSI k'ubwoko buvomererwa neza kari hagati ya 0.4 na 0.6, byemeza ko iki gipimo ari ikimenyetso cy'uko ubwoko bushobora guhangana n'amazi mu bihe by'imihangayiko. Byongeye kandi, Chavez-Arias et al. (2020) basuzumye akamaro k'ibintu bitera imihangayiko nk'ingamba zirambuye zo gucunga imihangayiko mu bimera bya C. elegans kandi basanga ibimera bisutsweho ibi binyabutabire byagaragaje RTI iri hejuru (65%). Hashingiwe ku byavuzwe haruguru, CK na BR bishobora gufatwa nk'ingamba z'ubuhinzi zigamije kongera ubushobozi bw'umuceri bwo kwihanganira ubushyuhe bwinshi, kuko izi ngaruka zigenzura imikurire y'ibimera zitera ingaruka nziza ku mikorere y'ibinyabutabire n'imikorere y'umubiri.
Mu myaka mike ishize, ubushakashatsi ku muceri muri Kolombiya bwibanze ku gusuzuma ubwoko bw’uturemangingo twihanganira ubushyuhe bwinshi bwa manywa cyangwa nijoro hakoreshejwe imiterere y’umubiri cyangwa imiterere ya biokemike (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2021). Ariko, mu myaka mike ishize, isesengura ry’ikoranabuhanga rifatika, rihendutse kandi ryunguka ryarushijeho kuba ingenzi kugira ngo haboneke uburyo bwo gucunga ibihingwa mu buryo buhuriweho kugira ngo hongerwe ingaruka z’ibihe bigoye by’ubushyuhe mu gihugu (Calderón-Páez et al., 2021; Quintero-Calderon et al., 2021). Bityo, uburyo ibimera by’umuceri byitabira ubushyuhe bukabije (40°C ku manywa/30°C nijoro) byagaragaye muri ubu bushakashatsi bigaragaza ko gutera CK cyangwa BR imbuto ku mashanyarazi bishobora kuba uburyo bwiza bwo gucunga ibihingwa kugira ngo bigabanye ingaruka mbi. Ingaruka z’ibihe bikomeye by’ubushyuhe buciriritse. Ubu buryo bwo kuvura bwazamuye ubushobozi bwo kwihanganira ubwoko bw’uturemangingo twombi tw’umuceri (CSI nke na RTI nyinshi), bigaragaza icyerekezo rusange mu buryo ibimera byitabira mu buryo bw’umubiri n’imiterere ya biokemike mu gihe cy’ubushyuhe buvanze. Igisubizo cy’ingenzi cy’ibimera by’umuceri cyari ukugabanuka kw’ingano ya GC, chlorophyll yose, chlorophyll α na β na carotenoids. Byongeye kandi, ibimera bigira ingaruka ku kwangirika kwa PSII (ibipimo bya chlorophyll fluorescence byagabanutse nka Fv/Fm ratio) no kwiyongera kwa lipide peroxidation. Ku rundi ruhande, igihe umuceri wavurwaga na CK na BR, izi ngaruka mbi zaragabanutse kandi ingano ya proline yiyongera (Ishusho ya 4).
Ishusho ya 4. Icyitegererezo cy'ingaruka z'ubushyuhe buvanze hamwe n'umuti ugabanya ubushyuhe ku bimera by'umuceri. Umwambi utukura n'ubururu ugaragaza ingaruka mbi cyangwa nziza z'imikoranire hagati y'ubushyuhe n'ikoreshwa rya BR (brassinosteroid) na CK (cytokinin) ku bimera by'umuceri n'ibimera by'umuceri, uko bikurikirana. gs: imikorere y'igifu; Chl yose: ingano ya chlorophyll yose; Chl α: ingano ya chlorophyll β; Cx+c: ingano ya carotenoid;
Muri make, ibisubizo by’imiterere y’umubiri n’ibijyanye n’ubuhanga muri ubu bushakashatsi bigaragaza ko ibimera by’umuceri bya Fedearroz 2000 bishobora kwibasirwa cyane n’igihe cy’ubushyuhe bukabije kurusha ibimera by’umuceri bya Fedearroz 67. Abashinzwe kugenzura imikurire bose bagenzuwe muri ubu bushakashatsi (auxins, gibberellins, cytokinins, cyangwa brassinosteroids) bagaragaje ku rwego runaka rwo kugabanya ubushyuhe. Ariko, cytokinin na brassinosteroids byatumye ibimera bihinduka neza kuko abashinzwe kugenzura imikurire y’ibimera bombi bongereye chlorophyll, alpha-chlorophyll fluorescence parameters, gs na RWC ugereranije n’ibimera by’umuceri bitakoreshejwe, ndetse binagabanya ingano ya MDA n’ubushyuhe bw’inyuma. Muri make, twanzura ko ikoreshwa ry’abashinzwe kugenzura imikurire y’ibimera (cytokinins na brassinosteroids) ari igikoresho cy’ingirakamaro mu gucunga imikurire y’ibihingwa by’umuceri iterwa n’ubushyuhe bwinshi mu bihe by’ubushyuhe bwinshi.
Ibikoresho by’umwimerere byatanzwe muri ubu bushakashatsi biri muri iyo nkuru, kandi ibindi bibazo bishobora koherezwa ku mwanditsi ubijyanye.
Igihe cyo kohereza: Kanama-08-2024