Imveliso yerayisi iyancipha ngenxa yokutshintsha kwemozulu kunye nokuguquguquka eColombia.Izilawuli zokukhula kwezityalozisetyenziswe njengeqhinga lokunciphisa uxinzelelo lobushushu kwizityalo ezahlukeneyo. Ngoko ke, injongo yolu phononongo yayikukuvavanya imiphumo ye-physiological (i-stomatal conductance, i-stomatal conductance, umxholo we-chlorophyll iyonke, i-Fv / Fm ratio ye-rayisi ezimbini ze-genotypes zorhwebo ezixhomekeke kuxinzelelo lobushushu obudibeneyo (ubushushu obuphezulu bemini nobusuku), ukushisa kwe-canopy kunye nomxholo wamanzi ohambelanayo) kunye ne-biochemical variables (malondialdehyde (i-MDA) kunye ne-prolinic acid content). Uvavanyo lokuqala nolwesibini lwenziwa kusetyenziswa izityalo ezimbini zerayisi genotypes Federrose 67 ("F67") kunye Federrose 2000 ("F2000"), ngokulandelanayo. Omabini amalinge ahlalutywa kunye njengoluhlu lwemifuniselo. Unyango olusekiweyo lube ngolu hlobo lulandelayo: ulawulo olupheleleyo (AC) (izityalo zerayisi ezikhule kumaqondo obushushu afanelekileyo (ubushushu bemini/ubusuku 30/25°C)), ulawulo loxinzelelo lobushushu (SC) [izityalo zerayisi ziphantsi koxinzelelo lobushushu oludibeneyo kuphela (40/25°C). 30 ° C)], kunye nezityalo zerayisi zigxininiswe kwaye zifafazwe ngabalawuli bokukhula kwezityalo (i-stress + AUX, i-stress + BR, i-stress + CK okanye i-stress + GA) kabini (iintsuku ezi-5 ngaphambi kunye neentsuku ezi-5 emva koxinzelelo lobushushu). Ukutshiza nge-SA kwandisa umxholo we-chlorophyll uwonke wazo zombini iindidi (ubunzima obutsha bezityalo zerayisi "F67" kunye ne "F2000" yayiyi-3.25 kunye ne-3.65 mg / g, ngokulandelanayo) xa kuthelekiswa nezityalo ze-SC (ubunzima obutsha be-"F67" yezityalo yayiyi-2.36 kunye ne-2.56 mg ye-rayisi "i-arF″" kunye ne-g-1 yesicelo″ ″ ye-0CK ″ ″ ″ ″ ″ ″ ″ ″ ″ ″ kunye ne-2.56 mg ngokubanzi iphucule i-stomatal conductance yerayisi "F2000" izityalo (499.25 vs. 150.60 mmol m-2 s) xa kuthelekiswa nolawulo loxinzelelo lobushushu. uxinzelelo lobushushu, ukushisa kwesithsaba sesityalo kuncipha ngo-2-3 °C, kwaye umxholo we-MDA kwizityalo uyancipha. Isalathisi sokunyamezelana sibonisa ukuba ukusetyenziswa kwe-foliar ye-CK (97.69%) kunye ne-BR (60.73%) kunokunceda ukunciphisa ingxaki yobushushu obudibeneyo. uxinzelelo ikakhulu F2000 izityalo irayisi. Ukuqukumbela, ukutshiza nge-foliar ye-BR okanye i-CK inokuthathwa njengeqhinga le-agronomic yokunceda ukunciphisa imiphumo emibi yeemeko ezidibeneyo zoxinzelelo lobushushu kwindlela yokuziphatha ye-physiological yezityalo zerayisi.
Irayisi (i-Oryza sativa) yeyosapho lakwaPoaceae kwaye yenye yezona cereal zilinywayo kwihlabathi kunye nombona kunye nengqolowa (Bajaj kunye noMohanty, 2005). Indawo ephantsi kokulinywa kwerayisi ziihektare ezingama-617,934, kwaye imveliso yelizwe ngo-2020 yayiziitoni ezingama-2,937,840 kunye nesivuno esiphakathi kweetoni ezingama-5.02/ha (Federarroz (Federación Nacional de Arroceros), ngo-2021).
Ukufudumala kwehlabathi kuchaphazela izityalo zerayisi, okukhokelela kwiindidi ezahlukeneyo zoxinzelelo lwe-abiotic njengamaqondo obushushu aphezulu kunye namaxesha embalela. Ukutshintsha kwemozulu kubangela ukuba amaqondo obushushu anyuke; Amaqondo obushushu aqikelelwa ukuba anyuke nge-1.0–3.7°C ngenkulungwane yama-21, nto leyo enokwandisa ukuphindaphindeka kunye nokuqina koxinzelelo lobushushu. Ukunyuka kwamaqondo obushushu okusingqongileyo kuchaphazele irayisi, nto leyo ebangela ukuba izivuno zezityalo zehle nge-6–7%. Kwelinye icala, utshintsho lwemozulu lukwakhokelela kwiimeko ezingathandekiyo zokusingqongileyo kwizityalo, njengamaxesha embalela eqatha okanye amaqondo obushushu aphezulu kwimimandla etshisayo neshushu . Ukongeza, iziganeko eziguquguqukayo ezifana ne-El Niño zinokukhokelela kuxinzelelo lobushushu kwaye zandise umonakalo wezityalo kwezinye iingingqi ezitshisayo. E-Colombia, amaqondo okushisa kwiindawo ezivelisa irayisi kulindeleke ukuba anyuke nge-2-2.5 ° C ngo-2050, ukunciphisa imveliso yerayisi kunye nokuchaphazela ukuhamba kwemveliso kwiimarike kunye neentambo zokubonelela.
Uninzi lwezityalo zerayisi zikhuliswa kwiindawo apho amaqondo obushushu akufutshane nowona luhlu lulungileyo lokukhula kwesityalo (Shah et al., 2011). Kuye kwaxelwa ukuba i-avareji ilungileyo amaqondo obushushu imini nobusuku forukukhula kwerayisi kunye nophuhlisongokuqhelekileyo i-28 ° C kunye ne-22 ° C, ngokulandelanayo (Kilasi et al., 2018; Calderón-Páez et al., 2021). Amaqondo obushushu angaphezulu kwale miqobo anokubangela amaxesha oxinzelelo oluphakathi ukuya koluqatha lobushushu ngexesha lezigaba ezinobuzaza zokukhula kwerayisi (i-tillering, i-anthesis, intyatyambo, kunye nokuzaliswa kweenkozo), ngaloo ndlela kuchaphazele kakubi isivuno sengqolowa . Oku kuncitshiswa kwesivuno kubangelwa kakhulu ixesha elide loxinzelelo lobushushu, oluchaphazela i-plant physiology . Ngenxa yentsebenziswano yezinto ezahlukeneyo, ezinje ngoxinzelelo lwexesha kunye nobushushu obuphezulu obufikelelweyo, uxinzelelo lobushushu lunokubangela uluhlu lomonakalo ongenakulungiseka kwimetabolism yezityalo kunye nophuhliso.
Uxinzelelo lobushushu luchaphazela iinkqubo ezahlukeneyo ze-physiological and biochemicals kwizityalo. Ifotosynthesis yamagqabi yenye yeenkqubo ezichanabeka kakhulu kuxinzelelo lobushushu kwizityalo zerayisi, njengoko izinga lefotosynthesis lehla nge-50% xa amaqondo obushushu emihla ngemihla edlula kuma-35°C. Iimpendulo zePhysiological zezityalo zerayisi ziyahluka ngokuxhomekeke kuhlobo loxinzelelo lobushushu. Umzekelo, amazinga e-photosynthetic kunye ne-stomatal conductance ayathintelwa xa izityalo zibonakaliswe kumaqondo aphezulu emini (33-40 ° C) okanye amaqondo aphezulu emini nasebusuku (35-40 ° C emini, 28-30 ° C). C ithetha ubusuku) (Lü et al., 2013; Fahad et al., 2016; Chaturvedi et al., 2017). Ukushisa okuphezulu ebusuku (30 ° C) kubangela ukuvinjelwa okuphakathi kwe-photosynthesis kodwa kwandisa ukuphefumla ebusuku (Fahad et al., 2016; Alvarado-Sanabria et al., 2017). Kungakhathaliseki ukuba ixesha loxinzelelo, uxinzelelo lobushushu luchaphazela umxholo we-chlorophyll yeqabunga, umlinganiselo we-chlorophyll variable fluorescence ukuya kwi-chlorophyll fluorescence ephezulu (Fv / Fm), kunye nokusebenza kweRubisco kwizityalo zerayisi (Cao et al. 2009; Yin et al. 2010). ) USanchez Reynoso et al., 2014).
Utshintsho lwe-Biochemicals luyinye inkalo yokulungiswa kwezityalo kuxinzelelo lobushushu (Wahid et al., 2007). Umxholo weProline usetyenziswe njengesalathisi se-biochemical yoxinzelelo lwezityalo (u-Ahmed no-Hassan 2011). I-Proline idlala indima ebalulekileyo kwi-metabolism yezityalo njengoko isebenza njengekhabhoni okanye umthombo we-nitrogen kunye ne-membrane stabilizer phantsi kweemeko eziphezulu zokushisa (i-Sánchez-Reinoso et al., 2014). Ukushisa okuphezulu kuchaphazela ukuzinza kwe-membrane ngokusebenzisa i-lipid peroxidation, ekhokelela ekubunjweni kwe-malondialdehyde (MDA) (Wahid et al., 2007). Ngoko ke, umxholo we-MDA uye wasetyenziselwa ukuqonda ukunyaniseka kwesakhiwo seembrane zeseli phantsi koxinzelelo lobushushu (Cao et al., 2009; Chavez-Arias et al., 2018). Ekugqibeleni, uxinzelelo lobushushu obudibeneyo [37/30 ° C (imini / ubusuku)] landa ipesenti yokuvuza kwe-electrolyte kunye nomxholo we-malondialdehyde kwilayisi (Liu et al., 2013).
Ukusetyenziswa kwabalawuli bokukhula kwezityalo (GRs) kuhlolwe ukunciphisa imiphumo emibi yoxinzelelo lobushushu, njengoko ezi zinto zibandakanyeka ngokusebenzayo kwiimpendulo zezityalo okanye iindlela zokukhusela umzimba malunga noxinzelelo olunjalo (Peleg noBlumwald, 2011; Yin et al., 2011; Ahmed et al., 2015). Ukusetyenziswa ngaphandle kwemithombo yofuzo kube nefuthe elihle ekunyamezeleni uxinzelelo lobushushu kwizityalo ezahlukeneyo. Ucwaningo luye lwabonisa ukuba i-phytohormones efana ne-gibberellins (GA), i-cytokinins (CK), i-auxins (AUX) okanye i-brassinosteroids (BR) ikhokelela ekunyuseni kweentlobo ezahlukeneyo ze-physiological and biochemical variables (Peleg kunye neBlumwald, i-2011; Yin et al. Ren, 2011; Mitler, Z201; 2014). E-Colombia, ukusetyenziswa kwangaphandle kwemithombo yofuzo kunye nefuthe layo kwizityalo zerayisi akukaqondwa ngokupheleleyo kwaye kufundwe. Nangona kunjalo, uphando lwangaphambili lubonise ukuba ukutshiza kwe-foliar ye-BR kunokuphucula ukunyamezela kwerayisi ngokuphucula iimpawu zokutshintshiselwa kwegesi, i-chlorophyll okanye umxholo we-proline wamagqabi e-rice seedling (Quintero-Calderón et al., 2021).
I-Cytokinins idibanisa iimpendulo zezityalo kwiingcinezelo ze-abiotic, kuquka uxinzelelo lobushushu (Ha et al., 2012). Ukongezelela, kuye kwaxelwa ukuba ukusetyenziswa kwangaphandle kwe-CK kunokunciphisa umonakalo wokushisa. Ngokomzekelo, ukusetyenziswa kwangaphandle kwe-zeatin kwandisa izinga le-photosynthetic, i-chlorophyll a kunye ne-b content, kunye nokusebenza kakuhle kwe-electron kwi-creeping bentgrass (Agrotis estolonifera) ngexesha loxinzelelo lobushushu (Xu no-Huang, 2009; Jespersen and Huang, 2015). Ukusetyenziswa kwangaphandle kwe-zeatin kunokuphucula umsebenzi we-antioxidant, ukuphucula ukuhlanganiswa kweeprotheni ezahlukeneyo, ukunciphisa iintlobo ze-oksijini ezisebenzayo (ROS) umonakalo kunye nokuveliswa kwe-malondialdehyde (MDA) kwizicubu zezityalo (Chernyadyev, 2009; Yang et al., 2009). , 2016; Kumar et al., 2020).
Ukusetyenziswa kwe-gibberellic acid kuye kwabonisa impendulo enhle kuxinzelelo lobushushu. Izifundo zibonise ukuba i-GA biosynthesis idibanisa iindlela ezahlukeneyo ze-metabolic kunye nokwandisa ukunyamezela phantsi kweemeko eziphezulu zokushisa (u-Alonso-Ramirez et al. 2009; Khan et al. 2020). Abdel-Nabi et al. (2020) yafumanisa ukuba ukutshiza kwe-foliar ye-GA exogenous (25 okanye 50 mg*L) kunokunyusa izinga le-photosynthetic kunye nomsebenzi we-antioxidant kwizityalo ezi-orenji ezigxininiswe bubushushu xa kuthelekiswa nezityalo zokulawula. Kuye kwaphawulwa ukuba ukusetyenziswa kwangaphandle kwe-HA kwandisa umxholo wokufuma, i-chlorophyll kunye neziqulatho ze-carotenoid kwaye kunciphisa i-lipid peroxidation kwi-date palm (Phoenix dactylifera) phantsi koxinzelelo lobushushu (Khan et al., 2020). I-Auxin nayo idlala indima ebalulekileyo ekulawuleni iimpendulo zokukhula eziguquguqukayo kwiimeko eziphezulu zokushisa (Sun et al., 2012; Wang et al., 2016). Lo mlawuli wokukhula usebenza njengomakishi we-biochemical kwiinkqubo ezahlukeneyo ezifana ne-proline synthesis okanye ukuthotywa phantsi koxinzelelo lwe-abiotic (Ali et al. 2007). Ukongeza, i-AUX iphinda ikhulise umsebenzi we-antioxidant, okhokelela ekunciphiseni kwe-MDA kwizityalo ngenxa yokuncipha kwe-lipid peroxidation (Bielach et al., 2017). USergeev et al. (2018) yabona ukuba kwizityalo ze-pea (i-Pisum sativum) phantsi koxinzelelo lobushushu, umxholo we-proline - i-dimethylaminoethoxycarbonylmethyl) i-naphthylchloromethyl ether (TA-14) iyanda. Kuvavanyo olufanayo, baye baqaphela amanqanaba asezantsi e-MDA kwizityalo ezinyangweyo xa kuthelekiswa nezityalo ezinganyangwanga nge-AUX.
I-Brassinosteroids yenye iklasi yokulawula ukukhula esetyenziselwa ukunciphisa iziphumo zoxinzelelo lobushushu. Ogweno et al. (2008) ingxelo ukuba exogenous BR isitshizi sonyuse isantya photosynthetic net, conductance stomatal kunye nesantya esiphezulu Rubisco carboxylation of tomato (Solanum lycopersicum) izityalo phantsi koxinzelelo lobushushu iintsuku ezisi-8. Ukutshiza nge-Foliar ye-epibrassinosteroids kunokunyusa izinga le-photosynthetic ye-cucumber (Cucumis sativus) izityalo phantsi koxinzelelo lobushushu (Yu et al., 2004). Ukongezelela, ukusetyenziswa kwangaphandle kwe-BR kulibaziseka ukuchithwa kwe-chlorophyll kunye nokwandisa ukusetyenziswa kwamanzi ngokufanelekileyo kunye nobuninzi besivuno se-PSII photochemistry kwizityalo eziphantsi koxinzelelo lobushushu (Holá et al., 2010; Toussagunpanit et al., 2015).
Ngenxa yokutshintsha kwemozulu kunye nokuguquguquka, izityalo zerayisi zijongene namaxesha amaqondo aphezulu emihla ngemihla (Lesk et al., 2016; Garcés, 2020; Federarroz (Federación Nacional de Arroceros), 2021). Kwi-phenotyping yezityalo, ukusetyenziswa kwe-phytonutrients okanye i-biostimulants ifundwe njengesicwangciso sokunciphisa uxinzelelo lobushushu kwiindawo ezikhulayo zerayisi (Alvarado-Sanabria et al., 2017; Calderón-Páez et al., 2021; Quintero-Calderón et al., 2021). Ukongezelela, ukusetyenziswa kwezinto eziguquguqukayo ze-biochemical kunye ne-physiological (ubushushu beqabunga, i-stomatal conductance, i-chlorophyll fluorescence parameters, i-chlorophyll kunye nomxholo wamanzi ahambelanayo, i-malondialdehyde kunye ne-proline synthesis) sisixhobo esithembekileyo sokuhlola izityalo zerayisi phantsi koxinzelelo lokushisa ngaphakathi nakwamanye amazwe (i-Sánchez -Reynoso2001 et al. 2017; Nangona kunjalo, uphando malunga nokusetyenziswa kwe-foliar phytohormonal sprays kwi-rayisi kwinqanaba lendawo lihlala linqabile, ngoko ke, ukufundwa kwe-physiological and biochemical reactions of the application of plant growth regulators kubaluleke kakhulu kwisiphakamiso seendlela ezisebenzayo ze-agronomic zale nto. iparameters zechlorophyll fluorescence kunye nomxholo wamanzi onxulumeneyo) kunye neziphumo zebhayokhemikhali zokusetyenziswa kwamagqabi kwizilawuli ezine zokukhula kwezityalo (AUX, CK, GA kunye neBR). (I-Photosynthetic pigments, malondialdehyde kunye neproline content) Izinto eziguquguqukayo kwiigenotypes ezimbini zerayisi yorhwebo eziphantsi koxinzelelo oludibeneyo lobushushu (ubushushu obuphezulu bemini/ebusuku).
Kolu phononongo, kwenziwa imifuniselo emibini ezimeleyo. I-genotypes Federrose 67 (F67: i-genotype ephuhliswe kumaqondo obushushu aphezulu kwiminyaka elishumi edlulileyo) kunye ne-Federrose 2000 (F2000: i-genotype eyaphuhliswa kwiminyaka elishumi yokugqibela yekhulu lama-20 ebonisa ukuchasana nentsholongwane yeqabunga elimhlophe) yasetyenziswa okokuqala. iimbewu. kunye novavanyo lwesibini, ngokulandelelanayo. Zombini i-genotypes zilinywa ngokubanzi ngamafama aseColombia. Imbewu ityalwe kwiitreyi ze-10-L (ubude obuyi-39.6 cm, ububanzi obuyi-28.8 cm, ubude obuyi-16.8 cm) equlethe umhlaba oyisanti ovunduvundu kunye ne-2% yezinto eziphilayo. Iimbewu ezintlanu ezintshulisiweyo zatyalwa kwitreyi nganye. Iipalati zifakwe kwi-greenhouse ye-Faculty of Agricultural Sciences yeYunivesithi yeSizwe yaseColombia, i-campus yaseBogotá (43 ° 50′56″ N, 74 ° 04′051" W), kwindawo ephakamileyo ye-2556 m ngaphezu kolwandle (asl). m.) kwaye zenziwa ukusuka ngo-Okthobha ukuya kuDisemba ka-2019. Uvavanyo olunye (Federroz 67) kunye novavanyo lwesibini (Federroz 2000) kwangexesha elinye lika-2020.
Iimeko zokusingqongileyo kwi-greenhouse ngexesha lonyaka lokutyala ngalinye zimi ngolu hlobo lulandelayo: iqondo lobushushu emini nasebusuku yi-30/25°C, ukufuma okuhambelanayo ne-60~80%, i-photoperiod yendalo iiyure ezili-12 (imitha ye-photosynthetically esebenzayo 1500 µmol (photons) m-2 s-). 1 emini). Izityalo zavundiswa ngokomxholo wento nganye emva kweentsuku ezingama-20 emva kokuvela kwembewu (DAE), ngokutsho kukaSánchez-Reinoso et al. (2019): 670 mg nitrogen kwisityalo ngasinye, 110 mg phosphorus kwisityalo ngasinye, 350 mg potassium kwisityalo ngasinye, 68 mg calcium kwisityalo ngasinye, 20 mg magnesium kwisityalo ngasinye, 20 mg sulphur kwisityalo ngasinye, 17 mg silicon ngesityalo ngasinye. Izityalo zine-10 mg ye-boron kwisityalo ngasinye, i-17 mg yobhedu kwisityalo ngasinye, kunye ne-44 mg ye-zinc kwisityalo ngasinye. Izityalo zerayisi zagcinwa ukuya kuthi ga kwi-47 DAE kumfuniselo ngamnye xa zifikelela kwisigaba se-phenological V5 ngeli xesha. Izifundo zangaphambili zibonise ukuba eli nqanaba le-phenological lixesha elifanelekileyo lokuqhuba izifundo zokuxinzezeleka kobushushu kwilayisi (i-Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2017).
Kuvavanyo ngalunye, izicelo ezibini ezahlukeneyo zolawulo lokukhula kwamagqabi zenziwa. Iseti yokuqala ye-foliar phytohormone sprays isetyenziswe kwiintsuku ezi-5 ngaphambi konyango loxinzelelo lobushushu (42 DAE) ukulungiselela izityalo zoxinzelelo lwendalo. Isitshizi sesibini samagqabi sanikwa iintsuku ezi-5 emva kokuba izityalo zichanabeke kwiimeko zoxinzelelo (52 DAE). I-phytohormones ezine zisetyenzisiwe kunye neempawu zesithako ngasinye esisebenzayo esifafaziweyo kwesi sifundo zidweliswe kwiThebhile eyoNgezelelweyo 1. Ugxininiso lwabalawuli bokukhula kweqabunga olusetyenzisiweyo lusetyenziswe ngolu hlobo lulandelayo: (i) i-Auxin (1-naphthylacetic acid: NAA) kwi-concentration ye-5 × 10-5 M (ii) 5 × 5 × gibberelli acid; GA3); (iii) Cytokinin (trans-zeatin) 1 × 10-5 M (iv) Brassinosteroids [Spirostan-6-one, 3,5-dihydroxy-, (3b,5a,25R)] 5 × 10-5; M. Ezi ngqwalasela zikhethwe ngenxa yokuba zibangela iimpendulo ezintle kwaye zikhulise ukuchasana kwezityalo kuxinzelelo lobushushu (Zahir et al., 2001; Wen et al., 2010; El-Bassiony et al., 2012; Salehifar et al., 2017). Izityalo zerayisi ngaphandle kwezitshizi zokulawula ukukhula kwezityalo zaphathwa ngamanzi adibeneyo kuphela. Zonke izityalo zerayisi zitshizwe ngesitshizi sesandla. Faka i-20 ml ye-H2O kwisityalo ukunyakamisa iindawo eziphezulu kunye nezisezantsi zamagqabi. Zonke izitshizi zefoliar zisebenzisa i-adjuvant yezolimo (Agrotin, Bayer CropScience, Colombia) kwi-0.1% (v/v). Umgama phakathi kwembiza kunye nesitshizi yi-30 cm.
Unyango loxinzelelo lobushushu lwaqhutywa kwiintsuku ezi-5 emva kwesitshizi sokuqala se-foliar (47 DAE) kumfuniselo ngamnye. Izityalo zerayisi zidluliselwe kwi-greenhouse ukuya kwigumbi lokukhula le-294 L (MLR-351H, Sanyo, IL, USA) ukuseka uxinzelelo lobushushu okanye ukugcina iimeko ezifanayo zokusingqongileyo (47 DAE). Unyango oludityanisiweyo loxinzelelo lobushushu lwenziwa ngokumisela igumbi kumaqondo obushushu alandelayo emini/ebusuku: ubushushu obuphezulu emini [40°C ngeeyure ezi-5 (ukususela ngo-11:00 ukuya ku-16:00)] nexesha lasebusuku [30°C iiyure ezi-5]. Iintsuku ezisi-8 zilandelelana (ukusuka ngo-19:00 ukuya ku-24:00). Ukushisa koxinzelelo kunye nexesha lokuchasa likhethwe ngokusekelwe kwizifundo zangaphambili (i-Sánchez-Reynoso et al. 2014; Alvarado-Sanabría et al. 2017). Ngakolunye uhlangothi, iqela lezityalo ezidluliselwe kwigumbi lokukhula ligcinwe kwi-greenhouse kwiqondo lokushisa elifanayo (30 ° C emini / 25 ° C ebusuku) iintsuku ezi-8 ezilandelelanayo.
Ekupheleni kovavanyo, la maqela alandelayo onyango afunyenwe: (i) imeko yobushushu bokukhula + ukusetyenziswa kwamanzi adityanisiweyo [Ulawulo olupheleleyo (AC)], (ii) imeko yoxinzelelo lobushushu + ukusetyenziswa kwamanzi adityanisiweyo [Ulawulo loxinzelelo lobushushu (SC)], (iii) iimeko zokuxinzezeleka kobushushu + isicelo se-auxin (AUX), (iv) imeko yoxinzelelo lobushushu + i-gibberellin application (GA) kunye nesicelo soxinzelelo sobushushu (i-vtoCK) + brassinosteroid (BR) Isihlomelo. La maqela onyango asetyenziselwa i-genotypes ezimbini (i-F67 kunye ne-F2000). Lonke unyango lwenziwa ngoyilo olungenamkhethe ngokupheleleyo oluneziphindaphindo ezintlanu, nganye ibandakanya isityalo esinye. Isityalo ngasinye sisetyenziselwe ukufunda izinto eziguquguqukayo ezigqitywe ekupheleni kovavanyo. Uvavanyo lwathatha i55 DAE.
I-Stomatal conductance (gs) yalinganiswa kusetyenziswa i-porosometer ephathekayo (SC-1, METER Group Inc., eU.SA) ukusuka kwi-0 ukuya kwi-1000 mmol m-2 s-1, kunye nesampulu yegumbi lokuvula i-6.35 mm. Imilinganiselo ithathwa ngokuncamathisela i-stomameter probe kwigqabi elisele likhulile kwaye ihlumelo eliphambili lesityalo landiswe ngokupheleleyo. Kunyango ngalunye, ufundo lwe-gs lwathathwa kumagqabi amathathu kwisityalo ngasinye phakathi kwe-11:00 kunye ne-16:00 kunye ne-avareji.
I-RWC yamiselwa ngokwendlela echazwe nguGhoulam et al. (2002). Iphepha elandiswe ngokupheleleyo elisetyenziselwe ukumisela g likwasetyenziswe ukulinganisa iRWC. Ubunzima obutsha (FW) bamiselwa ngoko nangoko emva kokuvuna kusetyenziswa isikali sedijithali. Emva koko amagqabi ayefakwa kwisitya seplastiki esizaliswe ngamanzi aze ashiywe ebumnyameni kumaqondo obushushu angaphakathi (ama-22 ngokukaCelsius) kangangeeyure ezingama-48. Emva koko ulinganise kwisikali sedijithali kwaye urekhode ubunzima obandisiweyo (TW). Amagqabi adumbileyo omisiwe kwi-oven kwi-75 ° C kwiiyure ezingama-48 kwaye ubunzima bawo obomileyo (DW) barekhodwa.
Isiqulatho seklorofili esinxulumeneyo simiselwe kusetyenziswa imitha yeklorofili (atLeafmeter, FT Green LLC, USA) kwaye ibonakaliswe kwiiyunithi zeLeaf (Dey et al., 2016). I-PSII ephezulu yokufundwa kwe-quantum esebenzayo (i-Fv / Fm ratio) yarekhodwa kusetyenziswa i-excitation eqhubekayo ye-chlorophyll fluorimeter (i-PeA Handy, i-Hansatech Instruments, e-UK). Amaqabunga ayemnyama-ayenziwe amnyama esebenzisa amaqabunga amaqabunga emizuzu engama-20 ngaphambi kwemilinganiselo yeFv / Fm (Restrepo-Diaz kunye neGarces-Varon, 2013). Emva kokuba amagqabi enziwe mnyama, isiseko (F0) kunye ne-fluorescence ephezulu (Fm) yalinganiswa. Ukusuka kwezi datha, i-fluorescence eguquguqukayo (Fv = Fm - F0), umlinganiselo we-fluorescence eguquguqukayo ukuya kwi-fluorescence ephezulu (Fv / Fm), isivuno esiphezulu se-quantum ye-PSII photochemistry (Fv / F0) kunye nomlinganiselo we-Fm / F0 zibalwe (Baker, 2008; Lee et al., 2017). Ufundo lweklorofili olunxulumeneyo kunye neklorofili ye-fluorescence luthathwe kumagqabi afanayo asetyenziselwa imilinganiselo ye-gs.
Malunga ne-800 mg yobunzima obutsha begqabi yaqokelelwa njengezinto eziguquguqukayo ze-biochemical. Iisampulu zamagqabi zaye zenziwa i-homogenized kwi-nitrogen engamanzi zaze zagcinwa ukuze zihlalutywe ngakumbi. Indlela ye-spectrometric esetyenziselwa ukuqikelela i-tissue chlorophyll a, b kunye nomxholo we-carotenoid isekelwe kwindlela kunye nokulinganisa okuchazwe nguWellburn (1994). Iisampulu zezicubu zamagqabi (30 mg) zaqokelelwa kwaye zenziwa i-homogenized kwi-3 ml ye-80% ye-acetone. Iisampuli zaye zenziwa i-centrifuged (imodeli 420101, i-Becton Dickinson Primary Care Diagnostics, USA) kwi-5000 rpm ye-10 min ukususa amaqhekeza. I-supernatant yahlanjululwa kumthamo wokugqibela we-6 ml ngokongeza i-80% ye-acetone (i-Sims kunye ne-Gamon, i-2002). Umxholo we-chlorophyll unqunywe kwi-663 (i-chlorophyll a) kunye ne-646 (i-chlorophyll b) nm, kunye ne-carotenoids kwi-470 nm usebenzisa i-spectrophotometer (Spectronic BioMate 3 UV-vis, Thermo, USA).
Indlela ye-thiobarbituric acid (TBA) echazwe nguHodges et al. (1999) yayisetyenziselwa ukuvavanya i-membrane lipid peroxidation (MDA). Malunga ne-0.3 g yethishu yamagqabi nayo yadityaniswa i-homogenized kwinitrogen engamanzi. Iisampulu zifakwe kwi-centrifuged kwi-5000 rpm kwaye i-absorbeance yalinganiswa kwi-spectrophotometer kwi-440, 532 kunye ne-600 nm. Ekugqibeleni, ugxininiso lwe-MDA lubalwe kusetyenziswa i-coefficient yokuphela (157 M mL-1).
Umxholo weProline wazo zonke unyango unqunywe ngokusebenzisa indlela echazwe nguBates et al. (1973). Yongeza i-10 ml ye-3% ye-aqueous solution ye-sulfosalicylic acid kwisampuli egciniweyo kunye nokucoca ngephepha le-Whatman filter (No. 2). Emva koko i-2 ml yale filtrate yaphendulwa kunye ne-2 ml ye-ninhydric acid kunye ne-2 ml ye-glacial acetic acid. Umxube ufakwe kwindawo yokuhlambela amanzi kwi-90 ° C ngeyure eli-1. Yeka ukusabela ngokufukamela emkhenkceni. Gcoba ityhubhu ngamandla usebenzisa i-vortex shaker kwaye uchithe isisombululo esiphumela kwi-4 ml ye-toluene. Ukufundwa kwe-Absorbance kunqunywe kwi-520 nm usebenzisa i-spectrophotometer efanayo esetyenziselwa ukulinganisa i-photosynthetic pigments (Spectronic BioMate 3 UV-Vis, Thermo, Madison, WI, USA).
Indlela echazwe nguGerhards et al. (2016) ukubala ubushushu becanopy kunye neCSI. Iifoto ze-Thermal zithathwe ngekhamera ye-FLIR 2 (FLIR Systems Inc., Boston, MA, USA) ngokuchaneka kwe-± 2 ° C ekupheleni kwexesha loxinzelelo. Beka indawo emhlophe emva kwesityalo ukuze ufote. Kwakhona, iifektri ezimbini zazijongwa njengemizekelo yereferensi. Izityalo zibekwe phezu komhlaba omhlophe; enye yayifakwe i-adjuvant yezolimo (i-Agrotin, i-Bayer CropScience, i-Bogotá, i-Colombia) ukulinganisa ukuvulwa kwayo yonke i-stomata [imodi emanzi (i-Twet)], kwaye enye yayiligqabi ngaphandle kwesicelo [imowudi eyomileyo (Tdry)] (Castro -Duque et al., 2020). Umgama phakathi kwekhamera kunye nembiza ngexesha lokufota yi-1 m.
Isalathisi sokunyamezelana sibalwa ngokungangqalanga kusetyenziswa i-stomatal conductance (gs) yezityalo eziphathwayo xa kuthelekiswa nezityalo zokulawula (izityalo ezingenazo unyango lwexinzelelo kunye nabalawuli bokukhula abasetyenziswayo) ukumisela ukunyamezela kwe-genotypes ephathwayo ehlolwe kolu cwaningo. I-RTI yafunyanwa ngokusebenzisa i-equation eguqulelwe kwi-Chávez-Arias et al. (2020).
Kuvavanyo ngalunye, zonke izinto eziguquguqukayo zefiziyoloji ezikhankanywe ngasentla zamiselwa kwaye zarekhodwa kwi-55 DAE kusetyenziswa amagqabi awandisiweyo ngokupheleleyo aqokelelwe kwicanopy ephezulu. Ukongezelela, imilinganiselo yenziwa kwigumbi lokukhula ukuze kugwenywe ukutshintsha iimeko zokusingqongileyo apho izityalo zikhula khona.
Idatha evela kwimifuniselo yokuqala neyesibini yahlalutywa kunye njengoluhlu lwemifuniselo. Iqela ngalinye lovavanyo libandakanya izityalo ezi-5, kwaye isityalo ngasinye senza iyunithi yovavanyo. Uhlalutyo lokuhluka (ANOVA) lwenziwa (P ≤ 0.05). Xa ukungafani okuphawulekayo kufunyenwe, uvavanyo lwe-post hoc lwe-Tukey lusetyenziswe kwi-P ≤ 0.05. Sebenzisa umsebenzi we-arcsine ukuguqula amaxabiso epesenti. Idatha yahlaziywa kusetyenziswa i-Statistix v 9.0 software (i-Analytical Software, Tallahassee, FL, USA) kwaye icwangciswe ngokusebenzisa i-SigmaPlot (inguqulo ye-10.0; i-Systat Software, i-San Jose, CA, USA). Uhlalutyo lwecandelo eliphambili lwenziwa kusetyenziswa isoftware ye-InfoStat 2016 (i-Analysis Software, iYunivesithi yeSizwe yaseCordoba, eArgentina) ukuchonga abalawuli ababalaseleyo bokukhula kwezityalo phantsi kophando.
Itheyibhile 1 ishwankathela i-ANOVA ebonisa imifuniselo, unyango olwahlukileyo, kunye nokusebenzisana kwabo kunye ne-photosynthetic pigments yamagqabi (i-chlorophyll a, b, iyonke, kunye ne-carotenoids), i-malondialdehyde (MDA) kunye nomxholo we-proline, kunye nokuziphatha kwe-stomatal. Isiphumo se-gs, isiqulatho samanzi esinxulumeneyo. (RWC), umxholo we-chlorophyll, i-chlorophyll alpha fluorescence parameters, ukushisa kwesithsaba (PCT) (° C), isalathisi soxinzelelo lwezityalo (CSI) kunye nesalathisi sokunyamezela kwezityalo zerayisi kwi-55 DAE.
Itheyibhile 1. Isishwankathelo sedatha ye-ANOVA kwirayisi ye-physiological and biochemical variables phakathi kwezilingo (i-genotypes) kunye nonyango loxinzelelo lobushushu.
Umahluko (P≤0.01) kwintsebenziswano ye-photosynthetic pigment yeqabunga, umxholo we-chlorophyll ohambelanayo (ukufundwa kwe-Atleaf), kunye ne-alpha-chlorophyll fluorescence parameters phakathi kovavanyo kunye nonyango kuboniswe kwiThebhile 2. Ukushisa okuphakamileyo emini nasebusuku kwandisa i-chlorophyll epheleleyo kunye nemixholo ye-carotenoid. Izithole zerayisi ezingenaso nasiphi na isitshizi samagqabi sephytohormones (2.36 mg g-1 for “F67″ kunye 2.56 mg g-1 for “F2000″) xa kuthelekiswa nezityalo ezikhuliswe phantsi kweemeko zobushushu obuphezulu (2.67 mg g -1)) zibonise isiqulatho esisezantsi seklorofili. Kuzo zombini iimvavanyo, "i-F67" yayingu-2.80 mg g-1 kwaye "F2000" yayingu-2.80 mg g-1. Ukongeza, izithole zerayisi ziphathwe ngendibaniselwano ye-AUX kunye ne-GA izitshizi phantsi koxinzelelo lobushushu nazo zibonise ukuncipha komxholo we-chlorophyll kuzo zombini i-genotypes (AUX = 1.96 mg g-1 kunye ne-GA = 1.45 mg g-1 ye-"F67"; AUX = 1.96 mg g-1 kunye ne-"F6" = 4 g-1 kunye ne-"F6" ye-GA = 1. 2.24 mg) g-1 kunye ne-GA = 1.43 mg g-1 (ye "F2000" ) phantsi kweemeko zoxinzelelo lobushushu. Phantsi kweemeko zoxinzelelo lobushushu, unyango lwe-foliar kunye ne-BR lubangele ukunyuka okuncinci kolu tshintsho kuzo zombini ii-genotypes. Ekugqibeleni, isitshizi se-CK foliar sabonisa awona maxabiso aphezulu e-photosynthetic pigment phakathi kwazo zonke unyango (AUX, GA, BR, SC kunye nonyango lwe-AC) kwi-genotypes F67 (3.24 mg g-1) kunye ne-F2000 (3.65 mg g-1). Isiqulatho esinxulumeneyo seklorofili (iyunithi ye-Atleaf) nayo yancitshiswa luxinzelelo oludityanisiweyo lobushushu. Awona maxabiso aphezulu aye abhalwa kwizityalo ezitshizwe ngeCC kuzo zombini iigenotypes (41.66 ye "F67" kunye ne-49.30 ye "F2000"). Umlinganiselo weFv kunye neFv/Fm ubonise umahluko omkhulu phakathi konyango kunye nemihlanganiselwa (Itheyibhile 2). Ngokubanzi, phakathi kwezi ziguquguqukayo, umhlanganisela weF67 wawungachaphazeleki kangako kuxinzelelo lobushushu kunomhlanganisela weF2000. Umlinganiselo weFv kunye neFv/Fm uhlupheke ngakumbi kuvavanyo lwesibini. Izithole ezigxininisiweyo ze-'F2000′ ezingazange zitshizwe ngayo nayiphi na i-phytohormones zinexabiso eliphantsi le-Fv (2120.15) kunye ne-Fv/Fm ratios (0.59), kodwa ukutshiza nge-foliar nge-CK kwanceda ukubuyisela la maxabiso (Fv: 2591, 89, Fv / Fm 3 ratio).: 0. , ukufumana ukufundwa okufana nezo zirekhodwe kwizityalo "ze-F2000" ezikhule phantsi kweemeko ezifanelekileyo zokushisa (Fv: 2955.35, Fv / Fm ratio: 0.73: 0.72). Kwakungekho nantlukwano ebalulekileyo kwi-fluorescence yokuqala (F0), i-fluorescence ephezulu (Fm), isivuno esiphezulu se-photochemical quantum ye-PSII (Fv / F0) kunye ne-Fm / F0. Ekugqibeleni, i-BR ibonise indlela efanayo njengoko ibonwa nge-CK (Fv 2545.06, Fv / Fm ratio 0.73).
Itheyibhile 2. Impembelelo yoxinzelelo lobushushu obudibeneyo (40°/30°C imini/ubusuku) kwi-photosynthetic pigments emagqabini [itotali ye-chlorophyll (i-Chl Total), i-chlorophyll a (Chl a), i-chlorophyll b (i-Chl b) kunye ne-carotenoids Cx+c] isiphumo], iyunithi ye-chlorophyll ehambelanayo (iyunithi ye-chlorophylliffophyll) iyunithi ye-chlorophyll i-fluorescence (F0), i-fluorescence ephezulu (Fm), i-fluorescence eguquguqukayo (Fv), ubuninzi be-PSII obusebenzayo (Fv / Fm), isivuno esiphezulu se-photochemical quantum ye-PSII (Fv / F0) kunye ne-Fm / F0 kwizityalo ezimbini ze-rayisi genotypes [Federrose 67 (F67) kunye ne-Federrose 000 emva kweentsuku ezingama-20 ze-20 (DAE)).
Isiqulatho samanzi esinxulumeneyo (i-RWC) yezityalo zerayisi eziphathwe ngokwahlukileyo zibonise umahluko (P ≤ 0.05) kwintsebenziswano phakathi konyango lovavanyo kunye ne-foliar (Umfanekiso we-1A). Xa uphathwa nge-SA, awona maxabiso aphantsi abhalwe kuzo zombini ii-genotypes (74.01% ye-F67 kunye ne-76.6% ye-F2000). Ngaphantsi kweemeko zokuxinzezeleka kobushushu, i-RWC yezityalo zerayisi zombini i-genotypes ephathwa nge-phytohormones eyahlukeneyo yanda kakhulu. Ngokubanzi, izicelo ze-Foliar ze-CK, GA, AUX, okanye i-BR zonyuse i-RWC ukuya kumaxabiso afana nezityalo ezikhuliswe phantsi kweemeko ezifanelekileyo ngexesha lovavanyo. Ulawulo olupheleleyo kunye nezityalo ezitshizwe ngamahlamvu ezirekhodiweyo malunga ne-83% yazo zombini i-genotypes. Ngakolunye uhlangothi, i-gs nayo ibonise ukungafani okuphawulekayo (P ≤ 0.01) kwi-experiment-treatment interaction (Fig. 1B). Isityalo solawulo olupheleleyo (AC) siphinde sabhala amaxabiso aphezulu kwi-genotype nganye (440.65 mmol m-2s-1 ye-F67 kunye ne-511.02 mmol m-2s-1 ye-F2000). Izityalo zerayisi eziphantsi koxinzelelo lobushushu oludityanisiweyo zizodwa zibonise awona maxabiso aphantsi e-gs kuzo zombini i-genotypes (150.60 mmol m-2s-1 ye-F67 kunye ne-171.32 mmol m-2s-1 ye-F2000). Unyango lwamagqabi kunye nazo zonke izilawuli zokukhula kwezityalo nalo lwandisiwe g. Kwizityalo zerayisi ze-F2000 ezitshizwe nge-CC, umphumo wokutshiza nge-foliar nge-phytohormones wawucace ngakumbi. Eli qela lezityalo alibonisi mahluko xa kuthelekiswa nezityalo zokulawula ngokupheleleyo (AC 511.02 kunye ne-CC 499.25 mmol m-2s-1).
Umzobo 1. Impembelelo yoxinzelelo lobushushu obudibeneyo (40 ° / 30 ° C imini / ubusuku) kumxholo wamanzi ohambelanayo (RWC) (A), i-stomatal conductance (gs) (B), i-malondialdehyde (MDA) imveliso (C), kunye nomxholo weproline. (D) kwizityalo ezimbini zerayisi genotypes (F67 kunye F2000) kwiintsuku ezingama-55 emva kokuvela (DAE). Unyango oluvavanyiweyo kwi-genotype nganye lubandakanya: ulawulo olupheleleyo (AC), ukulawula uxinzelelo lobushushu (SC), uxinzelelo lobushushu + auxin (AUX), uxinzelelo lobushushu + i-gibberellin (GA), uxinzelelo lobushushu + i-cell mitogen (CK), kunye noxinzelelo lobushushu + brassinosteroid. (BR). Ikholamu nganye imele intsingiselo ± impazamo eqhelekileyo yamanqaku amahlanu edatha (n = 5). Iikholomu ezilandelwa ngoonobumba abahlukeneyo zibonisa ukungafani okuphawulekayo ngokwezibalo ngokovavanyo lukaTukey (P ≤ 0.05). Iileta ezinophawu olulinganayo zibonisa ukuba intsingiselo ayibalulekanga ngokwezibalo (≤ 0.05).
I-MDA (P ≤ 0.01) kunye neproline (P ≤ 0.01) iziqulatho nazo zibonise ukungafani okuphawulekayo kwintsebenziswano phakathi kovavanyo kunye nonyango lwe-phytohormone (umzobo 1C, D). Ukunyuka kwe-lipid peroxidation kwabonwa ngonyango lwe-SC kuzo zombini i-genotypes (Umfanekiso we-1C), nangona kunjalo izityalo eziphathwe nge-spray yokulawula ukukhula kwamagqabi zibonise ukuhla kwe-lipid peroxidation kuzo zombini i-genotypes; Ngokuqhelekileyo, ukusetyenziswa kwe-phytohormones (CA, AUC, BR okanye GA) kukhokelela ekunciphiseni kwe-lipid peroxidation (umxholo we-MDA). Akukho mahluko ufunyenwe phakathi kwezityalo ze-AC ze-genotypes ezimbini kunye nezityalo phantsi koxinzelelo lobushushu kunye nokutshizwa nge-phytohormones (amaxabiso e-FW aqatshelweyo kwizityalo "F67" avela kwi-4.38-6.77 µmol g-1, kunye ne-FW "F2000" izityalo "amaxabiso agcinwe ukusuka kwi-9.18 ukuya kwi-18 g ukuya kwi-2. ngesandla, iproline synthesis kwi "F67" yezityalo yayingaphantsi kune "F2000" izityalo phantsi koxinzelelo oludibeneyo, olukhokelele ekwandeni kwemveliso yeproline kwizityalo zerayisi ezigxininiswe ngubushushu, kuzo zombini iimvavanyo, kwaphawulwa ukuba ukulawulwa kwezi hormone kwandisa kakhulu umxholo we-amino acid yezityalo ze-F2000 (AUX kunye ne-BR43l µ ne-118 g. (Umfanekiso 1G).
Iimpembelelo zesitshizi sokulawula ukukhula kwesityalo esine-foliar kunye noxinzelelo lobushushu oludityanisiweyo kwiqondo lobushushu besityalo kunye nesalathiso sokunyamezelana (RTI) kubonisiwe kwiMizobo 2A kunye no-B. Kuzo zombini iigenotypes, ubushushu becanopy yezityalo ze-AC babusondele kuma-27°C, kwaye obo bezityalo ze-SC babumalunga ne-28°C. KUNYE. Kwaye kwaphawulwa ukuba unyango lwe-foliar kunye ne-CK kunye ne-BR lubangele ukuhla kwe-2-3 ° C kubushushu be-canopy xa kuthelekiswa nezityalo ze-SC (Figure 2A). I-RTI ibonise ukuziphatha okufanayo kwezinye izinto eziguquguqukayo ze-physiological, ezibonisa ukungafani okuphawulekayo (P ≤ 0.01) ekusebenzisaneni phakathi kovavanyo kunye nonyango (Umfanekiso 2B). Izityalo ze-SC zibonise ukunyamezela kwezityalo ezisezantsi kuzo zombini i-genotypes (34.18% kunye ne-33.52% yezityalo zerayisi "F67" kunye ne "F2000", ngokulandelanayo). Ukondliwa kwe-foliar ye-phytohormones kuphucula i-RTI kwizityalo ezivezwe kuxinzelelo oluphezulu lobushushu. Esi siphumo sasibonakaliswe ngakumbi kwizityalo "ze-F2000" ezifafazwe nge-CC, apho i-RTI yayiyi-97.69. Ngakolunye uhlangothi, ukungafani okuphawulekayo kwabonwa kuphela kwisalathisi soxinzelelo lwesivuno (CSI) sezityalo zerayisi phantsi kweemeko ze-foliar factor spray stress (P ≤ 0.01) (Fig. 2B). Kuphela izityalo zerayisi eziphantsi koxinzelelo olunzima lobushushu zibonise ixabiso eliphezulu loxinzelelo (0.816). Xa izityalo zerayisi zitshizwe ngeephytohormones ezahlukeneyo, isalathisi soxinzelelo sasisezantsi (amaxabiso ukusuka ku-0.6 ukuya ku-0.67). Ekugqibeleni, isityalo selayisi esikhule phantsi kweemeko ezifanelekileyo sinexabiso le-0.138.
Umfanekiso 2. Iziphumo zoxinzelelo lobushushu obudibeneyo (40°/30°C imini/ebusuku) kubushushu obuphezulu (A), isalathiso sokunyamezelana (RTI) (B), kunye nesalathisi soxinzelelo lwezityalo (CSI) (C) zeentlobo ezimbini zezityalo. I-genotypes yerayisi yorhwebo (i-F67 kunye ne-F2000) yayiphantsi konyango oluhlukeneyo lobushushu. Unyango oluvavanyiweyo kwi-genotype nganye lubandakanya: ulawulo olupheleleyo (AC), ukulawula uxinzelelo lobushushu (SC), uxinzelelo lobushushu + auxin (AUX), uxinzelelo lobushushu + i-gibberellin (GA), uxinzelelo lobushushu + i-cell mitogen (CK), kunye noxinzelelo lobushushu + brassinosteroid. (BR). Uxinzelelo lobushushu oludityanisiweyo lubandakanya ukubeka izityalo zerayisi kumaqondo obushushu aphezulu emini/ebusuku (40°/30°C imini/ubusuku). Ikholamu nganye imele intsingiselo ± impazamo eqhelekileyo yamanqaku amahlanu edatha (n = 5). Iikholomu ezilandelwa ngoonobumba abahlukeneyo zibonisa ukungafani okuphawulekayo ngokwezibalo ngokovavanyo lukaTukey (P ≤ 0.05). Iileta ezinophawu olulinganayo zibonisa ukuba intsingiselo ayibalulekanga ngokwezibalo (≤ 0.05).
Uhlalutyo lwecandelo eliphambili (PCA) lubonise ukuba iinguqu ezihlolwe kwi-55 DAE zichaze i-66.1% yeempendulo ze-physiological and biochemical of the heat-stressed rice plantated treatment of growth regulator spray (Fig. 3). IiVectors zimela izinto eziguquguqukayo kwaye amachaphaza amele izilawuli zokukhula kwezityalo (GRs). I-vectors ye-gs, umxholo we-chlorophyll, ubuninzi be-quantum ye-PSII (Fv / Fm) kunye ne-biochemical parameters (TChl, MDA kunye ne-proline) i-angles esondeleyo kwimvelaphi, ebonisa ukuhambelana okuphezulu phakathi kokuziphatha kwe-physiological yezityalo kunye nabo. iyaguquguquka. Iqela elinye (V) libandakanya izithole zerayisi ezikhuliswe kubushushu obuphezulu (AT) kunye nezityalo ze-F2000 ezinyangwa nge-CK kunye ne-BA. Ngelo xesha, uninzi lwezityalo eziphathwe nge-GR zenza iqela elihlukeneyo (IV), kwaye unyango kunye ne-GA kwi-F2000 lwenza iqela elahlukileyo (II). Ngokuchaseneyo, izithole zerayisi ezigxininiswe bubushushu (amaqela I kunye ne-III) ngaphandle kokutshiza kwe-foliar ye-phytohormones (zombini i-genotypes yayiyi-SC) zazibekwe kwindawo echasene neqela V, ebonisa isiphumo soxinzelelo lobushushu kwi-physiology yezityalo. .
Umzobo 3. Uhlalutyo lwe-Bigraphical yemiphumo yoxinzelelo lobushushu obudibeneyo (40 ° / 30 ° C imini / ubusuku) kwizityalo ezimbini ze-rice genotypes (F67 kunye ne-F2000) kwiintsuku ze-55 emva kokuvela (DAE). Izifinyezo: AC F67, ulawulo olupheleleyo F67; SC F67, ukulawula uxinzelelo lobushushu F67; I-AUX F67, uxinzelelo lobushushu + auxin F67; GA F67, uxinzelelo lobushushu + gibberellin F67; CK F67, uxinzelelo lobushushu + ulwahlulo lweseli BR F67, uxinzelelo lobushushu + brassinosteroid. F67; AC F2000, ulawulo olupheleleyo F2000; SC F2000, Ulawulo loxinzelelo lobushushu F2000; I-AUX F2000, uxinzelelo lobushushu + auxin F2000; GA F2000, uxinzelelo lobushushu + gibberellin F2000; CK F2000, uxinzelelo lobushushu + cytokinin, BR F2000, uxinzelelo lobushushu + i-steroid yobhedu; F2000.
Iinguqu ezifana nomxholo we-chlorophyll, i-stomatal conductance, i-Fv / Fm ratio, i-CSI, i-MDA, i-RTI kunye nomxholo we-proline inokunceda ukuqonda ukulungelelaniswa kwe-rice genotypes kunye nokuvavanya impembelelo yezicwangciso ze-agronomic phantsi koxinzelelo lobushushu (Sarsu et al., 2018; Quintero-Calderon et al., 2021). Injongo yolu vavanyo yayikukuvavanya umphumo wokusetyenziswa kwabalawuli abane bokukhula kwiiparamitha ze-physiological kunye ne-biochemical yezithole zerayisi phantsi kweemeko ezinzima zoxinzelelo lobushushu. Ukuvavanywa kwezithole yindlela elula kwaye ekhawulezayo yokuvavanya ngaxeshanye izityalo zerayisi kuxhomekeke kubungakanani okanye imeko yeziseko ezikhoyo (Sarsu et al. 2018). Iziphumo zolu pho nonongo zibonise ukuba uxinzelelo lobushushu oludibeneyo lubangela iimpendulo ezahlukeneyo ze-physiological and biochemical in the two rice genotypes, ebonisa inkqubo yokulungelelanisa. Ezi ziphumo zikwabonisa ukuba izitshizi zokulawula ukukhula kwefoliar (ikakhulu iicytokinins kunye nebrassinosteroids) zinceda irayisi ukuba iqhelane noxinzelelo lobushushu oluntsonkothileyo njengoko ithandwa ichaphazela ikakhulu i-gs, RWC, Fv/Fm ratio, photosynthetic pigments kunye nomxholo weproline.
Ukusetyenziswa kwezilawuli zokukhula kunceda ekuphuculeni ubume bamanzi bezityalo zerayisi phantsi koxinzelelo lobushushu, ezinokunxulunyaniswa noxinzelelo oluphezulu kunye namaqondo asezantsi ekhenopi yezityalo. Olu phononongo lubonise ukuba phakathi kwezityalo "ze-F2000" (i-genotype) izityalo, izityalo zerayisi ziphathwa ngokuyintloko nge-CK okanye i-BR zinexabiso eliphezulu le-gs kunye nexabiso eliphantsi le-PCT kunezityalo eziphathwa nge-SC. Uphononongo lwangaphambili luye lwabonisa ukuba i-gs kunye ne-PCT zizibonakaliso ezichanekileyo ze-physiological ezinokuthi zichaze impendulo eguquguqukayo yezityalo zerayisi kunye nemiphumo yezicwangciso ze-agronomic kwixinzelelo lobushushu (Restrepo-Diaz kunye neGarces-Varon, 2013; Sarsu et al., 2018; Quintero). -Carr DeLong et al., 2021). I-Leaf CK okanye i-BR iphucula i-g phantsi koxinzelelo ngenxa yokuba ezi hormone zezityalo zinokukhuthaza ukuvulwa kwe-stomatal ngokusebenzisa intsebenziswano yokwenziwa kunye nezinye iimolekyuli zokubonisa ezifana ne-ABA (umgqugquzeli wokuvalwa kwe-stomatal phantsi koxinzelelo lwe-abiotic) (Macková et al., 2013; Zhou et al., 2013). 2013). ). , 2014). Ukuvulwa kwe-stomatal kukhuthaza ukupholisa kwamagqabi kwaye kunceda ukunciphisa ukushisa kwe-canopy (Sonjaroon et al., 2018; Quintero-Calderón et al., 2021). Ngezi zizathu, ubushushu becanopy yezityalo zerayisi etshizwe nge-CK okanye i-BR bunokuba sezantsi phantsi koxinzelelo lobushushu oludibeneyo.
Uxinzelelo oluphezulu lweqondo lokushisa kunokunciphisa umxholo we-photosynthetic pigment wamagqabi (Chen et al., 2017; Ahammed et al., 2018). Kolu phononongo, xa izityalo zerayisi ziphantsi koxinzelelo lobushushu kwaye zingatshizwanga naziphi na izilawuli zokukhula kwezityalo, iipigments ze-photosynthetic zithande ukuncipha kuzo zombini i-genotypes (Itheyibhile 2). Feng et al. (I-2013) iphinde ichaze ukwehla okukhulu kumxholo we-chlorophyll kumagqabi ee-genotypes ezimbini zengqolowa ezivezwe kuxinzelelo lobushushu. Ukuvezwa kwamaqondo obushushu aphezulu kudla ngokukhokelela ekunciphiseni komxholo we-chlorophyll, onokuthi kungenxa yokuncipha kwe-chlorophyll biosynthesis, ukuthotywa kwe-pigment, okanye iziphumo zabo ezidibeneyo phantsi koxinzelelo lobushushu (Fahad et al., 2017). Nangona kunjalo, izityalo zerayisi eziphathwa kakhulu nge-CK kunye ne-BA zonyuse ukuxinwa kwee-pigments ze-photosynthetic zamagqabi phantsi koxinzelelo lobushushu. Iziphumo ezifanayo zichazwe nguJespersen noHuang (2015) kunye no-Suchsagunpanit et al. (2015), owabona ukwanda komxholo we-chlorophyll yeqabunga emva kokusetyenziswa kwe-zeatin kunye ne-epibrassinosteroid hormones kwi-bentgrass egxininiswe ngubushushu kunye nerayisi, ngokulandelanayo. Inkcazo enengqiqo yokuba kutheni i-CK kunye ne-BR ikhuthaza ukunyuka kwe-chlorophyll yeqabunga phantsi koxinzelelo lobushushu obudibeneyo kukuba i-CK inokunyusa ukuqaliswa kwe-induction eqhubekayo yabakhuthazi bentetho (njenge-senescence-activating promoter (SAG12) okanye umgqugquzeli we-HSP18) kunye nokunciphisa ilahleko ye-chlorophyll kumagqabi. , ukulibazisa ukuvela kwamagqabi kunye nokwandisa ukuxhathisa kwezityalo kubushushu (Liu et al., 2020). I-BR inokukhusela i-chlorophyll yeqabunga kwaye ikhulise umxholo we-chlorophyll yeqabunga ngokusebenza okanye ukukhuthaza ukuhlanganiswa kwee-enzymes ezibandakanyekayo kwi-chlorophyll biosynthesis phantsi kweemeko zoxinzelelo (uSharma et al., 2017; Siddiqui et al., 2018). Ekugqibeleni, i-phytohormones ezimbini (i-CK kunye ne-BR) iphinda ikhuthaze ukubonakaliswa kweeprotheyini zokutshatyalaliswa kobushushu kunye nokuphucula iinkqubo ezahlukeneyo zokulungiswa kwe-metabolic, ezifana nokunyuka kwe-chlorophyll biosynthesis (uSharma et al., 2017; Liu et al., 2020).
I-Chlorophyll i-fluorescence parameters ibonelela ngendlela ekhawulezayo nengonakalisiyo enokuvavanya ukunyamezela kwezityalo okanye ukuziqhelanisa neemeko zoxinzelelo lwe-abiotic (Chaerle et al. 2007; Kalaji et al. 2017). Iiparamitha ezifana nomlinganiselo weFv / Fm zisetyenziswe njengezibonakaliso zokulungiswa kwezityalo kwiimeko zoxinzelelo (Alvarado-Sanabria et al. 2017; Chavez-Arias et al. 2020). Kolu phononongo, izityalo ze-SC zibonise awona maxabiso aphantsi kolu tshintsho, ikakhulu "F2000" izityalo zerayisi. Yin et al. (2010) kwakhona kwafumanisa ukuba umlinganiselo weFv/Fm welona gqabi liphezulu lokuhlakula irayisi lehle kakhulu kumaqondo obushushu angaphezu kwama-35°C. Ngokutsho kukaFeng et al. (2013), umlinganiselo ophantsi weFv/Fm phantsi koxinzelelo lobushushu ubonisa ukuba izinga lokubanjwa kwamandla okuvuselelayo nokuguqulwa liziko lokusabela le-PSII liyancitshiswa, nto leyo ebonisa ukuba iziko lokusabela le-PSII lichithakala phantsi koxinzelelo lobushushu. Olu qwalaselo lusivumela ukuba sigqibe kwelokuba iziphazamiso kwizixhobo zephotosynthetic zibonakala ngakumbi kwiintlobo ezibuthathaka (Fedearroz 2000) kuneentlobo ezixhathisayo (Fedearroz 67).
Ukusetyenziswa kwe-CK okanye i-BR ngokubanzi kuphuculwe ukusebenza kwe-PSII phantsi kweemeko ezinzima zoxinzelelo lobushushu. Iziphumo ezifanayo zafunyanwa ngu Suchsagunpanit et al. (2015), owabona ukuba isicelo se-BR sandise ukusebenza kakuhle kwe-PSII phantsi koxinzelelo lobushushu kwirayisi. Kumar et al. (I-2020) iphinde yafumanisa ukuba izityalo ze-chickpea eziphathwe nge-CK (i-6-benzyladenine) kwaye ziphantsi koxinzelelo lobushushu zanda umlinganiselo we-Fv / Fm, ukugqiba ukuba isicelo se-foliar se-CK ngokuvula umjikelo we-pigment ye-zeaxanthin wakhuthaza umsebenzi we-PSII. Ukongeza, i-BR leaf spray ithande i-PSII photosynthesis phantsi kweemeko zoxinzelelo ezidibeneyo, ebonisa ukuba ukusetyenziswa kwale phytohormone kubangele ukuncipha kokuchithwa kwamandla okuvuselela amandla eempondo ze-PSII kunye nokukhuthaza ukuqokelelwa kweeprotheyini zokutshatyalaliswa kobushushu kwi-chloroplasts (Ogweno et al. 2008; Kothari kunye neLachowitz). , 2021).
I-MDA kunye ne-proline contents zihlala zinyuka xa izityalo ziphantsi koxinzelelo lwe-abiotic xa kuthelekiswa nezityalo ezikhulile phantsi kweemeko ezifanelekileyo (Alvarado-Sanabria et al. 2017). Uphononongo lwangaphambili luye lwabonisa ukuba i-MDA kunye namanqanaba eproline zizibonakaliso ze-biochemical ezingasetyenziselwa ukuqonda inkqubo yokulungelelanisa okanye impembelelo yezenzo ze-agronomic kwirayisi phantsi kwemini okanye ebusuku ukushisa okuphezulu (Alvarado-Sanabria et al., 2017; Quintero-Calderón et al. . , 2021). Ezi zifundo zibonise kwakhona ukuba i-MDA kunye ne-proline contents ithande ukuba phezulu kwizityalo zerayisi ezivezwe kumaqondo aphezulu ebusuku okanye emini, ngokulandelanayo. Nangona kunjalo, ukutshizwa kwe-foliar ye-CK kunye ne-BR kube negalelo ekunciphiseni kwe-MDA kunye nokunyuka kwamanqanaba e-proline, ngokukodwa kwi-genotype enyamezelayo (Federroz 67). I-CK spray inokukhuthaza ukugqithiswa kwe-cytokinin oxidase / dehydrogenase, ngaloo ndlela ikhulise umxholo wezinto ezikhuselayo ezifana ne-betaine kunye neproline (Liu et al., 2020). I-BR ikhuthaza ukufakwa kwee-osmoprotectants ezifana ne-betaine, ishukela, kunye ne-amino acids (kubandakanywa ne-proline yamahhala), ukugcina ibhalansi ye-osmotic yeselula phantsi kweemeko ezininzi ezimbi zendalo (i-Kothari ne-Lachowiec, i-2021).
Isalathiso soxinzelelo lwezityalo (CSI) kunye nesalathiso sokunyamezelana (RTI) zisetyenziselwa ukufumanisa ukuba unyango oluvavanywayo lunceda ukunciphisa uxinzelelo olwahlukeneyo (i-abiotic kunye ne-biotic) kwaye lube nefuthe elihle kwi-physiology yezityalo (Castro-Duque et al., 2020; Chavez-Arias et al., 2020). Amaxabiso e-CSI anokuqala ukusuka kwi-0 ukuya kwi-1, emele iimeko zokungaxinzeki kunye noxinzelelo, ngokulandelanayo (Lee et al., 2010). Ixabiso le-CSI yezityalo ezigxininiswe ukushisa (SC) ukusuka kwi-0.8 ukuya kwi-0.9 (Umfanekiso we-2B), ebonisa ukuba izityalo zerayisi zazichatshazelwa kakubi ngoxinzelelo oludibeneyo. Nangona kunjalo, ukutshiza nge-foliar ye-BC (0.6) okanye i-CK (0.6) ikakhulu kukhokelele ekunciphiseni kwesi salathisi phantsi kweemeko zoxinzelelo lwe-abiotic xa kuthelekiswa nezityalo zerayisi ze-SC. Kwizityalo ze-F2000, i-RTI ibonise ukunyuka okuphezulu xa isebenzisa i-CA (97.69%) kunye ne-BC (60.73%) xa kuthelekiswa ne-SA (33.52%), ebonisa ukuba aba balawuli bokukhula kwezityalo nabo bafaka isandla ekuphuculeni impendulo yerayisi ekunyamezeleni ukubunjwa. Ubushushu obugqithisileyo. Ezi indices ziye zacetywa ukulawula iimeko zoxinzelelo kwiintlobo ezahlukeneyo. Uphononongo olwenziwe nguLee et al. (2010) ibonise ukuba i-CSI yeentlobo ezimbini zekotoni phantsi koxinzelelo lwamanzi oluphakathi malunga ne-0.85, ngelixa ixabiso le-CSI yeentlobo ezinkcenkceshelwe kakuhle zivela kwi-0.4 ukuya kwi-0.6, zigqiba ukuba esi salathisi sisalathisi sokulungelelaniswa kwamanzi kweentlobo. iimeko ezicinezelayo. Ngaphezu koko, uChavez-Arias et al. (2020) ivavanye ukusebenza kwe-elicitors zokwenziwa njengesicwangciso esibanzi solawulo loxinzelelo kwizityalo ze-C. elegans kwaye yafumanisa ukuba izityalo ezitshizwe ngezi zixhobo zibonise i-RTI ephezulu (65%). Ngokusekelwe apha ngasentla, i-CK kunye ne-BR inokuthathwa njengezicwangciso ze-agronomic ezijoliswe ekwandiseni ukunyamezela kwerayisi kuxinzelelo olunzima lobushushu, njengoko aba balawuli bokukhula kwezityalo babangela iimpendulo ezintle ze-biochemical kunye ne-physiological.
Kwiminyaka embalwa edlulileyo, uphando lwerayisi eColombia lugxininise ekuvavanyeni i-genotypes enyamezelayo ukuya kwimini ephezulu okanye ubushushu bobusuku usebenzisa iimpawu ze-physiological okanye ze-biochemical (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2021). Nangona kunjalo, kwiminyaka embalwa edlulileyo, uhlalutyo lobugcisa obusebenzayo, obunoqoqosho kunye nenzuzo luye lwaba lubaluleke kakhulu ukuphakamisa ukulawulwa kwezityalo ezihlangeneyo ukuphucula imiphumo yamaxesha anzima oxinzelelo lobushushu kwilizwe (Calderón-Páez et al., 2021; Quintero-Calderon et al., 2021) . Ngaloo ndlela, iimpendulo ze-physiological kunye ne-biochemical yezityalo zerayisi kuxinzelelo olunzima lobushushu (i-40 ° C imini / 30 ° C ebusuku) ebonwe kolu phononongo lubonisa ukuba ukutshiza nge-foliar nge-CK okanye i-BR ingaba yindlela efanelekileyo yokulawula isityalo ukunciphisa iziphumo ezibi. Isiphumo samaxesha oxinzelelo oluphakathi lobushushu. Olu nyango luphucule ukunyamezela kwazo zombini i-rice genotypes (i-CSI ephantsi kunye ne-RTI ephezulu), ebonisa indlela eqhelekileyo kwiimpendulo zezityalo ze-physiological and biochemicals phantsi koxinzelelo lobushushu obudibeneyo. Impendulo ephambili yezityalo zerayisi yayikukuncipha kumxholo we-GC, i-chlorophyll epheleleyo, i-chlorophylls α kunye ne-β kunye ne-carotenoids. Ukongezelela, izityalo zihlupheka ngenxa yomonakalo we-PSII (ukunciphisa i-chlorophyll fluorescence parameters ezifana ne-Fv / Fm ratio) kunye nokunyuka kwe-lipid peroxidation. Ngakolunye uhlangothi, xa irayisi iphathwa nge-CK kunye ne-BR, le miphumo emibi yancitshiswa kwaye umxholo weproline wanda (umzobo 4).
Umzobo 4. Imodeli yengqiqo yeziphumo zoxinzelelo oludibeneyo lobushushu kunye ne-foliar plant growth regulator isitshizi kwizityalo zerayisi. Iintolo ezibomvu neziluhlaza zibonisa imiphumo emibi okanye emihle yokusebenzisana phakathi koxinzelelo lobushushu kunye nokusetyenziswa kwe-foliar ye-BR (brassinosteroid) kunye ne-CK (cytokinin) kwiimpendulo ze-physiological and biochemical, ngokulandelanayo. gs: ukuqhuba kwesisu; Iyonke i-Chl: umxholo weklorofili iyonke; Chl α: umxholo we-chlorophyll β; Cx+c: umxholo we-carotenoid;
Isishwankathelo, iimpendulo ze-physiological kunye ne-biochemical kolu phononongo zibonisa ukuba izityalo zerayisi ze-Fedearroz 2000 zichaphazeleka ngakumbi kwixesha loxinzelelo olunzima lobushushu kuneFedearroz 67 izityalo zerayisi. Zonke izilawuli zokukhula ezihlolwe kolu phando (i-auxins, i-gibberellins, i-cytokinins, okanye i-brassinosteroids) ibonise iqondo elithile lokunciphisa uxinzelelo lobushushu obudibeneyo. Nangona kunjalo, i-cytokinin kunye ne-brassinosteroids zenze ngcono ukulungiswa kwezityalo njengoko zombini izilawuli zokukhula kwezityalo zandisa umxholo we-chlorophyll, i-alpha-chlorophyll fluorescence parameters, i-gs kunye ne-RWC xa kuthelekiswa nezityalo zerayisi ngaphandle kwesicelo, kunye nokunciphisa umxholo we-MDA kunye nobushushu be-canopy. Isishwankathelo, siphetha ukuba ukusetyenziswa kwezilawuli zokukhula kwezityalo (i-cytokinins kunye ne-brassinosteroids) sisixhobo esiluncedo ekulawuleni iimeko zoxinzelelo kwizityalo zerayisi ezibangelwa uxinzelelo olunzulu lobushushu ngexesha lobushushu obuphezulu.
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Ixesha lokuposa: Aug-08-2024