E jirila usoro ndị na-achịkwa uto osisi mee ihe dị ka atụmatụ iji belata nrụgide okpomọkụ n'ọtụtụ ihe ọkụkụ

Mmepụta osikapa na-ebelata n'ihi mgbanwe ihu igwe na mgbanwe dị iche iche na Colombia.Ndị na-achịkwa uto osisiA jirila ya mee ihe dị ka atụmatụ iji belata nrụgide okpomọkụ n'ọtụtụ ihe ubi. Ya mere, ebumnuche nke ọmụmụ ihe a bụ inyocha mmetụta nke anụ ahụ (nduzi stomatal, duzie stomatal, mkpokọta ọdịnaya chlorophyll, oke Fv/Fm nke ụdị osikapa azụmaahịa abụọ ejiri nrụgide okpomọkụ jikọta (okpomọkụ ehihie na abalị dị elu), okpomọkụ mkpuchi na ọdịnaya mmiri dị n'etiti) na mgbanwe biochemical (malondialdehyde (MDA) na ọdịnaya prolinic acid). Emere nnwale mbụ na nke abụọ site na iji osisi nke ụdị osikapa abụọ bụ Federrose 67 ("F67") na Federrose 2000 ("F2000"), n'otu n'otu. E nyochachara nnwale abụọ ahụ ọnụ dị ka usoro nnwale. Ọgwụgwọ ndị e guzobere bụ ndị a: njikwa zuru oke (AC) (osisi osikapa a kụrụ na okpomọkụ kacha mma (okpomọkụ ehihie/abalị 30/25°C)), njikwa nrụgide okpomọkụ (SC) [osisi osikapa ejiri nrụgide okpomọkụ jikọtara ọnụ naanị (40/25°C). 30°C)], a na-etinye nrụgide n'osisi osikapa ma fesa ya na ihe ndị na-achịkwa uto osisi (nrụgide+AUX, nrụgide+BR, nrụgide+CK ma ọ bụ nrụgide+GA) ugboro abụọ (ụbọchị 5 tupu na ụbọchị 5 mgbe nrụgide okpomọkụ gasịrị). Ịgbasa ya na SA mụbara mkpokọta ọdịnaya chlorophyll nke ụdị abụọ ahụ (ibu ọhụrụ nke osisi osikapa "F67" na "F2000" bụ 3.25 na 3.65 mg/g, n'otu n'otu) ma e jiri ya tụnyere osisi SC (ibu ọhụrụ nke osisi "F67" bụ 2.36 na 2.56 mg). g-1)" na osikapa "F2000", itinye CK n'ahụ na-emekwa ka ike stomatal nke osisi osikapa "F2000" (499.25 vs. 150.60 mmol m-2 s) ka mma ma e jiri ya tụnyere njikwa nrụgide okpomọkụ. nrụgide okpomọkụ, okpomọkụ nke okpueze osisi ahụ na-ebelata site na 2-3 °C, ọdịnaya MDA na osisi na-ebelatakwa. Ndekọ nnabata dị n'etiti na-egosi na itinye CK (97.69%) na BR (60.73%) n'ahụ nwere ike inye aka belata nsogbu nke nrụgide okpomọkụ. ọkachasị na osisi osikapa F2000. Na mmechi, a pụrụ iwere ịgbasa BR ma ọ bụ CK n'ahụ dị ka atụmatụ ọrụ ugbo iji nyere aka belata mmetụta ọjọọ nke ọnọdụ nrụgide okpomọkụ jikọtara ọnụ na omume anụ ahụ nke osisi osikapa.
Osikapa (Oryza sativa) sitere n'ezinụlọ Poaceae ma bụrụkwa otu n'ime ọka ndị a na-akọkarị n'ụwa yana ọka na ọka wit (Bajaj na Mohanty, 2005). Ebe a na-akọ osikapa bụ hekta 617,934, mmepụta mba na 2020 bụ tọn 2,937,840, nkezi mkpụrụ ya bụ tọn 5.02/ha (Federarroz (Federación Nacional de Arroceros), 2021).
Okpomọkụ ụwa na-emetụta ihe ọkụkụ osikapa, na-ebute ụdị nrụgide dị iche iche dịka oke okpomọkụ na oge ọkọchị. Mgbanwe ihu igwe na-eme ka okpomọkụ ụwa rịa elu; A na-atụ anya na okpomọkụ ga-arị elu site na 1.0–3.7°C na narị afọ nke 21, nke nwere ike ime ka ugboro na ike nke nrụgide okpomọkụ dịkwuo elu. Mmụba okpomọkụ gburugburu ebe obibi emetụtala osikapa, na-eme ka ihe ọkụkụ na-amịpụta belata site na 6–7%. N'aka nke ọzọ, mgbanwe ihu igwe na-edugakwa na ọnọdụ gburugburu ebe obibi na-adịghị mma maka ihe ọkụkụ, dị ka oge oké ọkọchị ma ọ bụ oke okpomọkụ na mpaghara okpomọkụ na nke dị n'okpuru okpomọkụ. Na mgbakwunye, ihe omume mgbanwe dị ka El Niño nwere ike ibute nrụgide okpomọkụ ma mee ka mmebi ihe ọkụkụ ka njọ na mpaghara ụfọdụ nke okpomọkụ. Na Colombia, a na-atụ anya na okpomọkụ na mpaghara ndị na-emepụta osikapa ga-amụba site na 2–2.5°C ka ọ na-erule 2050, na-ebelata mmepụta osikapa ma na-emetụta mbugharị ngwaahịa na ahịa na usoro ọkọnọ.
A na-akọ ọtụtụ ihe ubi osikapa n'ebe okpomọkụ dị nso na oke kachasị mma maka uto ihe ubi (Shah et al., 2011). Akọpụtala na okpomọkụ kacha mma maka ehihie na abalị makauto na mmepe osikapaỌ na-abụkarị 28°C na 22°C, n'otu n'otu (Kilasi et al., 2018; Calderón-Páez et al., 2021). Okpomọkụ dị n'elu oke ndị a nwere ike ibute oge nrụgide okpomọkụ dị oke elu ruo nke siri ike n'oge oge mmepe osikapa dị nro (ịkọ ihe ubi, anthesis, okooko osisi, na ijupụta ọka), si otú a na-emetụta mkpụrụ ọka n'ụzọ na-adịghị mma. Mbelata a na mkpụrụ bụ isi n'ihi ogologo oge nrụgide okpomọkụ, nke na-emetụta physiology osisi. N'ihi mmekọrịta nke ihe dị iche iche, dị ka oge nrụgide na oke okpomọkụ ruru, nrụgide okpomọkụ nwere ike ibute ọtụtụ mmebi na-enweghị mgbagha na metabolism na mmepe osisi.
Nrụgide okpomọkụ na-emetụta usoro dị iche iche nke ahụ mmadụ na ihe ndị dị ndụ na osisi. Fotosynthesis nke akwụkwọ bụ otu n'ime usoro ndị kachasị emetụta nrụgide okpomọkụ na osisi osikapa, ebe ọ bụ na ọnụego fotosynthesis na-ebelata site na 50% mgbe okpomọkụ kwa ụbọchị gafere 35°C. Mmeghachi omume anụ ahụ nke osisi osikapa na-adabere na ụdị nrụgide okpomọkụ. Dịka ọmụmaatụ, a na-egbochi ọnụego fotosynthesis na stomatal conductance mgbe osisi na-ekpughere okpomọkụ ehihie dị elu (33–40°C) ma ọ bụ okpomọkụ ehihie na abalị dị elu (35–40°C n'ehihie, 28–30°C). C pụtara abalị) (Lü et al., 2013; Fahad et al., 2016; Chaturvedi et al., 2017). Okpomọkụ abalị dị elu (30°C) na-akpata obere mgbochi nke photosynthesis mana ọ na-eme ka iku ume n'abalị dịkwuo mma (Fahad et al., 2016; Alvarado-Sanabria et al., 2017). N'agbanyeghị oge nrụgide ahụ, nrụgide okpomọkụ na-emetụtakwa ọdịnaya chlorophyll akwụkwọ, oke nke fluorescence variable chlorophyll ruo max chlorophyll fluorescence (Fv/Fm), na ọrụ Rubisco na osisi osikapa (Cao et al. 2009; Yin et al. 2010). ) Sanchez Reynoso et al., 2014).
Mgbanwe kemịkalụ nke ihe ọkụkụ bụ akụkụ ọzọ nke mgbanwe osisi na nrụgide okpomọkụ (Wahid et al., 2007). Ejirila ọdịnaya proline dị ka ihe ngosi kemịkalụ nke nrụgide osisi (Ahmed na Hassan 2011). Proline na-arụ ọrụ dị mkpa na metabolism osisi ebe ọ na-arụ ọrụ dị ka isi iyi carbon ma ọ bụ nitrogen yana dị ka ihe na-eme ka membrane guzosie ike n'okpuru ọnọdụ okpomọkụ dị elu (Sánchez-Reinoso et al., 2014). Okpomọkụ dị elu na-emetụtakwa nkwụsi ike nke akpụkpọ ahụ site na peroxidation lipid, na-eduga na nhazi nke malondialdehyde (MDA) (Wahid et al., 2007). Ya mere, ejirila ọdịnaya MDA ghọta iguzosi ike n'ezi ihe nke akpụkpọ ahụ sel n'okpuru nrụgide okpomọkụ (Cao et al., 2009; Chavez-Arias et al., 2018). N'ikpeazụ, nrụgide okpomọkụ jikọtara ọnụ [37/30°C (ehihie/abalị)] mụbara pasent nke ntapu electrolyte na ọdịnaya malondialdehyde na osikapa (Liu et al., 2013).
A tụlela ojiji nke ndị na-achịkwa uto osisi (GRs) iji belata mmetụta ọjọọ nke nrụgide okpomọkụ, ebe ihe ndị a na-etinye aka nke ọma na nzaghachi osisi ma ọ bụ usoro nchebe anụ ahụ megide nrụgide dị otú ahụ (Peleg na Blumwald, 2011; Yin et al. et al., 2011; Ahmed et al., 2015). Ngwa nke ihe ndị sitere na mkpụrụ ndụ ihe nketa nwere mmetụta dị mma na nnabata nrụgide okpomọkụ na ihe ọkụkụ dị iche iche. Ọmụmụ ihe egosila na phytohormones dị ka gibberellins (GA), cytokinins (CK), auxins (AUX) ma ọ bụ brassinosteroids (BR) na-eduga na mmụba na mgbanwe dị iche iche nke physiological na biochemical (Peleg na Blumwald, 2011; Yin et al. Ren, 2011; Mitler et al., 2012; Zhou et al., 2014). Na Colombia, a ghọtabeghị ma mụọ ojiji nke ihe ndị sitere na mkpụrụ ndụ ihe nketa na mmetụta ya na ihe ọkụkụ osikapa nke ọma. Agbanyeghị, nnyocha gara aga gosiri na ịgbasa akwụkwọ nri nke BR nwere ike ime ka ndidi osikapa ka mma site n'ime ka njirimara mgbanwe gas, ọdịnaya chlorophyll ma ọ bụ proline nke akwụkwọ mkpụrụ osisi osikapa ka mma (Quintero-Calderón et al., 2021).
Cytokinins na-eme ka mmeghachi omume osisi na nrụgide abiotic dị mfe, gụnyere nrụgide okpomọkụ (Ha et al., 2012). Na mgbakwunye, akọpụtala na itinye CK n'èzí nwere ike ibelata mmebi okpomọkụ. Dịka ọmụmaatụ, itinye zeatin n'èzí na-eme ka ọnụego photosynthesis, ọdịnaya chlorophyll a na b dịkwuo elu, yana arụmọrụ mbufe electron na bentgrass na-akpụ akpụ (Agrotis estolonifera) n'oge nrụgide okpomọkụ (Xu na Huang, 2009; Jespersen na Huang, 2015). Ntinye zeatin n'èzí nwekwara ike ime ka ọrụ antioxidant dịkwuo mma, melite njikọta nke protein dị iche iche, belata mmebi nke ụdị oxygen reactive (ROS) na mmepụta malondialdehyde (MDA) na anụ ahụ osisi (Chernyadyev, 2009; Yang et al., 2009)., 2016; Kumar et al., 2020).
Ojiji nke gibberellic acid egosikwala na ọ na-eme ka nrụgide okpomọkụ dị mma. Nnyocha egosila na biosynthesis GA na-akwalite ụzọ dị iche iche nke metabolic ma na-eme ka ndidi dịkwuo elu n'okpuru ọnọdụ okpomọkụ dị elu (Alonso-Ramirez et al. 2009; Khan et al. 2020). Abdel-Nabi et al. (2020) chọpụtara na ịgbasa akwụkwọ nke exogenous GA (25 ma ọ bụ 50 mg*L) nwere ike ime ka ọnụego photosynthetic na ọrụ antioxidant dịkwuo elu na osisi oroma nwere nrụgide okpomọkụ ma e jiri ya tụnyere osisi ndị na-achịkwa. A hụkwara na itinye HA n'èzí na-eme ka ọdịnaya mmiri dịkwuo elu, ọdịnaya chlorophyll na carotenoid dịkwuo elu ma na-ebelata lipid peroxidation na date palm (Phoenix dactylifera) n'okpuru nrụgide okpomọkụ (Khan et al., 2020). Auxin na-arụkwa ọrụ dị mkpa n'ịchịkwa mmeghachi omume uto na-agbanwe agbanwe na ọnọdụ okpomọkụ dị elu (Sun et al., 2012; Wang et al., 2016). Ihe na-achịkwa uto a na-arụ ọrụ dị ka ihe nrịbama kemịkalụ n'ọtụtụ usoro dịka njikọta proline ma ọ bụ mmebi n'okpuru nrụgide abiotic (Ali et al. 2007). Na mgbakwunye, AUX na-emekwa ka ọrụ antioxidant dịkwuo mma, nke na-eduga na mbelata na MDA na osisi n'ihi mbelata lipid peroxidation (Bielach et al., 2017). Sergeev et al. (2018) hụrụ na n'ime osisi pea (Pisum sativum) n'okpuru nrụgide okpomọkụ, ọdịnaya nke proline - dimethylaminoethoxycarbonylmethyl)naphthylchloromethyl ether (TA-14) na-abawanye. N'otu nnwale ahụ, ha hụkwara ọkwa MDA dị ala na osisi a gwọrọ ma e jiri ya tụnyere osisi a na-anaghị agwọ AUX.
Brassinosteroids bụ otu ụdị ihe ndị ọzọ na-achịkwa uto nke e ji ebelata mmetụta nke nrụgide okpomọkụ. Ogweno na ndị otu ya (2008) kọrọ na mmiri BR nke sitere na mpụga mụbara ọsọ photosynthesis net, stomatal conductance na oke ọsọ nke osisi Rubisco carboxylation nke tomato (Solanum lycopersicum) n'okpuru nrụgide okpomọkụ ruo ụbọchị 8. ​​Ịgbasa epibrassinosteroids nke akwụkwọ nri nwere ike ime ka ọnụego photosynthesis net nke osisi kukumba (Cucumis sativus) dịkwuo elu n'okpuru nrụgide okpomọkụ (Yu et al., 2004). Na mgbakwunye, itinye BR n'èzí na-egbu oge mmebi chlorophyll ma na-eme ka arụmọrụ ojiji mmiri na oke mmepụta quantum nke PSII photochemistry dịkwuo elu na osisi n'okpuru nrụgide okpomọkụ (Holá et al., 2010; Toussagunpanit et al., 2015).
N'ihi mgbanwe ihu igwe na mgbanwe, ihe ọkụkụ osikapa na-eche ihu oge okpomọkụ kwa ụbọchị (Lesk et al., 2016; Garcés, 2020; Federaroz (Federación Nacional de Arroceros), 2021). Na phenotyping osisi, a mụọla iji phytonutrients ma ọ bụ biostimulants dị ka atụmatụ iji belata nrụgide okpomọkụ na mpaghara osikapa (Alvarado-Sanabria et al., 2017; Calderón-Páez et al., 2021; Quintero-Calderón et al., 2021). Tinyere nke a, ojiji nke ihe ndị na-agbanwe agbanwe na ihe ndị na-agbanwe agbanwe (okpomọkụ akwụkwọ, ihe ndị na-eduzi stomatal, paramita fluorescence chlorophyll, chlorophyll na mmiri dị n'otu, malondialdehyde na proline synthesis) bụ ngwa a pụrụ ịtụkwasị obi maka inyocha osisi osikapa n'okpuru nrụgide okpomọkụ n'ógbè na mba ụwa (Sánchez -Reynoso et al., 2014; Alvarado-Sanabria et al., 2017; Agbanyeghị, nnyocha gbasara ojiji nke ọgwụ phytohormonal foliar na osikapa na ọkwa mpaghara ka dị ụkọ. Ya mere, ọmụmụ nke mmeghachi omume physiological na biochemical nke itinye ndị na-achịkwa uto osisi dị oke mkpa maka atụmatụ nke atụmatụ agronomic bara uru maka nke a. na-edozi mmetụta ọjọọ nke oge nrụgide okpomọkụ dị mgbagwoju anya na osikapa. Ya mere, ebumnuche nke ọmụmụ a bụ inyocha physiological (nduzi stomatal, paramita fluorescence chlorophyll na mmiri dị n'otu) na mmetụta biochemical nke itinye foliar nke ihe anọ na-achịkwa uto osisi (AUX, CK, GA na BR). (Pigments photosynthetic, malondialdehyde na proline ọdịnaya) Mgbanwe na ụdị osikapa azụmaahịa abụọ a na-etinye n'ọnọdụ okpomọkụ jikọtara ọnụ (oke okpomọkụ ehihie/abalị).
N'ọmụmụ ihe a, e mere nnwale abụọ n'onwe ha. E jiri mkpụrụ ndụ ihe nketa Federrose 67 (F67: mkpụrụ ndụ ihe nketa nke e mepụtara n'oge okpomọkụ dị elu n'ime afọ iri gara aga) na Federrose 2000 (F2000: mkpụrụ ndụ ihe nketa nke e mepụtara n'ime afọ iri gara aga nke narị afọ nke 20 nke na-egosi iguzogide nje akwụkwọ ọcha) mee ihe maka oge mbụ. mkpụrụ. na nnwale nke abụọ, n'otu n'otu. Ndị ọrụ ugbo Colombia na-akọ mkpụrụ abụọ ahụ nke ọma. A kụrụ mkpụrụ n'ime tray 10-L (ogologo 39.6 cm, obosara 28.8 cm, elu 16.8 cm) nke nwere ala ájá ájá nwere ihe organic 2% . A kụrụ mkpụrụ ise a kụrụ tupu oge eruo na tray ọ bụla. E tinyere pallets ndị ahụ n'ime ụlọ ntu nke Faculty of Agricultural Sciences nke National University of Colombia, Bogotá campus (43°50′56″ N, 74°04′051″ W), n'elu elu 2556 m n'elu oke osimiri (asl). m.) ma e mere ha site na Ọktoba ruo Disemba 2019. Otu nnwale (Federroz 67) na nnwale nke abụọ (Federroz 2000) n'otu oge ahụ nke 2020.
Ọnọdụ gburugburu ebe obibi n'ime griin haus n'oge ọ bụla a na-akụ ihe bụ ndị a: okpomọkụ ehihie na abalị 30/25°C, iru mmiri dị n'etiti 60~80%, oge foto eke awa 12 (radiation nke na-arụ ọrụ site na photosynthetic 1500 µmol (photons) m-2 s-). 1 n'ehihie). A na-eme fatịlaịza osisi dịka ọdịnaya nke ihe ọ bụla si dị ụbọchị 20 mgbe mkpụrụ pụtara (DAE), dịka Sánchez-Reinoso et al. (2019) si kwuo: nitrogen 670 mg kwa osisi, phosphorus 110 mg kwa osisi, potassium 350 mg kwa osisi, calcium 68 mg kwa osisi, magnesium 20 mg kwa osisi, sọlfọ 20 mg kwa osisi, silicon 17 mg kwa osisi. Osisi ndị ahụ nwere boron 10 mg kwa osisi, ọla kọpa 17 mg kwa osisi, na zinc 44 mg kwa osisi. A na-ejigide osisi osikapa ruo DAE 47 na nnwale ọ bụla mgbe ha ruru ọkwa phenological V5 n'oge a. Nnyocha ndị gara aga egosila na oge a nke phenological bụ oge kwesịrị ekwesị iji mee nnyocha maka nrụgide okpomọkụ na osikapa (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2017).
N'ime nnwale ọ bụla, e mere ngwa abụọ dị iche iche nke ihe na-achịkwa uto akwụkwọ. E tinyere usoro mbụ nke ọgwụ phytohormone foliar ụbọchị ise tupu ọgwụgwọ nrụgide okpomọkụ (42 DAE) iji kwadebe osisi maka nrụgide gburugburu ebe obibi. E nyere ọgwụ foliar nke abụọ ụbọchị ise mgbe osisi ndị ahụ kpughesịrị ọnọdụ nrụgide (52 DAE). E jiri phytohormones anọ mee ihe, e depụtakwara ihe onwunwe nke ihe ọ bụla na-arụ ọrụ nke a gbara n'ọmụmụ ihe a na Tebụl Mgbakwunye 1. Ọnụọgụ nke ihe ndị na-achịkwa uto akwụkwọ ejiri mee bụ ndị a: (i) Auxin (1-naphthylacetic acid: NAA) na mkpokọta nke 5 × 10−5 M (ii) 5 × 10–5 M gibberellin (gibberellic acid: NAA); GA3); (iii) Cytokinin (trans-zeatin) 1 × 10-5 M (iv) Brassinosteroids [Spirostan-6-one, 3,5-dihydroxy-, (3b,5a,25R)] 5 × 10-5; M. A họọrọ oke ndị a n'ihi na ha na-akpali nzaghachi dị mma ma na-eme ka osisi ghara iguzogide nrụgide okpomọkụ (Zahir et al., 2001; Wen et al., 2010; El-Bassiony et al., 2012; Salehifar et al., 2017). Osisi osikapa na-enweghị ihe ọ bụla na-achịkwa uto osisi ka ejiri mmiri a gwakọtara agwakọta gwọọ. A na-agba ahịhịa osikapa niile na ihe na-agba mmanụ aka. Tinye 20 ml H2O na osisi ahụ iji mee ka elu na ala nke akwụkwọ dị mmiri mmiri. Ihe niile e ji fesa akwụkwọ foliar mee jiri ihe na-enyere aka n'ọrụ ugbo (Agrotin, Bayer CropScience, Colombia) na 0.1% (v/v). Anya dị n'etiti ite na ihe na-agba mmanụ bụ 30 cm.
E nyere ọgwụgwọ nrụgide okpomọkụ ụbọchị ise ka e mesịrị ọgwụ mbụ e fesa foliar (47 DAE) na nnwale ọ bụla. E si na griin haus bufee osisi osikapa gaa na ụlọ uto 294 L (MLR-351H, Sanyo, IL, USA) iji guzobe nrụgide okpomọkụ ma ọ bụ nọgide na-enwe otu ọnọdụ gburugburu ebe obibi (47 DAE). E mere ọgwụgwọ nrụgide okpomọkụ jikọtara ọnụ site n'ịtọ ụlọ ahụ ka ọ dị na okpomọkụ ehihie/abalị ndị a: okpomọkụ dị elu n'ehihie [40°C maka awa ise (site na elekere 11:00 ruo elekere 16:00)] na oge abalị [30°C maka awa ise]. ụbọchị asatọ n'usoro (site na elekere 19:00 ruo elekere 24:00). A họọrọ okpomọkụ nrụgide na oge ikpughe dabere na ọmụmụ ihe gara aga (Sánchez-Reynoso et al. 2014; Alvarado-Sanabría et al. 2017). N'aka nke ọzọ, e tinyere otu osisi e bufere n'ime ụlọ uto n'ime ụlọ griin ha n'otu okpomọkụ ahụ (30°C n'ehihie/25°C n'abalị) ruo ụbọchị asatọ n'usoro.
Ná ngwụcha nnwale ahụ, e nwetara otu ọgwụgwọ ndị a: (i) ọnọdụ okpomọkụ uto + itinye mmiri a gwakọtara agwakọta [Njikwa zuru oke (AC)], (ii) ọnọdụ nrụgide okpomọkụ + itinye mmiri a gwakọtara agwakọta [Njikwa nrụgide okpomọkụ (SC)], (iii) ọnọdụ ọnọdụ nrụgide okpomọkụ + itinye auxin (AUX), (iv) ọnọdụ nrụgide okpomọkụ + itinye gibberellin (GA), (v) ọnọdụ nrụgide okpomọkụ + itinye cytokinin (CK), na (vi) ọnọdụ nrụgide okpomọkụ + brassinosteroid (BR) Mgbakwunye. E jiri otu ọgwụgwọ ndị a maka genotypes abụọ (F67 na F2000). Emere ọgwụgwọ niile n'usoro zuru oke yana ihe ise e ji mee ihe, nke ọ bụla nwere otu osisi. E jiri osisi ọ bụla gụọ ihe ndị a chọpụtara na njedebe nke nnwale ahụ. Nnwale ahụ were 55 DAE.
E jiri ihe eji ebugharị ebugharị (SC-1, METER Group Inc., USA) tụọ stomatal conductance (gs) site na 0 ruo 1000 mmol m-2 s-1, yana oghere ụlọ nlele nke 6.35 mm. A na-eme nha site na itinye ihe nyocha stomameter na akwụkwọ tozuru etozu nke nwere isi ihe ọkụkụ ahụ gbasaara nke ọma. Maka ọgwụgwọ ọ bụla, a na-agụ gs na akwụkwọ atọ nke osisi ọ bụla n'etiti elekere 11:00 na elekere 16:00 ma na-agbakọ nkezi.
A chọpụtara RWC dịka usoro Ghoulam et al. (2002) si kọwaa. E jikwa mpempe akwụkwọ gbasaara nke ọma e ji chọpụta g tụọ RWC. A na-eji usoro dijitalụ achọpụta ibu ọhụrụ (FW) ozugbo e wechara ya. E tinyeziri akwụkwọ ndị ahụ n'ime akpa plastik e ji mmiri jupụta ma hapụ ha n'ọchịchịrị na okpomọkụ ụlọ (22°C) ruo awa 48. Mgbe ahụ, tụọ ha na usoro dijitalụ ma dekọọ ibu gbasaara (TW). A kpọrọ nkụ akwụkwọ ndị ahụ sara mbara na oven na okpomọkụ 75°C ruo awa 48, e dekọọkwa ibu ha kpọrọ nkụ (DW).
E jiri mita chlorophyll (atLeafmeter, FT Green LLC, USA) chọpụta ọdịnaya chlorophyll dị n'etiti ha ma gosipụta ya na nkeji atLeaf (Dey et al., 2016). E jiri usoro chlorophyll fluorimeter na-aga n'ihu (Handy PEA, Hansatech Instruments, UK) dekọọ ọnụọgụgụ arụmọrụ PSII kachasị elu (Fv/Fm ratio). E jiri ihe mkpuchi akwụkwọ mee ka akwụkwọ ahụ gbaa ọchịchịrị ruo nkeji iri abụọ tupu nha Fv/Fm (Restrepo-Diaz na Garces-Varon, 2013). Mgbe akwụkwọ ahụ dị ọchịchịrị, a tụrụ ntọala (F0) na ọkụ kachasị elu (Fm). Site na data ndị a, a gbakọrọ mmụba ọkụ (Fv = Fm – F0), oke ọkụ ọkụ na oke ọkụ (Fv/Fm), oke ọkụ ọkụ na oke ọkụ (PSII photochemistry) kachasị (Fv/F0) na oke ọkụ ọkụ (Fm/F0) (Baker, 2008; Lee et al. ., 2017). E jiri otu akwụkwọ e ji atụ nha gs mee ihe mee ihe dịka ọkụ ọkụ na oke ọkụ chlorophyll.
A chịkọtara ihe dị ka 800 mg nke ibu ọhụrụ nke akwụkwọ dị ka ihe ndị na-agbanwe agbanwe na kemịkalụ. E mesịa, e tinyere ihe nlele akwụkwọ n'ime nitrogen mmiri mmiri ma chekwaa ya maka nyocha ọzọ. Usoro spectrometric eji eme atụmatụ ọdịnaya chlorophyll a, b na carotenoid nke anụ ahụ dabere na usoro na nha anya nke Wellburn (1994) kọwara. A chịkọtara ihe nlele anụ ahụ (30 mg) ma mee ka ha dị ka otu na 3 ml nke 80% acetone. E mesịa, e tinyere ihe nlele ndị ahụ na centrifuge (ụdị 420101, Becton Dickinson Primary Care Diagnostics, USA) na 5000 rpm maka nkeji 10 iji wepụ ihe ndị ahụ. A gwakọtara ihe dị n'elu ya ruo olu ikpeazụ nke 6 ml site na ịgbakwunye 80% acetone (Sims na Gamon, 2002). A chọpụtara ọdịnaya nke chlorophyll na 663 (chlorophyll a) na 646 (chlorophyll b) nm, na carotenoids na 470 nm site na iji spectrophotometer (Spectronic BioMate 3 UV-vis, Thermo, USA).
E jiri usoro thiobarbituric acid (TBA) nke Hodges et al. (1999) kọwara mee ihe iji chọpụta lipid peroxidation nke membrane (MDA). E mekwara ka ihe dị ka 0.3 g nke anụ ahụ akwụkwọ dị n'otu na nitrogen mmiri mmiri. E tinyere ihe nlele ndị ahụ na 5000 rpm ma tụọ absorbance na spectrophotometer na 440, 532 na 600 nm. N'ikpeazụ, e jiri ihe e ji ewepụ ihe (157 M mL−1) gbakọọ mkpokọta MDA.
E jiri usoro Bates et al. (1973) kọwara chọpụta ọdịnaya nke proline nke ọgwụgwọ niile. Tinye 10 ml nke mmiri sulfosalicylic acid 3% na ihe nlele echekwara ma tee ya site na akwụkwọ nzacha Whatman (Nke 2). E mesịa, e jiri 2 ml nke ninhydric acid na 2 ml nke glacial acetic acid meghachi omume na 2 ml nke filtrate a. E tinyere ngwakọta ahụ n'ime mmiri ịsa ahụ na 90°C maka otu awa. Kwụsị mmeghachi omume ahụ site na itinye ya na ice. Ma jijiji tube ahụ nke ọma site na iji vortex shaker wee gbazee ngwọta ahụ na 4 ml nke toluene. E kpebiri ịgụ absorbance na 520 nm site na iji otu spectrophotometer ejiri mee ihe maka nha nke photosynthetic pigments (Spectronic BioMate 3 UV-Vis, Thermo, Madison, WI, USA).
Usoro Gerhards et al. (2016) kọwara iji gbakọọ okpomọkụ mkpuchi na CSI. E sere foto okpomọkụ site na iji igwefoto FLIR 2 (FLIR Systems Inc., Boston, MA, USA) nke ziri ezi nke ±2°C na njedebe nke oge nrụgide ahụ. Debe elu ọcha n'azụ osisi ahụ maka foto. Ọzọkwa, a na-ewere ụlọ ọrụ mmepụta ihe abụọ dị ka ihe atụ ntụaka. E tinyere osisi ndị ahụ n'elu elu ọcha; e ji ihe na-enyere aka n'ọrụ ugbo (Agrotin, Bayer CropScience, Bogotá, Colombia) tee otu iji mee ka oghere nke stomata niile [ụdị mmiri mmiri (Twet)], nke ọzọ bụkwa akwụkwọ na-enweghị ihe ọ bụla e tinyere [ụdị mmiri kpọrọ nkụ (Tdry)] (Castro-Duque et al., 2020). Anya dị n'etiti igwefoto na ite ahụ n'oge ịse ihe nkiri bụ 1 m.
E jiri usoro stomatal conductance (gs) nke osisi e gwọrọ agwọ gbakọọ ihe nrịbama ndidi n'ụzọ na-apụtaghị ìhè ma e jiri ya tụnyere osisi ndị e gwọrọ agwọ ọrịa (osisi ndị na-enweghị ọgwụgwọ nrụgide na ndị na-achịkwa uto) iji chọpụta nnabata nke genotypes e gwọrọ agwọ ọrịa a. E nwetara RTI site na iji usoro nha nhata nke e si na Chávez-Arias et al. (2020) nweta.
N'ime nnwale ọ bụla, a chọpụtara ma dekọọ mgbanwe niile dị n'ahụ mmadụ nke a kpọtụrụ aha n'elu na 55 DAE site na iji akwụkwọ ndị a gbasapụrụ agbatị nke ọma nke a nakọtara site na elu mkpuchi. Na mgbakwunye, a na-eme nha n'ime ụlọ uto iji zere ịgbanwe ọnọdụ gburugburu ebe obibi nke osisi ndị ahụ na-eto.
E jiri usoro nnwale nke mbụ na nke abụọ nyochaa data sitere na nnwale mbụ na nke abụọ. Otu nnwale ọ bụla nwere osisi ise, osisi ọ bụla mekwara otu nnwale. E mere nyocha nke ọdịiche (ANOVA) (P ≤ 0.05). Mgbe achọpụtara nnukwu ọdịiche, e jiri ule ntụnyere nke Tukey na P ≤ 0.05. Jiri ọrụ arcsine gbanwee uru pasentị. E jiri ngwanrọ Statistix v 9.0 (Ngwanrọ Nyocha, Tallahassee, FL, USA) nyochaa data ma jiri SigmaPlot (ụdị 10.0; Systat Software, San Jose, CA, USA) see ya. E jiri ngwanrọ InfoStat 2016 (Ngwanrọ Nyocha, Mahadum Mba nke Cordoba, Argentina) mee nyocha akụkụ bụ isi iji chọpụta ndị na-achịkwa uto osisi kacha mma a na-amụ.
Tebụl 1 chịkọtara ANOVA nke na-egosi nnwale ndị ahụ, ọgwụgwọ dị iche iche, na mmekọrịta ha na pigments photosynthetic nke akwụkwọ (chlorophyll a, b, total, na carotenoids), malondialdehyde (MDA) na ọdịnaya proline, na stomatal conductance. Mmetụta nke gs, ọdịnaya mmiri dị n'otu. (RWC), ọdịnaya chlorophyll, paramita fluorescence chlorophyll alpha, okpomọkụ okpueze (PCT) (°C), ndeksi nrụgide ihe ubi (CSI) na ndeksi ndidi nke osisi osikapa na 55 DAE.
Tebụl 1. Nchịkọta nke data ANOVA na mgbanwe anụ ahụ na biochemical nke osikapa n'etiti nnwale (genotypes) na ọgwụgwọ nrụgide okpomọkụ.
E gosipụtara ọdịiche dị iche iche (P≤0.01) na mmekọrịta fotosynthetic pigment nke akwụkwọ, ọdịnaya chlorophyll dị n'etiti (agụmakwụkwọ Atleaf), na paramita fluorescence nke alpha-chlorophyll n'etiti nnwale na ọgwụgwọ na Tebụl 2. Okpomọkụ dị elu nke ehihie na abalị mụbara mkpokọta chlorophyll na carotenoid ọdịnaya. Mkpụrụ osisi osikapa na-enweghị ihe ọ bụla na-agba phytohormones (2.36 mg g-1 maka "F67" na 2.56 mg g-1 maka "F2000") ma e jiri ya tụnyere osisi ndị a kụrụ n'okpuru ọnọdụ okpomọkụ kacha mma (2.67 mg g-1)) gosiri obere mkpokọta chlorophyll ọdịnaya. Na nnwale abụọ ahụ, "F67" bụ 2.80 mg g-1 na "F2000" bụ 2.80 mg g-1. Na mgbakwunye, mkpụrụ osisi osikapa ejiri ngwakọta nke ọgwụ AUX na GA gwọọ n'okpuru nrụgide okpomọkụ gosikwara mbelata na ọdịnaya chlorophyll na genotypes abụọ ahụ (AUX = 1.96 mg g-1 na GA = 1.45 mg g-1 maka "F67"; AUX = 1.96 mg g-1 na GA = 1.45 mg g-1 maka "F67"; AUX = 2.24 mg) g-1 na GA = 1.43 mg g-1 (maka "F2000") n'okpuru ọnọdụ nrụgide okpomọkụ. N'okpuru ọnọdụ nrụgide okpomọkụ, ọgwụgwọ foliar na BR mere ka mmụba dị ntakịrị na mgbanwe a na genotypes abụọ ahụ. N'ikpeazụ, ọgwụ foliar CK gosiri uru kachasị elu nke agba photosynthetic n'etiti ọgwụgwọ niile (ọgwụgwọ AUX, GA, BR, SC na AC) na genotypes F67 (3.24 mg g-1) na F2000 (3.65 mg g-1). E mekwara ka nrụgide okpomọkụ jikọtara ọnụ belata ọdịnaya chlorophyll (otu Atleaf). E dekọkwara ọnụ ọgụgụ kachasị elu n'osisi ndị a na-agba CC n'ime genotypes abụọ ahụ (41.66 maka "F67" na 49.30 maka "F2000"). Oke Fv na Fv/Fm gosiri nnukwu ọdịiche dị n'etiti ọgwụgwọ na ụdị (Tebụl 2). N'ozuzu, n'etiti mgbanwe ndị a, ụdị F67 adịghị enwe mmetụta nrụgide okpomọkụ karịa ụdị F2000. Oke Fv na Fv/Fm tara ahụhụ karịa na nnwale nke abụọ. Mkpụrụ osisi 'F2000' nke a na-agbaghị na phytohormones ọ bụla nwere oke Fv kacha ala (2120.15) na oke Fv/Fm (0.59), mana ịgba foliar na CK nyere aka weghachite ụkpụrụ ndị a (Fv: 2591, 89, oke Fv/Fm: 0.73). , na-enweta ọgụgụ yiri nke e dekọrọ na osisi "F2000" a kụrụ n'okpuru ọnọdụ okpomọkụ kacha mma (Fv: 2955.35, Fv/Fm ratio: 0.73:0.72). Enweghị ọdịiche dị mkpa na fluorescence mbụ (F0), fluorescence kachasị (Fm), oke photochemical quantum mmepụta nke PSII (Fv/F0) na oke Fm/F0. N'ikpeazụ, BR gosiri usoro yiri nke ahụ dịka a hụrụ na CK (Fv 2545.06, oke Fv/Fm 0.73).
Tebụl 2. Mmetụta nke nrụgide okpomọkụ jikọtara ọnụ (40°/30°C n'ehihie/abalị) na agba fotosynthetic nke akwụkwọ [mkpokọta chlorophyll (Ngụkọta Chl), chlorophyll a (Chl a), chlorophyll b (Chl b) na carotenoids mmetụta Cx+c]], ọdịnaya chlorophyll dị n'otu (Atliff unit), paramita fluorescence chlorophyll (F0 mbụ), fluorescence kachasị (Fm), fluorescence agbanwe agbanwe (Fv), arụmọrụ PSII kachasị (Fv/Fm), mmepụta quantum kachasị fotochemical nke PSII (Fv/F0) na Fm/F0 na osisi nke genotypes osikapa abụọ [Federrose 67 (F67) na Federrose 2000 (F2000)] ụbọchị 55 mgbe ọ pụtasịrị (DAE)).
Mmiri dị n'etiti osisi osikapa e ji aka mee (RWC) gosiri ọdịiche dị n'etiti ha (P ≤ 0.05) na mmekọrịta dị n'etiti ọgwụgwọ nnwale na nke akwụkwọ (Foto 1A). Mgbe ejiri SA gwọọ ha, e dekọrọ uru kacha ala maka ụdị mkpụrụ ndụ ihe nketa abụọ ahụ (74.01% maka F67 na 76.6% maka F2000). N'okpuru ọnọdụ nrụgide okpomọkụ, RWC nke osisi osikapa nke ụdị mkpụrụ ndụ ihe nketa abụọ e ji phytohormones dị iche iche gwọọ mụbara nke ukwuu. N'ozuzu, ojiji nke mkpụrụ ndụ ihe nketa nke CK, GA, AUX, ma ọ bụ BR mụbara RWC ka ọ bụrụ uru yiri nke osisi ndị a kụrụ n'okpuru ọnọdụ kacha mma n'oge nnwale ahụ. Njikwa zuru oke na osisi ndị a gbasasịrị akwụkwọ akụkọ ruru ihe dị ka 83% maka ụdị mkpụrụ ndụ ihe nketa abụọ ahụ. N'aka nke ọzọ, gs gosikwara ọdịiche dị mkpa (P ≤ 0.01) na mmekọrịta nnwale na ọgwụgwọ (Foto 1B). Osisi njikwa zuru oke (AC) dekọkwara ụkpụrụ kachasị elu maka ụdị mkpụrụ ndụ ihe nketa ọ bụla (440.65 mmol m-2s-1 maka F67 na 511.02 mmol m-2s-1 maka F2000). Osisi osikapa ndị a na-etinye n'ọnọdụ nrụgide okpomọkụ naanị gosiri ụkpụrụ gs kacha ala maka ụdị mkpụrụ ndụ ihe nketa abụọ ahụ (150.60 mmol m-2s-1 maka F67 na 171.32 mmol m-2s-1 maka F2000). Ọgwụgwọ akwụkwọ nri na ndị na-achịkwa uto osisi niile mụbakwara g. Na osisi osikapa F2000 nke a na-agba CC, mmetụta nke ịgba foliar na phytohormones pụtara ìhè karịa. Otu osisi a egosighi ọdịiche ọ bụla ma e jiri ya tụnyere osisi njikwa zuru oke (AC 511.02 na CC 499.25 mmol m-2s-1).
Foto 1. Mmetụta nke nrụgide okpomọkụ jikọtara ọnụ (40°/30°C n'ehihie/abalị) na ọdịnaya mmiri dị n'otu (RWC) (A), njikwa stomatal (gs) (B), mmepụta malondialdehyde (MDA) (C), na ọdịnaya proline. (D) na osisi nke ụdị mkpụrụ ndụ osikapa abụọ (F67 na F2000) na ụbọchị 55 mgbe ọ pụtasịrị (DAE). Ọgwụgwọ ndị a tụlere maka ụdị mkpụrụ ndụ ọ bụla gụnyere: njikwa zuru oke (AC), njikwa nrụgide okpomọkụ (SC), nrụgide okpomọkụ + auxin (AUX), nrụgide okpomọkụ + gibberellin (GA), nrụgide okpomọkụ + mitogen cell (CK), na nrụgide okpomọkụ + brassinosteroid. (BR). Kọlụm ọ bụla na-anọchite anya njehie ọkọlọtọ nkezi ± nke isi data ise (n = 5). Kọlụm ndị mkpụrụedemede dị iche iche sochiri na-egosi ọdịiche dị mkpa na ọnụ ọgụgụ dịka ule Tukey si dị (P ≤ 0.05). Mkpụrụ akwụkwọ nwere akara nhata na-egosi na nkezi abụghị ihe dị mkpa na ọnụ ọgụgụ (≤ 0.05).
Ihe dị n'ime MDA (P ≤ 0.01) na proline (P ≤ 0.01) gosikwara nnukwu ọdịiche dị na mmekọrịta dị n'etiti nnwale na ọgwụgwọ phytohormone (Foto 1C, D). A hụrụ mmụba nke lipid peroxidation site na ọgwụgwọ SC na genotypes abụọ ahụ (Foto 1C), agbanyeghị, osisi ejiri ọgwụ fesa nke na-achịkwa uto akwụkwọ gwọọ gosiri mbelata nke lipid peroxidation na genotypes abụọ ahụ; N'ozuzu, ojiji nke phytohormones (CA, AUC, BR ma ọ bụ GA) na-eduga na mbelata nke lipid peroxidation (ọdịnaya MDA). A hụghị ọdịiche ọ bụla n'etiti osisi AC nke genotypes abụọ na osisi ndị nọ n'okpuru nrụgide okpomọkụ ma fesa ha na phytohormones (uru FW a hụrụ na osisi "F67" sitere na 4.38–6.77 µmol g-1, na osisi FW "F2000" "uru a hụrụ sitere na 2.84 ruo 9.18 µmol g-1 (osisi). N'aka nke ọzọ, mmepụta proline na osisi "F67" dị ala karịa na osisi "F2000" n'okpuru nrụgide jikọtara ọnụ, nke butere mmụba na mmepụta proline. Na osisi osikapa ndị nwere nrụgide okpomọkụ, na nnwale abụọ ahụ, a hụrụ na nchịkwa nke homonụ ndị a mụbara ọdịnaya amino acid nke osisi F2000 nke ukwuu (AUX na BR bụ 30.44 na 18.34 µmol g-1) n'otu n'otu (Foto 1G).
E gosiri mmetụta nke ọgwụ fesa nke na-achịkwa uto osisi foliar na nrụgide okpomọkụ jikọtara ọnụ na okpomọkụ mkpuchi osisi na ihe nrịbama ndidi (RTI) na Foto 2A na B. Maka ụdị genotypes abụọ ahụ, okpomọkụ mkpuchi osisi AC dị nso na 27°C, nke osisi SC dịkwa ihe dị ka 28°C. NA. A hụkwara na ọgwụgwọ akwụkwọ osisi na CK na BR mere ka okpomọkụ mkpuchi dị na 2–3°C belata ma e jiri ya tụnyere osisi SC (Foto 2A). RTI gosipụtara omume yiri nke ahụ na mgbanwe anụ ahụ ndị ọzọ, na-egosi ọdịiche dị mkpa (P ≤ 0.01) na mmekọrịta dị n'etiti nnwale na ọgwụgwọ (Foto 2B). Osisi SC gosipụtara obere ndidi osisi na ụdị genotypes abụọ ahụ (34.18% na 33.52% maka osisi osikapa "F67" na "F2000", n'otu n'otu). Nri akwụkwọ phytohormones na-eme ka RTI dịkwuo mma na osisi ndị a na-ekpughere nrụgide okpomọkụ dị elu. Mmetụta a pụtara ìhè karịa n'osisi "F2000" a na-agba CC, nke RTI bụ 97.69. N'aka nke ọzọ, a hụrụ nnukwu ọdịiche naanị na ndepụta nrụgide mkpụrụ (CSI) nke osisi osikapa n'okpuru ọnọdụ nrụgide spray foliar factor (P ≤ 0.01) (Fig. 2B). Naanị osisi osikapa ndị a na-etinye nrụgide okpomọkụ dị mgbagwoju anya gosiri uru ndepụta nrụgide kachasị elu (0.816). Mgbe a na-agba osisi osikapa na phytohormones dị iche iche, ndepụta nrụgide dị ala (uru sitere na 0.6 ruo 0.67). N'ikpeazụ, osisi osikapa a kụrụ n'okpuru ọnọdụ kacha mma nwere uru nke 0.138.
Foto nke 2. Mmetụta nke nrụgide okpomọkụ jikọtara ọnụ (40°/30°C ehihie/abalị) na okpomọkụ mkpuchi (A), ndeksi ndidi dị n'otu (RTI) (B), na ndeksi nrụgide ihe ubi (CSI) (C) nke ụdị osisi abụọ. E tinyere ụdị mkpụrụ ndụ osikapa azụmaahịa (F67 na F2000) ọgwụgwọ okpomọkụ dị iche iche. Ọgwụgwọ ndị a tụlere maka ụdị mkpụrụ ndụ ọ bụla gụnyere: njikwa zuru oke (AC), njikwa nrụgide okpomọkụ (SC), nrụgide okpomọkụ + auxin (AUX), nrụgide okpomọkụ + gibberellin (GA), nrụgide okpomọkụ + mitogen cell (CK), na nrụgide okpomọkụ + brassinosteroid. (BR). Nrụgide okpomọkụ ejikọtara ọnụ gụnyere ikpughe osisi osikapa na okpomọkụ ehihie/abalị dị elu (40°/30°C ehihie/abalị). Kọlụm ọ bụla na-anọchite anya njehie ọkọlọtọ nkezi ± nke isi data ise (n = 5). Kọlụm ndị sochiri ya na mkpụrụedemede dị iche iche na-egosi ọdịiche dị mkpa na ọnụọgụgụ dịka ule Tukey si dị (P ≤ 0.05). Mkpụrụ akwụkwọ nwere akara nhata na-egosi na nkezi abụghị ihe dị mkpa na ọnụọgụgụ (≤ 0.05).
Nnyocha ihe mejupụtara ya (PCA) gosiri na ihe ndị a tụlere na 55 DAE kọwara 66.1% nke nzaghachi physiological na biochemical nke osisi osikapa ndị nwere nrụgide okpomọkụ nke ejiri mmiri ọgwụ na-achịkwa uto gwọọ (Foto 3). Vektọ na-anọchite anya ihe ndị na-agbanwe agbanwe na ntụpọ na-anọchite anya ihe ndị na-achịkwa uto osisi (GRs). Vektọ nke gs, ọdịnaya chlorophyll, arụmọrụ kachasị nke PSII (Fv/Fm) na paramita biochemical (TChl, MDA na proline) dị nso na mmalite, na-egosi njikọ dị elu n'etiti omume physiological nke osisi na ha. agbanwe agbanwe. Otu otu (V) gụnyere mkpụrụ osisi osikapa a kụrụ na okpomọkụ kacha mma (AT) na osisi F2000 ejiri CK na BA gwọọ. N'otu oge ahụ, ọtụtụ osisi ejiri GR gwọọ mepụtara otu dị iche (IV), ọgwụgwọ na GA na F2000 guzobere otu dị iche (II). N'ụzọ dị iche, mkpụrụ osisi osikapa ndị nwere nrụgide okpomọkụ (otu I na III) na-enweghị ihe ọ bụla na-agba phytohormones (genotypes abụọ ahụ bụ SC) dị na mpaghara dị iche na otu V, na-egosi mmetụta nke nrụgide okpomọkụ na physiology osisi.
Foto nke 3. Nyocha akụkọ ndụ nke mmetụta nke nrụgide okpomọkụ jikọtara ọnụ (40°/30°C ehihie/abalị) na osisi nke ụdị osikapa abụọ (F67 na F2000) na ụbọchị 55 mgbe ọ pụtasịrị (DAE). Nchịkọta: AC F67, njikwa zuru oke F67; SC F67, njikwa nrụgide okpomọkụ F67; AUX F67, nrụgide okpomọkụ + auxin F67; GA F67, nrụgide okpomọkụ + gibberellin F67; CK F67, nrụgide okpomọkụ + nkewa sel BR F67, nrụgide okpomọkụ + brassinosteroid. F67; AC F2000, njikwa zuru oke F2000; SC F2000, Njikwa nrụgide okpomọkụ F2000; AUX F2000, nrụgide okpomọkụ + auxin F2000; GA F2000, nrụgide okpomọkụ + gibberellin F2000; CK F2000, nrụgide okpomọkụ + cytokinin, BR F2000, nrụgide okpomọkụ + steroid ọla; F2000.
Ihe ndị dị iche iche dịka ọdịnaya chlorophyll, stomatal conductance, Fv/Fm ratio, CSI, MDA, RTI na ọdịnaya proline nwere ike inyere aka ịghọta mgbanwe nke mkpụrụ ndụ ihe nketa osikapa ma nyochaa mmetụta nke atụmatụ agronomic n'okpuru nrụgide okpomọkụ (Sarsu et al., 2018; Quintero-Calderon et al., 2021). Ebumnuche nke nnwale a bụ inyocha mmetụta nke itinye ihe nhazi uto anọ na paramita physiological na biochemical nke mkpụrụ osisi osikapa n'okpuru ọnọdụ nrụgide okpomọkụ dị mgbagwoju anya. Nnwale mkpụrụ osisi bụ ụzọ dị mfe ma dị ngwa maka nyocha osisi osikapa n'otu oge dabere na nha ma ọ bụ ọnọdụ nke akụrụngwa dị (Sarsu et al. 2018). Nsonaazụ nke ọmụmụ a gosiri na nrụgide okpomọkụ jikọtara ọnụ na-akpali nzaghachi physiological na biochemical dị iche iche na mkpụrụ ndụ ihe nketa osikapa abụọ ahụ, na-egosi usoro mgbanwe. Nsonaazụ ndị a na-egosikwa na ọgwụ ndị na-achịkwa uto foliar (karịsịa cytokinins na brassinosteroids) na-enyere osikapa aka ịgbanwe nrụgide okpomọkụ dị mgbagwoju anya n'ihi na amara na-emetụtakarị gs, RWC, Fv/Fm ratio, pigments photosynthetic na ọdịnaya proline.
Iji ihe ndị na-achịkwa uto na-enyere aka ime ka ọnọdụ mmiri nke osisi osikapa ka mma n'okpuru nrụgide okpomọkụ, nke nwere ike ịbụ njikọ na nrụgide dị elu na obere okpomọkụ nke osisi. Ọmụmụ ihe a gosiri na n'etiti osisi "F2000" (genotype nwere ike ịdị ndụ), osisi osikapa ejiri CK ma ọ bụ BR gwọọ nke ọma nwere uru GS ​​dị elu na uru PCT dị ala karịa osisi ejiri SC gwọọ. Ọmụmụ ihe ndị gara aga egosikwala na gs na PCT bụ ihe ngosi ahụike ziri ezi nke nwere ike ikpebi mmeghachi omume mgbanwe nke osisi osikapa na mmetụta nke atụmatụ agronomic na nrụgide okpomọkụ (Restrepo-Diaz na Garces-Varon, 2013; Sarsu et al., 2018; Quintero). -Carr DeLong et al., 2021). Akwụkwọ CK ma ọ bụ BR na-eme ka g dịkwuo mma n'okpuru nrụgide n'ihi na homonụ osisi ndị a nwere ike ịkwalite oghere stomatal site na mmekọrịta sịntetik na molekul ndị ọzọ na-egosi ihe dịka ABA (onye na-akwalite mmechi stomatal n'okpuru nrụgide abiotic) (Macková et al., 2013; Zhou et al., 2013). 2013). , 2014). Mmepe stomatal na-eme ka akwụkwọ jụọ oyi ma na-enyere aka belata okpomọkụ mkpuchi (Sonjaroon et al., 2018; Quintero-Calderón et al., 2021). N'ihi ihe ndị a, okpomọkụ mkpuchi nke osisi osikapa a na-agba CK ma ọ bụ BR nwere ike ịdị ala n'okpuru nrụgide okpomọkụ ejikọtara ọnụ.
Nrụgide okpomọkụ dị elu nwere ike ibelata ọdịnaya agba fotosynthetic nke akwụkwọ (Chen et al., 2017; Ahammed et al., 2018). N'ọmụmụ ihe a, mgbe osisi osikapa nọ n'okpuru nrụgide okpomọkụ ma ghara ịgbasa ihe ọ bụla na-achịkwa uto osisi, agba photosynthetic na-ebelata na genotypes abụọ ahụ (Tebụl 2). Feng et al. (2013) kọkwara mbelata dị ukwuu na ọdịnaya chlorophyll na akwụkwọ nke genotypes abụọ nke wit na-enweta nrụgide okpomọkụ. Ikpughe na okpomọkụ dị elu na-ebutekarị mbelata ọdịnaya chlorophyll, nke nwere ike ịbụ n'ihi mbelata biosynthesis chlorophyll, mmebi nke pigments, ma ọ bụ mmetụta ha jikọtara ọnụ n'okpuru nrụgide okpomọkụ (Fahad et al., 2017). Agbanyeghị, osisi osikapa ejiri CK na BA gwọọ nke ukwuu mụbara mkpokọta nke pigments photosynthetic akwụkwọ n'okpuru nrụgide okpomọkụ. Jespersen na Huang (2015) na Suchsagunpanit et al. kọkwara nsonaazụ ndị yiri ya. (2015), onye hụrụ mmụba nke ọdịnaya chlorophyll akwụkwọ mgbe etinyere homonụ zeatin na epibrassinosteroid na bentgrass na osikapa nke nwere nrụgide okpomọkụ, n'otu n'otu. Nkọwa ezi uche dị na ya maka ihe mere CK na BR ji kwalite mmụba nke ọdịnaya chlorophyll akwụkwọ n'okpuru nrụgide okpomọkụ jikọtara ọnụ bụ na CK nwere ike ime ka mmalite nke mkpali na-aga n'ihu nke ndị na-akwalite ngosipụta (dịka onye na-akwalite senescence-activating promoter (SAG12) ma ọ bụ onye na-akwalite HSP18) dịkwuo mma ma belata mfu chlorophyll na akwụkwọ. , na-egbu oge ịka nká akwụkwọ ma na-eme ka iguzogide okpomọkụ dịkwuo elu (Liu et al., 2020). BR nwere ike ichebe chlorophyll akwụkwọ ma mụbaa ọdịnaya chlorophyll akwụkwọ site na ịgbalite ma ọ bụ ịkpalite njikọ nke enzymes metụtara biosynthesis chlorophyll n'okpuru ọnọdụ nrụgide (Sharma et al., 2017; Siddiqui et al., 2018). N'ikpeazụ, phytohormones abụọ (CK na BR) na-akwalitekwa ngosipụta nke protein ndị na-akpata okpomọkụ ma na-emeziwanye usoro mgbanwe metabolic dị iche iche, dịka mmụba nke biosynthesis chlorophyll (Sharma et al., 2017; Liu et al., 2020).
Paramita fluorescence nke Chlorophyll na-enye usoro ngwa ngwa na nke na-anaghị emebi emebi nke nwere ike inyocha nnabata osisi ma ọ bụ mgbanwe na ọnọdụ nrụgide abiotic (Chaerle et al. 2007; Kalaji et al. 2017). Ejirila paramita dịka oke Fv/Fm dị ka ihe ngosi nke mgbanwe osisi na ọnọdụ nrụgide (Alvarado-Sanabria et al. 2017; Chavez-Arias et al. 2020). N'ọmụmụ ihe a, osisi SC gosipụtara uru kachasị ala nke mgbanwe a, nke kachasị bụ osisi osikapa "F2000". Yin et al. (2010) chọpụtakwara na oke Fv/Fm nke akwụkwọ osikapa kachasị elu belatara nke ukwuu na okpomọkụ karịrị 35°C. Dịka Feng et al. (2013) si kwuo, obere oke Fv/Fm n'okpuru nrụgide okpomọkụ na-egosi na ọnụego njide ike mkpali na mgbanwe site na ebe mmeghachi omume PSII na-ebelata, na-egosi na ebe mmeghachi omume PSII na-agbaze n'okpuru nrụgide okpomọkụ. Ihe a a chọpụtara na-eme ka anyị kwubie na nsogbu dị na ngwa fotosynthesis na-apụta ìhè karịa na ụdị dị iche iche na-adịghị ike (Fedearroz 2000) karịa na ụdị dị iche iche na-eguzogide ọgwụ (Fedearroz 67).
Ojiji nke CK ma ọ bụ BR na-eme ka arụmọrụ PSII ka mma n'okpuru ọnọdụ nrụgide okpomọkụ dị mgbagwoju anya. Suchsagunpanit et al. (2015) nwetara ihe ndị yiri ya, bụ ndị hụrụ na itinye BR mụbara arụmọrụ nke PSII n'okpuru nrụgide okpomọkụ na osikapa. Kumar et al. (2020) chọpụtakwara na osisi chickpea ejiri CK (6-benzyladenine) gwọọ ma nwee nrụgide okpomọkụ mụbara oke Fv/Fm, na-ekwu na itinye foliar nke CK site na ịgbalite okirikiri pigment zeaxanthin kwalitere ọrụ PSII. Na mgbakwunye, ọgwụ fesa akwụkwọ BR kwadoro fotosynthesis PSII n'okpuru ọnọdụ nrụgide jikọtara ọnụ, na-egosi na itinye phytohormone a butere mbelata mgbasa nke ike mkpali nke antennae PSII ma kwalite mkpokọta obere protein ọkụ ọkụ na chloroplasts (Ogweno et al. 2008; Kothari na Lachowitz)., 2021).
Ọdịnaya MDA na proline na-abawanye mgbe osisi nọ n'okpuru nrụgide abiotic ma e jiri ya tụnyere osisi ndị a kụrụ n'okpuru ọnọdụ kacha mma (Alvarado-Sanabria et al. 2017). Ọmụmụ ihe ndị gara aga egosikwala na ọkwa MDA na proline bụ ihe ngosi biochemical nke enwere ike iji ghọta usoro mgbanwe ma ọ bụ mmetụta nke omume agronomic na osikapa n'okpuru okpomọkụ ehihie ma ọ bụ n'abalị (Alvarado-Sanabria et al., 2017; Quintero-Calderón et al.., 2021). Ọmụmụ ihe ndị a gosikwara na ọdịnaya MDA na proline na-adịkarị elu na osisi osikapa ndị a na-ekpughere na okpomọkụ dị elu n'abalị ma ọ bụ n'ehihie, n'otu n'otu. Agbanyeghị, ịgba CK na BR tinyere aka na mbelata na MDA na mmụba na ọkwa proline, ọkachasị na genotype na-anabata (Federroz 67). Ịgba CK nwere ike ịkwalite oke ngosipụta nke cytokinin oxidase/dehydrogenase, si otú a na-amụba ọdịnaya nke ihe nchebe dịka betaine na proline (Liu et al., 2020). BR na-akwalite mmụba nke osmoprotectants dịka betaine, shuga, na amino acid (gụnyere proline efu), na-ejigide nguzozi osmotic cellular n'okpuru ọtụtụ ọnọdụ gburugburu ebe obibi na-adịghị mma (Kothari na Lachowiec, 2021).
A na-eji ihe nrịbama nrụgide ihe ubi (CSI) na ihe nrịbama ndidi (RTI) chọpụta ma ọgwụgwọ ndị a na-enyocha na-enyere aka belata nrụgide dị iche iche (abiotic na biotic) ma nwee mmetụta dị mma na physiology osisi (Castro-Duque et al., 2020; Chavez-Arias et al., 2020). Ụkpụrụ CSI nwere ike ịdị site na 0 ruo 1, nke na-anọchite anya ọnọdụ ndị na-abụghị nrụgide na nrụgide, n'otu n'otu (Lee et al., 2010). Ụkpụrụ CSI nke osisi ndị nwere nrụgide okpomọkụ (SC) sitere na 0.8 ruo 0.9 (Foto 2B), na-egosi na nrụgide jikọtara ọnụ metụtara osisi osikapa. Agbanyeghị, ịgbasa akwụkwọ nke BC (0.6) ma ọ bụ CK (0.6) bụ isi butere mbelata na ihe ngosi a n'okpuru ọnọdụ nrụgide abiotic ma e jiri ya tụnyere osisi osikapa SC. N'ime osisi F2000, RTI gosiri mmụba dị elu mgbe ejiri CA (97.69%) na BC (60.73%) ma e jiri ya tụnyere SA (33.52%), nke na-egosi na ndị na-achịkwa uto osisi ndị a na-enyekwa aka ime ka mmeghachi omume osikapa dịkwuo mma na nnabata nke ihe mejupụtara ya. Okpomọkụ gabigara ókè. Atụpụtala ihe ngosi ndị a iji jikwaa ọnọdụ nrụgide n'ụdị dị iche iche. Nnyocha nke Lee et al. (2010) mere gosiri na CSI nke ụdị owu abụọ n'okpuru nrụgide mmiri dị ala dị ihe dị ka 0.85, ebe uru CSI nke ụdị owu a gbara mmiri nke ọma sitere na 0.4 ruo 0.6, na-ekwubi na ihe ngosi a bụ ihe na-egosi mgbanwe mmiri nke ụdị dị iche iche. Ọnọdụ nrụgide. Ọzọkwa, Chavez-Arias et al. (2020) tụlere irè nke ihe ndị na-akpalite ihe dị ka atụmatụ njikwa nrụgide zuru oke na osisi C. elegans wee chọpụta na osisi ndị a na-agbaze ihe ndị a gosipụtara RTI dị elu (65%). Dabere na ihe ndị dị n'elu, a pụrụ iwere CK na BR dị ka atụmatụ ọrụ ugbo nke ebumnuche ya bụ ịbawanye nnabata nke osikapa na nrụgide okpomọkụ dị mgbagwoju anya, ebe ọ bụ na ndị na-achịkwa uto osisi ndị a na-akpali mmeghachi omume biochemical na physiological dị mma.
N'ime afọ ole na ole gara aga, nnyocha osikapa na Colombia elekwasịla anya n'inyocha ụdị mkpụrụ ndụ ihe nketa ndị na-anagide okpomọkụ ehihie ma ọ bụ abalị site na iji àgwà physiological ma ọ bụ biochemical (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2021). Agbanyeghị, n'ime afọ ole na ole gara aga, nyocha nke teknụzụ bara uru, akụ na ụba na uru aghọwo ihe dị mkpa iji tụọ aro njikwa ihe ọkụkụ agbakwunyere iji melite mmetụta nke oge nrụgide okpomọkụ dị mgbagwoju anya na mba ahụ (Calderón-Páez et al., 2021; Quintero-Calderon et al., 2021). Ya mere, mmeghachi omume physiological na biochemical nke osisi osikapa na nrụgide okpomọkụ dị mgbagwoju anya (40°C ụbọchị/30°C abalị) a hụrụ na ọmụmụ ihe a na-egosi na ịgbasa foliar na CK ma ọ bụ BR nwere ike ịbụ ụzọ njikwa ihe ọkụkụ kwesịrị ekwesị iji belata mmetụta ọjọọ. Mmetụta nke oge nrụgide okpomọkụ dị obere. Ọgwụgwọ ndị a mere ka ndidi nke mkpụrụ ndụ ihe nketa osikapa (CSI dị ala na RTI dị elu) dịkwuo mma, na-egosi usoro izugbe na nzaghachi physiological na biochemical osisi n'okpuru nrụgide okpomọkụ jikọtara ọnụ. Isi ihe ndị sitere n'osisi osikapa bụ mbelata na ọdịnaya GC, mkpokọta chlorophyll, chlorophylls α na β na carotenoids. Na mgbakwunye, osisi na-enwe mmebi PSII (ebelatara paramita fluorescence chlorophyll dịka Fv/Fm ratio) na mmụba nke lipid peroxidation. N'aka nke ọzọ, mgbe ejiri CK na BR gwọọ osikapa, mmetụta ọjọọ ndị a belatara ma ọdịnaya proline mụbara (Foto 4).
Foto nke 4. Ihe nlereanya echiche nke mmetụta nke nrụgide okpomọkụ jikọtara ọnụ na ihe na-achịkwa uto osisi foliar na osisi osikapa. Akụ uhie na acha anụnụ anụnụ na-egosi mmetụta ọjọọ ma ọ bụ dị mma nke mmekọrịta dị n'etiti nrụgide okpomọkụ na itinye foliar nke BR (brassinosteroid) na CK (cytokinin) na nzaghachi physiological na biochemical, n'otu n'otu. gs: stomatal conductance; Ngụkọta Cl: mkpokọta ọdịnaya chlorophyll; Chl α: ọdịnaya chlorophyll β; Cx+c: ọdịnaya carotenoid;
Na nchịkọta, mmeghachi omume nke anụ ahụ na biochemical na ọmụmụ ihe a na-egosi na osisi osikapa Fedearroz 2000 nwere ike ịnwe oge nrụgide okpomọkụ dị mgbagwoju anya karịa osisi osikapa Fedearroz 67. Ndị niile na-achịkwa uto n'ọmụmụ ihe a (auxins, gibberellins, cytokinins, ma ọ bụ brassinosteroids) gosipụtara ụfọdụ oke mbelata nrụgide okpomọkụ. Agbanyeghị, cytokinin na brassinosteroids kpalitere mgbanwe osisi ka mma ka ndị na-achịkwa uto osisi na-amụba ọdịnaya chlorophyll, paramita fluorescence alpha-chlorophyll, gs na RWC ma e jiri ya tụnyere osisi osikapa na-enweghị ngwa ọ bụla, ma belatakwa ọdịnaya MDA na okpomọkụ mkpuchi. Na nchịkọta, anyị na-ekwubi na ojiji nke ndị na-achịkwa uto osisi (cytokinins na brassinosteroids) bụ ngwaọrụ bara uru n'ịchịkwa ọnọdụ nrụgide na ihe ọkụkụ osikapa nke nrụgide okpomọkụ siri ike kpatara n'oge okpomọkụ dị elu.
E tinyere ihe ndị e dere n'ọmụmụ ihe ahụ n'isiokwu ahụ, a pụkwara ịjụ ajụjụ ndị ọzọ nye onye dere ya.