Abalawuli bokukhula kwezityalo basetyenziswe njengecebo lokunciphisa uxinzelelo lobushushu kwizityalo ezahlukeneyo

Imveliso yerayisi iyehla ngenxa yotshintsho lwemozulu kunye nokuguquguquka eColombia.Abalawuli bokukhula kwezityaloziye zasetyenziswa njengecebo lokunciphisa uxinzelelo lobushushu kwizityalo ezahlukeneyo. Ke ngoko, injongo yolu phononongo yayikukuvavanya iziphumo zomzimba (ukuqhuba kwesisu, ukuqhuba kwesisu, umxholo we-chlorophyll iyonke, umlinganiselo we-Fv/Fm weentlobo ezimbini zerayisi zorhwebo eziphantsi koxinzelelo lobushushu oludibeneyo (ubushushu obuphezulu basemini nasebusuku), ubushushu bekhenophi kunye nomxholo wamanzi ohambelanayo) kunye neenguqu zebhayoloji (i-malondialdehyde (MDA) kunye nomxholo we-prolinic acid). Uvavanyo lokuqala nolwesibini lwenziwe kusetyenziswa izityalo zeentlobo ezimbini zerayisi iFederrose 67 (“F67”) kunye neFederrose 2000 (“F2000”), ngokwahlukeneyo. Zombini iimvavanyo zahlalutywa kunye njengoluhlu lwezilingo. Unyango olumiselweyo lwalulandelayo: ulawulo olupheleleyo (AC) (izityalo zerayisi ezikhuliswe kumaqondo obushushu afanelekileyo (ubushushu basemini/ebusuku 30/25°C)), ulawulo loxinzelelo lobushushu (SC) [izityalo zerayisi ziphantsi koxinzelelo lobushushu oludibeneyo kuphela (40/ 25°C). 30°C)], kwaye izityalo zerayisi zaxinzelelwa zaza zatshizwa ngabalawuli bokukhula kwezityalo (uxinzelelo+AUX, uxinzelelo+BR, uxinzelelo+CK okanye uxinzelelo+GA) kabini (iintsuku ezi-5 ngaphambi nangeentsuku ezi-5 emva koxinzelelo lobushushu). Ukutshiza nge-SA kwandisa umxholo we-chlorophyll iyonke kuzo zombini iintlobo (ubunzima obutsha bezityalo zerayisi “F67″ kunye ne-“F2000″” yayiyi-3.25 kunye ne-3.65 mg/g, ngokulandelelana) xa kuthelekiswa nezityalo ze-SC (ubunzima obutsha bezityalo “F67″ babuyi-2.36 kunye ne-2.56 mg). g-1)” kunye nerayisi “F2000″, ukusetyenziswa kwamagqabi e-CK nako kuphucule ngokubanzi ukuhamba kwesisu kwezityalo zerayisi “F2000″ (499.25 vs. 150.60 mmol m-2 s) xa kuthelekiswa nolawulo loxinzelelo lobushushu. uxinzelelo lobushushu, ubushushu besithsaba sesityalo buyancipha nge-2–3 °C, kwaye umxholo we-MDA kwizityalo uyancipha. Isalathisi sokunyamezelana okuqhelekileyo sibonisa ukuba ukusetyenziswa kwe-CK (97.69%) kunye ne-BR (60.73%) kunokunceda ukunciphisa ingxaki yoxinzelelo lobushushu obudibeneyo. ikakhulu kwizityalo zerayisi ze-F2000. Ukuqukumbela, ukutshiza i-BR okanye i-CK ngamagqabi kunokuthathwa njengecebo lezolimo lokunceda ukunciphisa iziphumo ezimbi zeemeko zoxinzelelo lobushushu obudibeneyo kwindlela yokuziphatha kwezityalo zerayisi.
Irayisi (Oryza sativa) yeyosapho lwePoaceae kwaye yenye yezona nkozo zilinywa kakhulu emhlabeni kunye nombona nengqolowa (Bajaj kunye noMohanty, 2005). Indawo ephantsi kokulima irayisi yi-617,934 yeehektare, kwaye imveliso yesizwe ngo-2020 yayiyi-2,937,840 yeetoni kunye nesivuno esiphakathi seetoni eziyi-5.02/ha (Federarroz (Federación Nacional de Arroceros), 2021).
Ukufudumala kwehlabathi kuchaphazela izityalo zerayisi, okukhokelela kwiintlobo ezahlukeneyo zoxinzelelo lwe-abiotic ezifana namaqondo obushushu aphezulu kunye namaxesha embalela. Utshintsho lwemozulu lubangela ukuba amaqondo obushushu ehlabathi anyuke; Amaqondo obushushu aqikelelwa ukuba aza kunyuka nge-1.0–3.7°C kwinkulungwane yama-21, nto leyo enokunyusa ukuphindaphindeka kunye nobukhali boxinzelelo lobushushu. Ukunyuka kwamaqondo obushushu okusingqongileyo kuchaphazele irayisi, okubangela ukuba isivuno sezityalo sehle nge-6–7%. Kwelinye icala, utshintsho lwemozulu lukwakhokelela kwiimeko ezingathandekiyo zendalo kwizityalo, ezinje ngamaxesha embalela enzima okanye amaqondo obushushu aphezulu kwimimandla yetropiki neyasetropiki. Ukongeza, iziganeko ezahlukeneyo ezifana ne-El Niño zinokukhokelela kuxinzelelo lobushushu kwaye zibangele umonakalo omkhulu kwizityalo kwezinye iindawo zetropiki. EColombia, amaqondo obushushu kwiindawo ezivelisa irayisi aqikelelwa ukuba aza kunyuka nge-2–2.5°C ngo-2050, ehlisa imveliso yerayisi kwaye ichaphazele ukuhamba kwemveliso kwiimarike nakwiikhonkco zokubonelela.
Uninzi lwezityalo zerayisi zikhuliswa kwiindawo apho amaqondo obushushu asondelelene nomgangatho ofanelekileyo wokukhula kwesityalo (Shah et al., 2011). Kuye kwaxelwa ukuba amaqondo obushushu aphakathi emini nasebusuku afanelekileyoukukhula nophuhliso lwerayisingokubanzi zingama-28°C kunye nama-22°C, ngokulandelelana (Kilasi et al., 2018; Calderón-Páez et al., 2021). Amaqondo obushushu angaphezulu kwala manqanaba anokubangela amaxesha oxinzelelo lobushushu oluphakathi ukuya kolunzima ngexesha lamanqanaba abuthathaka ophuhliso lwerayisi (ukulinywa, ukuhluma, ukudubula, kunye nokuzaliswa kweenkozo), ngaloo ndlela kuchaphazela kakubi isivuno seenkozo. Oku kuncipha kwesivuno kungenxa kakhulu yexesha elide loxinzelelo lobushushu, oluchaphazela i-physiology yezityalo. Ngenxa yokusebenzisana kwezinto ezahlukeneyo, njengobude bexesha loxinzelelo kunye nobushushu obuphezulu obufikelelweyo, uxinzelelo lobushushu lunokubangela umonakalo omkhulu ongenakuphikiswa kwi-metabolism yezityalo kunye nophuhliso.
Uxinzelelo lobushushu luchaphazela iinkqubo ezahlukeneyo ze-physiological kunye ne-biochemical kwizityalo. I-photosynthesis yamagqabi yenye yeenkqubo ezichaphazeleka kakhulu kuxinzelelo lobushushu kwizityalo zerayisi, njengoko izinga le-photosynthesis lincipha ngama-50% xa amaqondo obushushu emihla ngemihla edlula ama-35°C. Iimpendulo ze-physiological zezityalo zerayisi ziyahluka ngokuxhomekeke kuhlobo loxinzelelo lobushushu. Umzekelo, amazinga e-photosynthetic kunye nokuhanjiswa kwe-stomatal kuyathintelwa xa izityalo zichaphazeleka kumaqondo obushushu aphezulu emini (33–40°C) okanye amaqondo obushushu aphezulu emini nasebusuku (35–40°C emini, 28–30°C). I-C ithetha ubusuku) (Lü et al., 2013; Fahad et al., 2016; Chaturvedi et al., 2017). Amaqondo obushushu aphezulu ebusuku (30°C) abangela uthintelo oluphakathi lwe-photosynthesis kodwa andisa ukuphefumla ebusuku (Fahad et al., 2016; Alvarado-Sanabria et al., 2017). Nokuba lixesha loxinzelelo, uxinzelelo lobushushu luchaphazela nomxholo we-chlorophyll yamagqabi, umlinganiselo we-chlorophyll variable fluorescence ukuya kwi-fluorescence ephezulu ye-chlorophyll (Fv/Fm), kunye nokusebenza kweRubisco kwizityalo zerayisi (Cao et al. 2009; Yin et al. 2010). ) Sanchez Reynoso et al., 2014).
Utshintsho lwebhayoloji lolunye umba wokulungelana kwezityalo noxinzelelo lobushushu (Wahid et al., 2007). Umxholo weproline usetyenziswe njengesalathisi sebhayoloji soxinzelelo lwezityalo (Ahmed noHassan 2011). Iproline idlala indima ebalulekileyo kwimetabolism yezityalo njengoko isebenza njengomthombo wekhabhoni okanye initrogen kwaye njenge-membrane stabilizer phantsi kweemeko zobushushu obuphezulu (Sánchez-Reinoso et al., 2014). Ubushushu obuphezulu buchaphazela nokuzinza kwe-membrane nge-lipid peroxidation, nto leyo ekhokelela ekwakhiweni kwe-malondialdehyde (MDA) (Wahid et al., 2007). Ke ngoko, umxholo we-MDA usetyenzisiwe ukuqonda ukuthembeka kwesakhiwo se-membranes zeseli phantsi koxinzelelo lobushushu (Cao et al., 2009; Chavez-Arias et al., 2018). Ekugqibeleni, uxinzelelo lobushushu oludibeneyo [37/30°C (emini/ebusuku)] lonyuse ipesenti yokuvuza kwe-electrolyte kunye nomxholo we-malondialdehyde kwirayisi (Liu et al., 2013).
Ukusetyenziswa kwezixhobo zokulawula ukukhula kwezityalo (ii-GR) kuye kwavavanywa ukunciphisa iziphumo ezimbi zoxinzelelo lobushushu, njengoko ezi zinto zibandakanyeka ngokukhutheleyo kwiimpendulo zezityalo okanye kwiindlela zokuzikhusela zomzimba nxamnye noxinzelelo olunjalo (uPeleg noBlumald, 2011; uYin et al. et al., 2011; uAhmed et al., 2015). Ukusetyenziswa kwezixhobo zemfuza ngaphandle kwelizwe kube nefuthe elihle ekunyamezeleni uxinzelelo lobushushu kwizityalo ezahlukeneyo. Izifundo zibonise ukuba ii-phytohormones ezifana ne-gibberellins (GA), i-cytokinins (CK), i-auxins (AUX) okanye i-brassinosteroids (BR) zikhokelela ekwandeni kwezinto ezahlukeneyo zemfuza kunye ne-biochemical (uPeleg noBlumald, 2011; uYin et al. Ren, 2011; uMitler et al., 2012; uZhou et al., 2014). EColombia, ukusetyenziswa kwezixhobo zemfuza ngaphandle kwelizwe kunye nefuthe lazo kwizityalo zerayisi akukaqondwa ngokupheleleyo kwaye kufundwe. Nangona kunjalo, uphando lwangaphambili lubonise ukuba ukutshiza i-BR ngamahlamvu kunokuphucula ukunyamezela irayisi ngokuphucula iimpawu zokutshintshiselana kwegesi, i-chlorophyll okanye umxholo we-proline wamagqabi eembewu zerayisi (Quintero-Calderón et al., 2021).
IiCytokinins zilawula iimpendulo zezityalo kwiingcinezelo ze-abiotic, kuquka uxinzelelo lobushushu (Ha et al., 2012). Ukongeza, kuye kwaxelwa ukuba ukusetyenziswa kwe-CK ngaphandle kunokunciphisa umonakalo wobushushu. Umzekelo, ukusetyenziswa kwe-zeatin ngaphandle kwandisa izinga le-photosynthetic, umxholo we-chlorophyll a kunye no-b, kunye nokusebenza kakuhle kokuthutha kwe-electron kwi-creeping bentgrass (Agrotis estolonifera) ngexesha loxinzelelo lobushushu (Xu kunye noHuang, 2009; Jespersen kunye noHuang, 2015). Ukusetyenziswa kwe-zeatin ngaphandle kunokuphucula umsebenzi we-antioxidant, kuphucule ukwenziwa kweeproteni ezahlukeneyo, kunciphise umonakalo we-reactive oxygen species (ROS) kunye nemveliso ye-malondialdehyde (MDA) kwizicubu zezityalo (Chernyadyev, 2009; Yang et al., 2009). , 2016; Kumar et al., 2020).
Ukusetyenziswa kwe-gibberellic acid kukwabonakalise impendulo entle kuxinzelelo lobushushu. Izifundo zibonise ukuba i-GA biosynthesis ilawula iindlela ezahlukeneyo ze-metabolic kwaye yonyusa ukunyamezela phantsi kweemeko zobushushu obuphezulu (Alonso-Ramirez et al. 2009; Khan et al. 2020). U-Abdel-Nabi et al. (2020) bafumanise ukuba ukutshiza i-GA yangaphandle (25 okanye 50 mg*L) ngefoliar kunokunyusa izinga le-photosynthetic kunye nomsebenzi we-antioxidant kwizityalo ze-orenji ezixinezelekileyo ngobushushu xa kuthelekiswa nezityalo ezilawulayo. Kukwabonwe ukuba ukusetyenziswa kwe-HA ngaphandle kwandisa umxholo womswakama, i-chlorophyll kunye nomxholo we-carotenoid kwaye kunciphisa i-lipid peroxidation kwi-date palm (Phoenix dactylifera) phantsi koxinzelelo lobushushu (Khan et al., 2020). I-Auxin ikwadlala indima ebalulekileyo ekulawuleni iimpendulo zokukhula ezihambelanayo kwiimeko zobushushu obuphezulu (Sun et al., 2012; Wang et al., 2016). Lo mlawuli wokukhula usebenza njengophawu lwe-biochemical kwiinkqubo ezahlukeneyo ezifana nokwenziwa kwe-proline okanye ukubola phantsi koxinzelelo lwe-abiotic (Ali et al. 2007). Ukongeza, i-AUX ikwaphucula umsebenzi we-antioxidant, okhokelela ekunciphiseni i-MDA kwizityalo ngenxa yokuncipha kwe-lipid peroxidation (Bielach et al., 2017). USergeev et al. (2018) baqaphele ukuba kwizityalo ze-pea (Pisum sativum) phantsi koxinzelelo lobushushu, umxholo we-proline - dimethylaminoethoxycarbonylmethyl)naphthylchloromethyl ether (TA-14) uyanda. Kuvavanyo olufanayo, baqaphele amanqanaba aphantsi e-MDA kwizityalo ezinyangiweyo xa kuthelekiswa nezityalo ezinganyangwanga nge-AUX.
IiBrassinosteroids lolunye udidi lwabalawuli bokukhula abasetyenziselwa ukunciphisa imiphumo yoxinzelelo lobushushu. U-Ogweno et al. (2008) baxele ukuba i-BR spray engaphandle yonyusa izinga le-photosynthetic, i-stomatal conductance kunye nezinga eliphezulu le-Rubisco carboxylation yezityalo zetamatato (Solanum lycopersicum) phantsi koxinzelelo lobushushu kangangeentsuku ezi-8. Ukutshiza nge-epibrassinosteroids ngefoliar kunokunyusa izinga le-photosynthetic lezityalo zekhukhamba (Cucumis sativus) phantsi koxinzelelo lobushushu (Yu et al., 2004). Ukongeza, ukusetyenziswa kwe-BR ngaphandle kuyalibazisa ukubola kwe-chlorophyll kwaye kwandisa ukusebenza kakuhle kokusetyenziswa kwamanzi kunye ne-quantum yield ye-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). Ukongeza, ukusetyenziswa kwezinto eziphilayo kunye nezinto eziphilayo (ubushushu bamagqabi, ukuhanjiswa kwesisu, iiparameter ze-chlorophyll fluorescence, i-chlorophyll kunye nomxholo wamanzi ohambelanayo, i-malondialdehyde kunye nokuhlanganiswa kwe-proline) sisixhobo esithembekileyo sokuhlola izityalo zerayisi phantsi koxinzelelo lobushushu ekuhlaleni nakwamanye amazwe (Sánchez -Reynoso et al., 2014; Alvarado-Sanabria et al., 2017; Nangona kunjalo, uphando malunga nokusetyenziswa kwe-foliar phytohormonal sprays kwirayisi kwinqanaba lendawo luhlala lunqabile. Ke ngoko, uphando lwempendulo yomzimba kunye neye-biochemical yokusetyenziswa kwabalawuli bokukhula kwezityalo lubaluleke kakhulu kwisiphakamiso sezicwangciso zezolimo ezisebenzayo kule nto. ukujongana nemiphumo emibi yexesha loxinzelelo lobushushu oluyinkimbinkimbi kwirayisi. Ke ngoko, injongo yolu phononongo yayikukuvavanya i-physiological (ukuhanjiswa kwesisu, iiparameter ze-chlorophyll fluorescence kunye nomxholo wamanzi ohambelanayo) kunye nemiphumo ye-biochemical yokusetyenziswa kwamagqabi kwabalawuli bokukhula kwezityalo abane (AUX, CK, GA kunye ne-BR). (Iipigmenti ze-Photosynthetic, i-malondialdehyde kunye nomxholo we-proline) Iinguqu kwiintlobo ezimbini zerayisi zorhwebo ezijongene noxinzelelo lobushushu oludibeneyo (amaqondo obushushu aphezulu emini/ebusuku).
Kolu phononongo, kwenziwe iimvavanyo ezimbini ezizimeleyo. Ii-genotypes iFederrose 67 (F67: i-genotype ephuhliswe kumaqondo obushushu aphezulu kwiminyaka elishumi edlulileyo) kunye neFederrose 2000 (F2000: i-genotype ephuhliswe kwiminyaka elishumi yokugqibela yenkulungwane yama-20 ebonisa ukumelana nentsholongwane yamagqabi amhlophe) zasetyenziswa okokuqala. imbewu. kunye novavanyo lwesibini, ngokulandelelana. Zombini ii-genotypes zityalwa kakhulu ngabalimi baseColombia. Imbewu ityalwe kwiitreyi ezili-10-L (ubude buyi-39.6 cm, ububanzi buyi-28.8 cm, ukuphakama buyi-16.8 cm) equlethe umhlaba wesanti onomhlwa one-2% yezinto eziphilayo. Imbewu ezintlanu ezisandula ukuhluma zatyalwa kwitreyi nganye. Iipalethi zabekwa kwindawo eluhlaza yeFakhalthi yeSayensi yezoLimo kwiYunivesithi yeSizwe yaseColombia, ikhampasi yaseBogotá (43°50′56″ N, 74°04′051″ W), kubude obuyi-2556 m ngaphezu kolwandle (asl). m.) kwaye zenziwe ukusuka ngo-Okthobha ukuya kuDisemba ka-2019. Uvavanyo olunye (iFederroz 67) kunye novavanyo lwesibini (iFederroz 2000) kwixesha elifanayo lonyaka ka-2020.
Iimeko zokusingqongileyo kwigreenhouse ngexesha ngalinye lokutyala zezi zilandelayo: ubushushu basemini nasebusuku yi-30/25°C, umswakama ohambelanayo yi-60~80%, ixesha le-photoperiod yendalo iiyure ezili-12 (imitha esebenzayo ye-photosynthetically 1500 µmol (photons) m-2 s-). 1 emini). Izityalo zatyandwa ngokomxholo wento nganye kwiintsuku ezingama-20 emva kokuvela kwembewu (DAE), ngokutsho kukaSánchez-Reinoso et al. (2019): i-670 mg ye-nitrogen ngesityalo ngasinye, i-110 mg ye-phosphorus ngesityalo ngasinye, i-350 mg ye-potassium ngesityalo ngasinye, i-68 mg ye-calcium ngesityalo ngasinye, i-20 mg ye-magnesium ngesityalo ngasinye, i-20 mg ye-sulfur ngesityalo ngasinye, i-17 mg ye-silicon ngesityalo ngasinye. Izityalo ziqulathe i-10 mg ye-boron ngesityalo ngasinye, i-17 mg ye-copper ngesityalo ngasinye, kunye ne-44 mg ye-zinc ngesityalo ngasinye. Izityalo zerayisi zigcinwe ukuya kuthi ga kwi-47 DAE kuvavanyo ngalunye xa zifikelela kwinqanaba le-phenological V5 ngeli xesha. Izifundo zangaphambili zibonise ukuba eli nqanaba le-phenological lixesha elifanelekileyo lokwenza izifundo zoxinzelelo lobushushu kwirayisi (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2017).
Kwilinge ngalinye, kwenziwe iindlela ezimbini ezahlukeneyo zokulawula ukukhula kwamagqabi. Iseti yokuqala ye-phytohormone sprays yafakwa kwiintsuku ezi-5 ngaphambi konyango loxinzelelo lobushushu (42 DAE) ukulungiselela izityalo uxinzelelo lokusingqongileyo. I-spray yesibini yanikwa emva kweentsuku ezi-5 emva kokuba izityalo zichatshazelwe kwiimeko zoxinzelelo (52 DAE). Kusetyenziswe ii-phytohormones ezine kwaye iimpawu zesithako ngasinye esisebenzayo esitshiziweyo kolu phononongo zidweliswe kwiTheyibhile eyoNgezelelweyo 1. Uxinzelelo lwe-Auxin (1-naphthylacetic acid: NAA) kuxinzelelo lwe-5 × 10−5 M (ii) 5 × 10–5 M gibberellin (gibberellic acid: NAA); GA3); (iii) I-Cytokinin (i-trans-zeatin) 1 × 10-5 M (iv) IiBrassinosteroids [Spirostan-6-one, 3,5-dihydroxy-, (3b,5a,25R)] 5 × 10-5; M. La manqanaba akhethwe kuba abangela iimpendulo ezilungileyo kwaye andisa ukumelana kwezityalo noxinzelelo lobushushu (Zahir et al., 2001; Wen et al., 2010; El-Bassiony et al., 2012; Salehifar et al., 2017). Izityalo zerayisi ezingenazo naziphi na izitshizi zokulawula ukukhula kwezityalo ziphathwe ngamanzi acocekileyo kuphela. Zonke izityalo zerayisi zitshizwe ngesitshizi sesandla. Faka i-20 ml H2O kwisityalo ukuze umanzise umphezulu ophezulu nongaphantsi kwamagqabi. Zonke izitshizi zefoliar zisebenzise i-adjuvant yezolimo (Agrotin, Bayer CropScience, Colombia) kwi-0.1% (v/v). Umgama phakathi kwembiza nesitshizi yi-30 cm.
Unyango loxinzelelo lobushushu lwenziwa kwiintsuku ezi-5 emva kokutshizwa kwefoliar yokuqala (47 DAE) kuvavanyo ngalunye. Izityalo zerayisi zidluliselwe ukusuka kwindawo yokukhulisa izityalo ukuya kwigumbi lokukhula elingama-294 L (MLR-351H, Sanyo, IL, USA) ukuze kusekwe uxinzelelo lobushushu okanye kugcinwe iimeko ezifanayo zokusingqongileyo (47 DAE). Unyango loxinzelelo lobushushu oludibeneyo lwenziwe ngokubeka igumbi kubushushu obulandelayo basemini/ebusuku: ubushushu obuphezulu basemini [40°C iiyure ezi-5 (ukusuka ngo-11:00 ukuya ku-16:00)] kunye nexesha lasebusuku [30°C iiyure ezi-5]. Iintsuku ezi-8 zilandelelana (ukusuka ngo-19:00 ukuya ku-24:00). Ubushushu boxinzelelo kunye nexesha lokuvezwa zikhethwe ngokusekelwe kwizifundo zangaphambili (Sánchez-Reynoso et al. 2014; Alvarado-Sanabía et al. 2017). Kwelinye icala, iqela lezityalo ezifuduselwe kwigumbi lokukhula lagcinwa kwigreenhouse kubushushu obufanayo (30°C emini/25°C ebusuku) kangangeentsuku ezisi-8 ezilandelelanayo.
Ekupheleni kovavanyo, kufunyenwe la maqela onyango alandelayo: (i) imeko yobushushu bokukhula + ukusetyenziswa kwamanzi acocekileyo [ulawulo olupheleleyo (AC)], (ii) imeko yoxinzelelo lobushushu + ukusetyenziswa kwamanzi acocekileyo [ulawulo loxinzelelo lobushushu (SC)], (iii) imeko yoxinzelelo lobushushu + ukusetyenziswa kwe-auxin (AUX), (iv) imeko yoxinzelelo lobushushu + ukusetyenziswa kwe-gibberellin (GA), (v) imeko yoxinzelelo lobushushu + ukusetyenziswa kwe-cytokinin (CK), kunye (vi) imeko yoxinzelelo lobushushu + i-brassinosteroid (BR) Isihlomelo. La maqela onyango asetyenziselwe ii-genotypes ezimbini (F67 kunye ne-F2000). Zonke iindlela zonyango zenziwe ngoyilo olucwangcisiweyo ngokupheleleyo oluneekopi ezintlanu, nganye inesityalo esinye. Isityalo ngasinye sasetyenziselwa ukufunda iinguqu ezichazwe ekupheleni kovavanyo. Uvavanyo luthathe i-55 DAE.
Ukuhanjiswa kwesisu (gs) kulinganiswe kusetyenziswa i-portable porosometer (SC-1, METER Group Inc., USA) ukusuka kwi-0 ukuya kwi-1000 mmol m-2 s-1, kunye nomgodi wegumbi lesampulu oyi-6.35 mm. Umlinganiselo uthathwa ngokuncamathisela i-stomameter probe kwigqabi elivuthiweyo kunye nehlumela eliphambili lesityalo elandisiweyo ngokupheleleyo. Kunyango ngalunye, ukufundwa kwe-gs kuthathwe kumagqabi amathathu esityalo ngasinye phakathi kwe-11:00 kunye ne-16:00 kwaye kulinganiswe ngokomyinge.
I-RWC yamiselwa ngokwendlela echazwe nguGhoulam et al. (2002). Iphepha elandisiweyo ngokupheleleyo elisetyenziselwa ukumisela i-g lasetyenziswa nokulinganisa i-RWC. Ubunzima obutsha (FW) bamiselwa kwangoko emva kokuvuna kusetyenziswa isikali sedijithali. Amagqabi emva koko afakwa kwisikhongozeli seplastiki esizaliswe ngamanzi aze ashiywe ebumnyameni kubushushu begumbi (22°C) iiyure ezingama-48. Emva koko linganisa kwisikali sedijithali uze ubhale phantsi ubunzima obubanzi (TW). Amagqabi avuvukileyo omiswa kwi-oven kwi-75°C iiyure ezingama-48 kwaye ubunzima bawo obomileyo (DW) babhalwa phantsi.
Umxholo we-chlorophyll ohambelanayo uchongiwe kusetyenziswa imitha ye-chlorophyll (atLeafmeter, FT Green LLC, USA) kwaye uchazwe kwiiyunithi ze-atLeaf (Dey et al., 2016). Ukufundwa kwe-PSII maximum quantum efficiency (Fv/Fm ratio) kubhalwe kusetyenziswa i-continuous excitation chlorophyll fluorimeter (Handy PEA, Hansatech Instruments, UK). Amagqabi ahlengahlengiswa ngobumnyama kusetyenziswa ii-leaf clamps imizuzu engama-20 ngaphambi kokulinganiswa kwe-Fv/Fm (Restrepo-Diaz kunye noGarces-Varon, 2013). Emva kokuba amagqabi eqhelene nobumnyama, i-baseline (F0) kunye ne-maximum fluorescence (Fm) zilinganisiwe. Ukusuka kwezi datha, i-variable fluorescence (Fv = Fm – F0), umlinganiselo we-variable fluorescence ukuya kwi-maximum fluorescence (Fv/Fm), i-maximum quantum yield ye-PSII photochemistry (Fv/F0) kunye nomlinganiselo we-Fm/F0 zibalwe (Baker, 2008; Lee et al. ., 2017). Ukufundwa kwe-chlorophyll kunye ne-chlorophyll fluorescence kuthathwe kumagqabi afanayo asetyenziselwa ukulinganisa i-gs.
Malunga ne-800 mg yobunzima bamagqabi amatsha yaqokelelwa njengezinto eziguquguqukayo zebhayoloji. Iisampulu zamagqabi zaze zafakwa kwi-nitrogen engamanzi zaza zagcinwa ukuze kuhlalutywe ngakumbi. Indlela ye-spectrometric esetyenziselwa ukuqikelela umxholo we-chlorophyll a, b kunye ne-carotenoid yezicubu isekelwe kwindlela kunye nezibalo ezichazwe nguWellburn (1994). Iisampulu zamagqabi (30 mg) zaqokelelwa zaza zafakwa kwi-3 ml ye-80% ye-acetone. Iisampulu zaze zafakwa kwi-centrifuge (model 420101, Becton Dickinson Primary Care Diagnostics, USA) kwi-5000 rpm kangangemizuzu eli-10 ukususa amasuntswana. I-supernatant yancitshiswa yaya kumthamo wokugqibela we-6 ml ngokongeza i-80% ye-acetone (Sims and Gamon, 2002). Umxholo we-chlorophyll ufunyenwe kwi-663 (chlorophyll a) kunye ne-646 (chlorophyll b) nm, kunye nee-carotenoids kwi-470 nm kusetyenziswa i-spectrophotometer (Spectronic BioMate 3 UV-vis, Thermo, USA).
Indlela ye-thiobarbituric acid (TBA) echazwe nguHodges et al. (1999) yasetyenziswa ukuvavanya i-membrane lipid peroxidation (MDA). Malunga ne-0.3 g yezicubu zamagqabi nayo yafakwa kwi-homogenized kwi-nitrogen engamanzi. Iisampuli zafakwa kwi-centrifuge kwi-5000 rpm kwaye ukufunxwa kwalinganiswa kwi-spectrophotometer kwi-440, 532 kunye ne-600 nm. Ekugqibeleni, uxinzelelo lwe-MDA lwabalwa kusetyenziswa i-extinction coefficient (157 M mL−1).
Umxholo weproline kuzo zonke iindlela zonyango uchongiwe kusetyenziswa indlela echazwe nguBates et al. (1973). Yongeza i-10 ml yesisombululo samanzi se-3% se-sulfosalicylic acid kwisampuli egciniweyo kwaye ucofe iphepha lesihluzo seWhatman (No. 2). Emva koko i-2 ml yesi sihluzo ixutywe ne-2 ml ye-ninhydric acid kunye ne-2 ml ye-glacial acetic acid. Umxube ubekwe kwindawo yokuhlambela ngamanzi kwi-90°C iyure e-1. Misa impendulo ngokuyifunxa emkhenkceni. Shukumisa ityhubhu ngamandla usebenzisa i-vortex shaker kwaye unyibilikise isisombululo esiphumayo kwi-4 ml ye-toluene. Ukufundwa kokufunxwa kuchongiwe kwi-520 nm kusetyenziswa i-spectrophotometer efanayo esetyenziselwa ukulinganisa i-pigment ye-photosynthetic (Spectronic BioMate 3 UV-Vis, Thermo, Madison, WI, USA).
Indlela echazwe nguGerhards et al. (2016) yokubala ubushushu bekhanopu kunye neCSI. Iifoto ezishushu zithathwe ngekhamera yeFLIR 2 (FLIR Systems Inc., eBoston, MA, e-USA) ngokuchanekileyo kwe-±2°C ekupheleni kwexesha loxinzelelo. Beka umphezulu omhlophe emva kwesityalo ukuze kuthathwe imifanekiso. Kwakhona, iifektri ezimbini zathathwa njengeemodeli zokubhekisa. Izityalo zabekwa kumphezulu omhlophe; enye yayigqunywe nge-adjuvant yezolimo (Agrotin, Bayer CropScience, Bogotá, eColombia) ukulinganisa ukuvulwa kwazo zonke iistomata [imo emanzi (Twet)], kwaye enye yayiyigqabi elingenalo naluphi na uhlobo lokusetyenziswa [imo eyomileyo (Tdry)] (Castro -Duque et al., 2020). Umgama phakathi kwekhamera kunye nembiza ngexesha lokudubula yayiyi-1 m.
Isalathisi sokunyamezelana okunxulumeneyo sibalwe ngokungathanga ngqo kusetyenziswa i-stomatal conductance (gs) yezityalo ezinyangiweyo xa kuthelekiswa nezityalo zolawulo (izityalo ezingenazo iindlela zonyango loxinzelelo kunye nabalawuli bokukhula abasetyenzisiweyo) ukumisela ukunyamezela kwe-genotypes ezinyangiweyo ezivavanyiweyo kolu phononongo. I-RTI ifunyenwe kusetyenziswa i-equation eguqulwe kwiChávez-Arias et al. (2020).
Kuvavanyo ngalunye, zonke izinto eziguquguqukayo zomzimba ezikhankanyiweyo apha ngasentla zichongiwe kwaye zarekhodwa kwi-55 DAE kusetyenziswa amagqabi akhuliswe ngokupheleleyo aqokelelwe kwikhabhathi ephezulu. Ukongeza, izilinganiso zenziwe kwigumbi lokukhula ukuze kuthintelwe ukutshintsha iimeko zendalo apho izityalo zikhula khona.
Idatha evela kuvavanyo lokuqala nolwesibini yahlalutywa kunye njengoluhlu lwezilingo. Iqela ngalinye lovavanyo lalinazo izityalo ezi-5, kwaye isityalo ngasinye sakha iyunithi yovavanyo. Uhlalutyo lokwahluka (ANOVA) lwenziwa (P ≤ 0.05). Xa kwafunyanwa umahluko obalulekileyo, uvavanyo lokuthelekisa lwe-post hoc lukaTukey lwasetyenziswa kwi-P ≤ 0.05. Sebenzisa umsebenzi we-arcsine ukuguqula amaxabiso epesenti. Idatha yahlalutywa kusetyenziswa isoftware ye-Statistix v 9.0 (i-Analytical Software, eTallahassee, FL, e-USA) yaza yadweliswa kusetyenziswa i-SigmaPlot (inguqulelo 10.0; iSystat Software, eSan Jose, CA, e-USA). Uhlalutyo oluphambili lwenziwe kusetyenziswa isoftware ye-InfoStat 2016 (i-Analysis Software, kwiYunivesithi yeSizwe yaseCordoba, eArgentina) ukuchonga abalawuli bokukhula kwezityalo ababalaseleyo abaphantsi kophando.
Itheyibhile 1 ishwankathela i-ANOVA ebonisa iimvavanyo, unyango olwahlukeneyo, kunye nokusebenzisana kwazo neepigment ze-photosynthetic zamagqabi (i-chlorophyll a, b, iyonke, kunye nee-carotenoids), i-malondialdehyde (MDA) kunye nomxholo we-proline, kunye nokuhanjiswa kwe-stomatal. Isiphumo se-gs, umxholo wamanzi ohambelanayo. (RWC), umxholo we-chlorophyll, iiparameter ze-chlorophyll alpha fluorescence, ubushushu be-crown (PCT) (°C), isalathisi soxinzelelo lwesivuno (CSI) kunye nesalathisi sokunyamezelana okuhambelanayo kwezityalo zerayisi kwi-55 DAE.
Itheyibhile 1. Isishwankathelo sedatha ye-ANOVA kwiinguqu ze-rice physiological kunye ne-biochemical phakathi kovavanyo (ii-genotypes) kunye nonyango loxinzelelo lobushushu.
Umahluko (P≤0.01) kwiindlela zokudibanisa imibala ye-photosynthetic leaf, umxholo we-chlorophyll ohambelanayo (ukufundwa kwe-Atleaf), kunye neeparameter ze-alpha-chlorophyll fluorescence phakathi kovavanyo kunye nonyango ziboniswe kwiTheyibhile 2. Amaqondo obushushu aphezulu emini nasebusuku anyuse umxholo we-chlorophyll kunye ne-carotenoid iyonke. Izithole zerayisi ezingenayo nayiphi na i-spray ye-phytohormones yamagqabi (2.36 mg g-1 ye-"F67″ kunye ne-2.56 mg g-1 ye-"F2000″) xa kuthelekiswa nezityalo ezikhuliswe phantsi kweemeko zobushushu ezifanelekileyo (2.67 mg g -1)) zibonise umxholo we-chlorophyll ophantsi. Kuzo zombini iimvavanyo, i-"F67" yayiyi-2.80 mg g-1 kwaye i-"F2000" yayiyi-2.80 mg g-1. Ukongeza, izithole zerayisi eziphathwe ngokudibanisa izitshizi ze-AUX kunye ne-GA phantsi koxinzelelo lobushushu nazo zibonise ukwehla komxholo we-chlorophyll kuzo zombini ii-genotypes (AUX = 1.96 mg g-1 kunye ne-GA = 1.45 mg g-1 kwi-"F67"; AUX = 1.96 mg g-1 kunye ne-GA = 1.45 mg g-1 kwi-"F67″; AUX = 2.24 mg) g-1 kunye ne-GA = 1.43 mg g-1 (kwi-"F2000″) phantsi kweemeko zoxinzelelo lobushushu. Phantsi kweemeko zoxinzelelo lobushushu, unyango lwe-foliar nge-BR lubangele ukwanda okuncinci kolu tshintsho kuzo zombini ii-genotypes. Ekugqibeleni, i-CK foliar spray ibonise amaxabiso aphezulu e-photosynthetic pigment ​​phakathi kwazo zonke iindlela zonyango (AUX, GA, BR, SC kunye nonyango lwe-AC) kwii-genotypes F67 (3.24 mg g-1) kunye ne-F2000 (3.65 mg g-1). Umxholo we-chlorophyll (iyunithi ye-Atleaf) nawo wehlisiwe kukuxinzelelwa kobushushu okudibeneyo. Amaxabiso aphezulu abhalwe kwizityalo ezitshizwe nge-CC kuzo zombini ii-genotypes (41.66 kwi-"F67" kunye ne-49.30 kwi-"F2000"). Umlinganiselo we-Fv kunye ne-Fv/Fm ubonise umahluko omkhulu phakathi konyango kunye nee-cultivars (Itheyibhile 2). Lilonke, phakathi kwezi zinto ziguquguqukayo, i-cultivar F67 yayingachaphazeleki kakhulu kukuxinzelelwa kobushushu kune-cultivar F2000. Umlinganiselo we-Fv kunye ne-Fv/Fm wonzakele ngakumbi kuvavanyo lwesibini. Izithole ezixinzekileyo ze-'F2000′ ezingatshizwanga nge-phytohormones zazinexabiso eliphantsi le-Fv (2120.15) kunye nomlinganiselo we-Fv/Fm (0.59), kodwa ukutshiza nge-foliar nge-CK kuncede ukubuyisela la maxabiso (Fv: 2591, 89, umlinganiselo we-Fv/Fm: 0.73). , ifumana ukufundwa okufana noko kubhalwe kwizityalo ze-“F2000” ezikhuliswe phantsi kweemeko zobushushu ezifanelekileyo (Fv: 2955.35, umlinganiselo we-Fv/Fm: 0.73:0.72). Akukho mahluko abalulekileyo kwi-initial fluorescence (F0), maximum fluorescence (Fm), maximum photochemical quantum yield ye-PSII (Fv/F0) kunye nomlinganiselo we-Fm/F0. Ekugqibeleni, i-BR ibonise umkhwa ofanayo njengoko kubonwe kwi-CK (Fv 2545.06, Fv/Fm ratio 0.73).
Itheyibhile 2. Isiphumo soxinzelelo lobushushu oludibeneyo (40°/30°C imini/ubusuku) kwimibala ye-photosynthetic yamagqabi [i-chlorophyll iyonke (i-Chl Total), i-chlorophyll a (i-Chl a), i-chlorophyll b (i-Chl b) kunye ne-carotenoids isiphumo se-Cx+c] ], umxholo we-chlorophyll ohambelanayo (iyunithi ye-Atliff), iiparameter ze-chlorophyll fluorescence (i-fluorescence yokuqala (i-F0), i-fluorescence ephezulu (i-Fm), i-fluorescence eguquguqukayo (i-Fv), ukusebenza kakuhle kwe-PSII (i-Fv/Fm), i-photochemical maximum quantum yield ye-PSII (Fv/F0 ) kunye ne-Fm/F0 kwizityalo ze-genotypes ezimbini zerayisi [i-Federrose 67 (i-F67) kunye ne-Federrose 2000 (i-F2000)] kwiintsuku ezingama-55 emva kokuvela (i-DAE)).
Umxholo wamanzi (RWC) wezityalo zerayisi eziphathwe ngokwahlukileyo ubonise umahluko (P ≤ 0.05) kunxibelelwano phakathi konyango lovavanyo nolwe-foliar (Umzobo 1A). Xa kunyangwa nge-SA, amaxabiso aphantsi arekhodwe kuzo zombini ii-genotypes (74.01% kwi-F67 kunye ne-76.6% kwi-F2000). Phantsi kweemeko zoxinzelelo lobushushu, i-RWC yezityalo zerayisi zeentlobo zombini ze-genotypes eziphathwe ngee-phytohormones ezahlukeneyo yanda kakhulu. Ngokubanzi, ukusetyenziswa kwe-CK, GA, AUX, okanye i-BR kwizityalo ze-foliar kwandise i-RWC ukuya kumaxabiso afana nalawo ezityalo ezikhuliswe phantsi kweemeko ezifanelekileyo ngexesha lovavanyo. Ulawulo olupheleleyo kunye nezityalo ezitshiziweyo kwizityalo ze-foliar zirekhode amaxabiso amalunga ne-83% kuzo zombini ii-genotypes. Kwelinye icala, ii-gs nazo zibonise umahluko obalulekileyo (P ≤ 0.01) kunxibelelwano lovavanyo nonyango (Umzobo 1B). Isityalo esilawula ngokupheleleyo (AC) sikwabhale amaxabiso aphezulu kwi-genotype nganye (440.65 mmol m-2s-1 kwi-F67 kunye ne-511.02 mmol m-2s-1 kwi-F2000). Izityalo zerayisi eziphantsi koxinzelelo lobushushu oludibeneyo zodwa zibonise amaxabiso aphantsi kakhulu kwi-genotypes zombini (150.60 mmol m-2s-1 kwi-F67 kunye ne-171.32 mmol m-2s-1 kwi-F2000). Unyango lwe-foliar ngazo zonke i-plant regulators nalo lwanda g. Kwizityalo zerayisi ze-F2000 ezitshizwe nge-CC, isiphumo sokutshiza nge-foliar nge-phytohormones sasicacile ngakumbi. Eli qela lezityalo alibonisanga mahluko xa kuthelekiswa nezityalo ezilawula ngokupheleleyo (AC 511.02 kunye ne-CC 499.25 mmol m-2s-1).
Umfanekiso 1. Isiphumo soxinzelelo lobushushu oludibeneyo (40°/30°C imini/ubusuku) kumxholo wamanzi ohambelanayo (RWC) (A), ukuhanjiswa kwesisu (gs) (B), imveliso ye-malondialdehyde (MDA) (C), kunye nomxholo we-proline. (D) kwizityalo ze-genotypes ezimbini zerayisi (F67 kunye ne-F2000) kwiintsuku ezingama-55 emva kokuvela (DAE). Unyango oluvavanyiweyo kwi-genotype nganye luquka: ulawulo olupheleleyo (AC), ulawulo loxinzelelo lobushushu (SC), uxinzelelo lobushushu + auxin (AUX), uxinzelelo lobushushu + gibberellin (GA), uxinzelelo lobushushu + i-cell mitogen (CK), kunye noxinzelelo lobushushu + i-brassinosteroid. (BR). Ikholamu nganye imele impazamo ye-± standard yamanqaku edatha amahlanu (n = 5). Iikholamu ezilandelwa ngoonobumba abahlukeneyo zibonisa umahluko obalulekileyo ngokwezibalo ngokovavanyo lukaTukey (P ≤ 0.05). Oonobumba abanophawu olulinganayo babonisa ukuba i-mean ayibalulekanga ngokwezibalo (≤ 0.05).
Umxholo we-MDA (P ≤ 0.01) kunye ne-proline (P ≤ 0.01) nawo ubonise umahluko omkhulu kunxibelelwano phakathi kovavanyo kunye nonyango lwe-phytohormone (Umzobo 1C, D). Ukwanda kwe-lipid peroxidation kubonwe ngonyango lwe-SC kuzo zombini ii-genotypes (Umfanekiso 1C), nangona kunjalo izityalo ezinyangwe nge-leaf growth regulator spray zibonise ukwehla kwe-lipid peroxidation kuzo zombini ii-genotypes; Ngokubanzi, ukusetyenziswa kwe-phytohormones (CA, AUC, BR okanye GA) kukhokelela ekunciphiseni kwe-lipid peroxidation (umxholo we-MDA). Akukho mahluko afunyenweyo phakathi kwezityalo ze-AC zeentlobo ezimbini ze-genotypes kunye nezityalo eziphantsi koxinzelelo lobushushu kwaye zatshizwa nge-phytohormones (kubonwe amaxabiso e-FW kwizityalo ze-"F67" ukusuka kwi-4.38–6.77 µmol g-1, kwaye kwizityalo ze-FW "F2000" "kubonwe amaxabiso ukusuka kwi-2.84 ukuya kwi-9.18 µmol g-1 (izityalo). Kwelinye icala, ukwenziwa kwe-proline kwizityalo ze-"F67" kwakuphantsi kunezityalo ze-"F2000" eziphantsi koxinzelelo oludibeneyo, nto leyo ekhokelele ekwandeni kwemveliso ye-proline. kwizityalo zerayisi ezixinezelekileyo kubushushu, kuzo zombini iimvavanyo, kwabonwa ukuba ukulawulwa kwezi hormones kwandisa kakhulu umxholo we-amino acid kwizityalo ze-F2000 (i-AUX kunye ne-BR zaziyi-30.44 kunye ne-18.34 µmol g-1) ngokwahlukeneyo (Umzobo 1G).
Iziphumo ze-foliar plant growth regulator spray kunye noxinzelelo lobushushu oludibeneyo kubushushu be-canopy yezityalo kunye ne-relative tolerance index (RTI) ziboniswe kwiMifanekiso 2A kunye no-B. Kuzo zombini ii-genotypes, ubushushu be-canopy yezityalo ze-AC babusondele kwi-27°C, kwaye obo bezityalo ze-SC babumalunga ne-28°C. WITH. Kwakhona kwabonwa ukuba unyango lwe-foliar nge-CK kunye ne-BR lubangele ukwehla kwe-canopy nge-2–3°C xa kuthelekiswa nezityalo ze-SC (Umfanekiso 2A). I-RTI ibonakalise ukuziphatha okufanayo nezinye izinto eziphilayo, ibonisa umahluko obalulekileyo (P ≤ 0.01) kwintsebenziswano phakathi kovavanyo kunye nonyango (Umfanekiso 2B). Izityalo ze-SC zibonise ukunyamezela kwezityalo okuphantsi kuzo zombini ii-genotypes (34.18% kunye ne-33.52% kwizityalo zerayisi ze-"F67" kunye ne-"F2000", ngokulandelelana). Ukondla nge-phytohormones nge-foliar kuphucula i-RTI kwizityalo ezivezwe kuxinzelelo lobushushu obuphezulu. Esi siphumo sibonakale ngakumbi kwizityalo ze-“F2000” ezitshizwe nge-CC, apho i-RTI yayiyi-97.69. Kwelinye icala, umahluko omkhulu ubonwe kuphela kwi-index yoxinzelelo lwesivuno (CSI) sezityalo zerayisi phantsi kweemeko zoxinzelelo lwe-foliar factor (P ≤ 0.01) (Umzobo 2B). Zizityalo zerayisi kuphela eziphantsi koxinzelelo lobushushu oluntsonkothileyo ezibonise ixabiso eliphezulu le-index yoxinzelelo (0.816). Xa izityalo zerayisi zitshizwe ngee-phytohormones ezahlukeneyo, i-index yoxinzelelo yayiphantsi (amaxabiso ukusuka kwi-0.6 ukuya kwi-0.67). Ekugqibeleni, isityalo zerayisi esikhuliswe phantsi kweemeko ezifanelekileyo sasinexabiso le-0.138.
Umfanekiso 2. Iziphumo zoxinzelelo lobushushu oludibeneyo (40°/30°C emini/ebusuku) kubushushu bekhenophi (A), isalathisi sokunyamezelana okunxulumeneyo (RTI) (B), kunye nesalathisi soxinzelelo lwesivuno (CSI) (C) seentlobo ezimbini zezityalo. Ii-genotype zerayisi zorhwebo (F67 kunye ne-F2000) ziphantsi konyango lobushushu olwahlukeneyo. Unyango oluvavanyiweyo kwi-genotype nganye luquka: ulawulo olupheleleyo (AC), ulawulo loxinzelelo lobushushu (SC), uxinzelelo lobushushu + auxin (AUX), uxinzelelo lobushushu + gibberellin (GA), uxinzelelo lobushushu + cell mitogen (CK), kunye noxinzelelo lobushushu + brassinosteroid. (BR). Uxinzelelo lobushushu oludibeneyo lubandakanya ukuveza izityalo zerayisi kubushushu obuphezulu bemini/ebusuku (40°/30°C emini/ebusuku). Ikholamu nganye imele impazamo eqhelekileyo ± yamanqaku edatha amahlanu (n = 5). Iikholamu ezilandelwa ngoonobumba abahlukeneyo zibonisa umahluko obalulekileyo ngokwezibalo ngokovavanyo lukaTukey (P ≤ 0.05). Oonobumba abanophawu olulinganayo babonisa ukuba i-mean ayibalulekanga ngokwezibalo (≤ 0.05).
Uhlalutyo lwecandelo eliphambili (PCA) luveze ukuba iinguqu ezivavanyiweyo kwi-55 DAE zichaze i-66.1% yeempendulo ze-physiological kunye ne-biochemical zezityalo zerayisi ezixinezelekileyo ngobushushu eziphathwe nge-growth regulator spray (Umzobo 3). Ii-vectors zimele iinguqu kwaye amachaphaza amele abalawuli bokukhula kwezityalo (ii-GR). Ii-vectors ze-gs, umxholo we-chlorophyll, ukusebenza kakuhle kwe-quantum ye-PSII (Fv/Fm) kunye neeparameter ze-biochemical (TChl, MDA kunye ne-proline) zikufutshane nemvelaphi, nto leyo ebonisa ulwalamano oluphezulu phakathi kokuziphatha kwe-physiological kwezityalo kunye nazo. eziguquguqukayo. Iqela elinye (V) liquke izithole zerayisi ezikhuliswe kubushushu obuphezulu (AT) kunye nezityalo ze-F2000 eziphathwe nge-CK kunye ne-BA. Kwangaxeshanye, uninzi lwezityalo eziphathwe nge-GR zenze iqela elahlukileyo (IV), kwaye unyango nge-GA kwi-F2000 zenze iqela elahlukileyo (II). Ngokwahlukileyo koko, izithole zerayisi ezixinene nobushushu (amaqela oku-1 kunye nelesi-III) ezingenazo ii-phytohormones ezitshizwa ngamahlamvu (zombini ii-genotypes zaziyi-SC) zazikwindawo echasene neqela V, nto leyo ebonisa impembelelo yoxinzelelo lobushushu kwi-physiology yezityalo.
Umfanekiso 3. Uhlalutyo lwebhayografi yeziphumo zoxinzelelo lobushushu oludibeneyo (40°/30°C imini/ubusuku) kwizityalo zeentlobo ezimbini zerayisi (F67 kunye ne-F2000) kwiintsuku ezingama-55 emva kokuvela (DAE). Izifinyezo: AC F67, ulawulo olupheleleyo F67; SC F67, ulawulo loxinzelelo lobushushu F67; AUX F67, uxinzelelo lobushushu + auxin F67; GA F67, uxinzelelo lobushushu + gibberellin F67; CK F67, uxinzelelo lobushushu + ukwahlulwa kweeseli BR F67, uxinzelelo lobushushu + brassinosteroid. F67; AC F2000, ulawulo olupheleleyo F2000; SC F2000, ulawulo loxinzelelo lobushushu F2000; AUX F2000, uxinzelelo lobushushu + auxin F2000; GA F2000, uxinzelelo lobushushu + gibberellin F2000; I-CK F2000, uxinzelelo lobushushu + i-cytokinin, i-BR F2000, uxinzelelo lobushushu + i-steroid yobhedu; F2000.
Izinto eziguquguqukayo ezifana nomxholo we-chlorophyll, i-stomatal conductance, i-Fv/Fm ratio, i-CSI, i-MDA, i-RTI kunye nomxholo we-proline zinokunceda ukuqonda ukulungelelaniswa kwee-genotypes zerayisi kunye nokuvavanya impembelelo yezicwangciso zezolimo phantsi koxinzelelo lobushushu (Sarsu et al., 2018; Quintero-Calderon et al., 2021). Injongo yolu vavanyo yayikukuvavanya impembelelo yokusetyenziswa kwabalawuli bokukhula abane kwimilinganiselo ye-physiological kunye ne-biochemical yezityalo zerayisi phantsi kweemeko ezinzima zoxinzelelo lobushushu. Uvavanyo lwezithombo yindlela elula nekhawulezayo yokuvavanya ngaxeshanye izityalo zerayisi kuxhomekeke kubukhulu okanye imeko yeziseko ezikhoyo (Sarsu et al. 2018). Iziphumo zolu phononongo zibonise ukuba uxinzelelo lobushushu oludibeneyo lubangela iimpendulo ezahlukeneyo ze-physiological kunye ne-biochemical kwii-genotypes ezimbini zerayisi, nto leyo ebonisa inkqubo yokulungelelanisa. Ezi ziphumo zibonisa nokuba izitshizi zokulawula ukukhula kwamagqabi (ikakhulu ii-cytokinins kunye ne-brassinosteroids) zinceda irayisi ukuba ivumelane noxinzelelo lobushushu oluntsonkothileyo njengoko i-favor ichaphazela kakhulu i-gs, i-RWC, umlinganiselo we-Fv/Fm, iipigment ze-photosynthetic kunye nomxholo we-proline.
Ukusetyenziswa kwabalawuli bokukhula kunceda ukuphucula imeko yamanzi yezityalo zerayisi phantsi koxinzelelo lobushushu, nto leyo enokunxulunyaniswa noxinzelelo oluphezulu kunye nobushushu obuphantsi bezityalo. Olu phononongo lubonise ukuba phakathi kwezityalo "ze-F2000" (i-genotype enokuchaphazeleka), izityalo zerayisi ezinyangwe ikakhulu nge-CK okanye i-BR zazinamaxabiso aphezulu e-gs kunye namaxabiso aphantsi e-PCT kunezityalo ezinyangwe nge-SC. Izifundo zangaphambili zikwabonise ukuba ii-gs kunye ne-PCT zizikhombisi ezichanekileyo ze-physiological ezinokumisela impendulo eguquguqukayo yezityalo zerayisi kunye nemiphumo yezicwangciso zezolimo kuxinzelelo lobushushu (Restrepo-Diaz kunye noGarces-Varon, 2013; Sarsu et al., 2018; Quintero). -Carr DeLong et al., 2021). I-Leaf CK okanye i-BR yonyusa i-g phantsi koxinzelelo kuba ezi hormone zezityalo zinokukhuthaza ukuvulwa kwe-stomatal ngokusebenzisana okwenziweyo nezinye iimolekyuli zesignali ezifana ne-ABA (umkhuthazi wokuvalwa kwe-stomatal phantsi koxinzelelo lwe-abiotic) (Macková et al., 2013; Zhou et al., 2013). 2013). ). , 2014). Ukuvulwa kwesisu kukhuthaza ukupholisa amagqabi kwaye kunceda ukunciphisa amaqondo obushushu be-canopy (Sonjaroon et al., 2018; Quintero-Calderón et al., 2021). Ngenxa yezi zizathu, amaqondo obushushu be-canopy yezityalo zerayisi ezitshizwe nge-CK okanye i-BR anokuba phantsi xa kudityaniswe uxinzelelo lobushushu.
Uxinzelelo lobushushu obuphezulu lunokunciphisa umxholo we-pigment ye-photosynthetic yamagqabi (Chen et al., 2017; Ahammed et al., 2018). Kolu phononongo, xa izityalo zerayisi zaziphantsi koxinzelelo lobushushu kwaye zingatshizwanga ngabalawuli bokukhula kwezityalo, imibala ye-photosynthetic yayidla ngokuncipha kuzo zombini ii-genotypes (Itheyibhile 2). UFeng et al. (2013) nabo babike ukwehla okukhulu komxholo we-chlorophyll kumagqabi eentlobo ezimbini zengqolowa ezivezwe kuxinzelelo lobushushu. Ukuvezwa kubushushu obuphezulu kudla ngokubangela ukwehla komxholo we-chlorophyll, okunokubangelwa kukwehla kwe-biosynthesis ye-chlorophyll, ukonakala kwee-pigment, okanye iziphumo zazo ezidibeneyo phantsi koxinzelelo lobushushu (Fahad et al., 2017). Nangona kunjalo, izityalo zerayisi ezinyangwe kakhulu nge-CK kunye ne-BA zonyusa uxinzelelo lwee-pigment ze-photosynthetic zamagqabi phantsi koxinzelelo lobushushu. Iziphumo ezifanayo nazo zibikwe nguJespersen noHuang (2015) kunye noSuchsagunpanit et al. (2015), obone ukwanda komxholo we-chlorophyll yamagqabi emva kokusetyenziswa kwee-hormone ze-zeatin kunye ne-epibrassinosteroid kwi-bentgrass kunye nerayisi exinezelekileyo ngobushushu, ngokulandelelana. Ingcaciso efanelekileyo yokuba kutheni i-CK kunye ne-BR zikhuthaza ukwanda komxholo we-chlorophyll yamagqabi phantsi koxinzelelo lobushushu oludibeneyo kukuba i-CK inokuphucula ukuqaliswa kokungeniswa okuqhubekayo kwezikhuthazi zokubonakalisa (ezifana ne-senescence-activating promoter (SAG12) okanye i-HSP18 promoter) kwaye inciphise ukulahleka kwe-chlorophyll kumagqabi. , ukulibazisa ukuguga kwamagqabi kunye nokunyusa ukumelana nezityalo kubushushu (Liu et al., 2020). I-BR inokukhusela i-chlorophyll yamagqabi kwaye yonyuse umxholo we-chlorophyll yamagqabi ngokusebenzisa okanye ukukhuthaza ukwenziwa kwee-enzymes ezibandakanyeka kwi-chlorophyll biosynthesis phantsi kweemeko zoxinzelelo (Sharma et al., 2017; Siddiqui et al., 2018). Okokugqibela, ii-phytohormones ezimbini (i-CK kunye ne-BR) nazo zikhuthaza ukubonakaliswa kweeproteni zokutshisa ubushushu kwaye ziphucula iinkqubo ezahlukeneyo zohlengahlengiso lwe-metabolic, njengokwanda kwe-chlorophyll biosynthesis (Sharma et al., 2017; Liu et al., 2020).
Iiparameter ze-Chlorophyll, i-fluorescence, zibonelela ngendlela ekhawulezayo nengonakalisiyo enokuvavanya ukunyamezela kwezityalo okanye ukuziqhelanisa neemeko zoxinzelelo lwe-abiotic (Chaerle et al. 2007; Kalaji et al. 2017). Iiparameter ezifana nomlinganiselo we-Fv/Fm zisetyenzisiwe njengezalathisi zokuziqhelanisa nezityalo kwiimeko zoxinzelelo (Alvarado-Sanabria et al. 2017; Chavez-Arias et al. 2020). Kolu phononongo, izityalo ze-SC zibonise amaxabiso aphantsi kakhulu ale nguqu, ikakhulu izityalo zerayisi ze-"F2000". U-Yin et al. (2010) bakwafumanise ukuba umlinganiselo we-Fv/Fm wamagqabi erayisi aphezulu kakhulu unciphile kakhulu kumaqondo obushushu angaphezu kwama-35°C. Ngokutsho kukaFeng et al. (2013), umlinganiselo ophantsi we-Fv/Fm phantsi koxinzelelo lobushushu ubonisa ukuba izinga lokubamba amandla okuchukumisa kunye nokuguqulwa yiziko lokusabela le-PSII liyancipha, nto leyo ebonisa ukuba iziko lokusabela le-PSII liyaqhekeka phantsi koxinzelelo lobushushu. Olu qwalaselo lusenza sigqibe kwelokuba ukuphazamiseka kwizixhobo ze-photosynthetic kubonakala ngakumbi kwiintlobo ezibuthathaka (Fedearroz 2000) kunakwiintlobo ezixhathisayo (Fedearroz 67).
Ukusetyenziswa kwe-CK okanye i-BR ngokubanzi kuphucule ukusebenza kwe-PSII phantsi kweemeko ezinzima zoxinzelelo lobushushu. Iziphumo ezifanayo zifunyenwe nguSuchsagunpanit et al. (2015), abaqaphele ukuba ukusetyenziswa kwe-BR kwandisa ukusebenza kwe-PSII phantsi koxinzelelo lobushushu kwirayisi. UKumar et al. (2020) bakwafumanise ukuba izityalo ze-chickpea ezinyangwe nge-CK (6-benzyladine) kwaye ziphantsi koxinzelelo lobushushu zonyusa umlinganiselo we-Fv/Fm, zigqiba kwelokuba ukusetyenziswa kwe-CK ngokusebenzisa umjikelo we-zeaxanthin pigment cycle kukhuthaza umsebenzi we-PSII. Ukongeza, i-BR leaf spray ithanda i-PSII photosynthesis phantsi kweemeko ezidibeneyo zoxinzelelo, nto leyo ebonisa ukuba ukusetyenziswa kwale phytohormone kubangele ukuncipha kokusasazeka kwamandla okuvuselela ii-antennae ze-PSII kwaye kwakhuthaza ukuqokelelwa kweeproteni ezincinci zoxinzelelo lobushushu kwi-chloroplasts (Ogweno et al. 2008; Kothari kunye neLachowitz). , 2021).
Umxholo we-MDA kunye ne-proline uhlala unyuka xa izityalo ziphantsi koxinzelelo lwe-abiotic xa kuthelekiswa nezityalo ezikhuliswe phantsi kweemeko ezifanelekileyo (Alvarado-Sanabria et al. 2017). Izifundo zangaphambili zikwabonise ukuba amanqanaba e-MDA kunye ne-proline ziimpawu zebhayoloji ezinokusetyenziselwa ukuqonda inkqubo yohlengahlengiso okanye impembelelo yeendlela zezolimo kwirayisi phantsi kobushushu obuphezulu basemini okanye ebusuku (Alvarado-Sanabria et al., 2017; Quintero-Calderón et al. . , 2021). Ezi zifundo zikwabonise ukuba umxholo we-MDA kunye ne-proline wawuthanda ukuba phezulu kwizityalo zerayisi ezivezwe kubushushu obuphezulu ebusuku okanye emini, ngokulandelelana. Nangona kunjalo, ukutshiza nge-foliar ye-CK kunye ne-BR kube negalelo ekunciphiseni kwe-MDA kunye nokunyuka kwamanqanaba e-proline, ikakhulu kwi-genotype enyamezelayo (Federroz 67). I-CK spray inokukhuthaza ukuvezwa kakhulu kwe-cytokinin oxidase/dehydrogenase, ngaloo ndlela inyusa umxholo weekhompawundi ezikhuselayo ezifana ne-betaine kunye ne-proline (Liu et al., 2020). I-BR ikhuthaza ukuqaliswa kwee-osmoprotectants ezifana ne-betaine, iiswekile, kunye nee-amino acids (kuquka i-free proline), igcina ibhalansi ye-cellular osmotic phantsi kweemeko ezininzi ezingalunganga zokusingqongileyo (Kothari kunye noLachowiec, 2021).
Isalathisi soxinzelelo lwezityalo (CSI) kunye nesalathisi sokunyamezelana okunxulumeneyo (RTI) zisetyenziselwa ukumisela ukuba unyango oluvavanywayo lunceda ekunciphiseni uxinzelelo olwahlukeneyo (olungaxutywanga kunye nolwe-biotic) kwaye lunefuthe elihle kwi-physiology yezityalo (Castro-Duque et al., 2020; Chavez-Arias et al., 2020). Amaxabiso e-CSI anokuqala ukusuka ku-0 ukuya ku-1, amele iimeko ezingezizo zoxinzelelo kunye noxinzelelo, ngokwahlukeneyo (Lee et al., 2010). Amaxabiso e-CSI ezityalo ezixinezelekileyo ngobushushu (SC) aqala ku-0.8 ukuya ku-0.9 (Umfanekiso 2B), nto leyo ebonisa ukuba izityalo zerayisi zichaphazeleke kakubi luxinzelelo oludibeneyo. Nangona kunjalo, ukutshiza nge-BC (0.6) okanye i-CK (0.6) ngokuyintloko kukhokelele ekwehleni kwesi salathisi phantsi kweemeko zoxinzelelo olungafaniyo xa kuthelekiswa nezityalo zerayisi ze-SC. Kwizityalo ze-F2000, i-RTI ibonise ukwanda okuphezulu xa kusetyenziswa i-CA (97.69%) kunye ne-BC (60.73%) xa kuthelekiswa ne-SA (33.52%), nto leyo ebonisa ukuba ezi zixhobo zokulawula ukukhula kwezityalo nazo zinegalelo ekuphuculeni impendulo yerayisi ekunyamezeleni umxube. Ubushushu obugqithisileyo. Ezi zikhokelo ziye zacetyiswa ukulawula iimeko zoxinzelelo kwiintlobo ezahlukeneyo. Uphononongo olwenziwe nguLee et al. (2010) lubonise ukuba i-CSI yeentlobo ezimbini zomqhaphu phantsi koxinzelelo lwamanzi oluphakathi yayimalunga ne-0.85, ngelixa amaxabiso e-CSI eentlobo ezinkcenkceshelwe kakuhle ayeqala kwi-0.4 ukuya kwi-0.6, egqiba kwelokuba olu luhlu luphawu lokulungelelaniswa kwamanzi kweentlobo. iimeko zoxinzelelo. Ngaphezu koko, uChavez-Arias et al. (2020) bavavanye ukusebenza kakuhle kwe-exicitors zokwenziwa njengecebo elibanzi lokulawula uxinzelelo kwizityalo ze-C. elegans kwaye bafumanisa ukuba izityalo ezitshizwe ngezi zinto zibonakalise i-RTI ephezulu (65%). Ngokusekelwe koku kungasentla, i-CK kunye ne-BR zinokuthathwa njengezicwangciso zezolimo ezijolise ekwandiseni ukunyamezela irayisi kuxinzelelo lobushushu oluntsonkothileyo, njengoko ezi zixhobo zokulawula ukukhula kwezityalo zikhuthaza iimpendulo ezilungileyo ze-biochemical kunye ne-physiological.
Kwiminyaka embalwa edlulileyo, uphando ngerayisi eColombia lugxile ekuvavanyeni ii-genotypes ezinyamezela amaqondo aphezulu emini okanye ebusuku kusetyenziswa iimpawu ze-physiological okanye ze-biochemical (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2021). Nangona kunjalo, kwiminyaka embalwa edlulileyo, uhlalutyo lwetekhnoloji esebenzayo, eyongayo nenenzuzo luye lwabaluleka ngakumbi ekucebiseni ulawulo oludibeneyo lwezityalo ukuphucula imiphumo yamaxesha anzima oxinzelelo lobushushu kweli lizwe (Calderón-Páez et al., 2021; Quintero-Calderon et al., 2021). Ke ngoko, iimpendulo ze-physiological kunye ne-biochemical zezityalo zerayisi kuxinzelelo lobushushu oluntsonkothileyo (40°C imini/30°C ebusuku) ezibonwe kolu phononongo zibonisa ukuba ukutshiza nge-CK okanye i-BR kusenokuba yindlela efanelekileyo yolawulo lwezityalo ukunciphisa iziphumo ezimbi. Isiphumo samaxesha oxinzelelo lobushushu oluphakathi. Olu nyango luphucule ukunyamezela zombini ii-genotypes zerayisi (i-CSI ephantsi kunye ne-RTI ephezulu), ebonisa umkhwa jikelele kwiimpendulo ze-physiological kunye ne-biochemical zezityalo phantsi koxinzelelo lobushushu oludibeneyo. Impendulo ephambili yezityalo zerayisi yayikukwehla komxholo we-GC, i-chlorophyll iyonke, ii-chlorophylls α kunye ne-β kunye nee-carotenoids. Ukongeza, izityalo zichaphazeleka kumonakalo we-PSII (iiparameter ze-fluorescence ye-chlorophyll ezinciphileyo ezifana nomlinganiselo we-Fv/Fm) kunye nokwanda kwe-lipid peroxidation. Kwelinye icala, xa irayisi inyangwa nge-CK kunye ne-BR, ezi ziphumo zimbi zancitshiswa kwaye umxholo we-proline wanda (Umzobo 4).
Umfanekiso 4. Imodeli yengcamango yemiphumo yoxinzelelo lobushushu oludibeneyo kunye nesitshizi sokulawula ukukhula kwezityalo kwizityalo zerayisi. Iintolo ezibomvu neziluhlaza okwesibhakabhaka zibonisa iziphumo ezimbi okanye ezilungileyo zonxibelelwano phakathi koxinzelelo lobushushu kunye nokusetyenziswa kwe-BR (brassinosteroid) kunye ne-CK (cytokinin) kwimpendulo ye-physiological kunye ne-biochemical, ngokulandelelana. gs: stomatal conductance; I-Chl iyonke: umxholo we-chlorophyll iyonke; I-Chl α: umxholo we-chlorophyll β; I-Cx+c: umxholo we-carotenoid;
Ngamafutshane, iimpendulo ze-physiological kunye ne-biochemical kolu phononongo zibonisa ukuba izityalo zerayisi zeFedearroz 2000 zisengozini enkulu kwixesha loxinzelelo lobushushu oluntsonkothileyo kunezityalo zerayisi zeFedearroz 67. Bonke abalawuli bokukhula abavavanyiweyo kolu phononongo (ii-auxins, ii-gibberellins, ii-cytokinins, okanye ii-brassinosteroids) babonise inqanaba elithile lokunciphisa uxinzelelo lobushushu oludibeneyo. Nangona kunjalo, ii-cytokinin kunye nee-brassinosteroids zibangele uhlengahlengiso olungcono lwezityalo njengoko zombini abalawuli bokukhula kwezityalo bonyusa umxholo we-chlorophyll, iiparameter ze-alpha-chlorophyll fluorescence, ii-gs kunye ne-RWC xa kuthelekiswa nezityalo zerayisi ngaphandle kokusetyenziswa, kwaye kwanokunciphisa umxholo we-MDA kunye nobushushu bekhanopu. Ngamafutshane, sigqiba kwelokuba ukusetyenziswa kwabalawuli bokukhula kwezityalo (ii-cytokinins kunye nee-brassinosteroids) sisixhobo esiluncedo ekulawuleni iimeko zoxinzelelo kwizityalo zerayisi ezibangelwa luxinzelelo lobushushu obukhulu ngexesha lobushushu obuphezulu.
Izinto zokuqala ezivezwe kolu phando zifakiwe kwinqaku, kwaye imibuzo engaphezulu ingathunyelwa kumbhali ohambelanayo.