Wax soo saarka bariiska ayaa hoos u dhacaya sababtoo ah isbeddelka cimilada iyo kala duwanaanshaha Colombia.Xakamaynta koritaanka dhirtaayaa loo isticmaalay sidii xeelad si loo yareeyo diiqada kulaylka ee dalagyada kala duwan. Sidaa darteed, ujeeddada daraasaddan waxay ahayd in la qiimeeyo saameynta jireed (dhaqdhaqaaqa stomatal, habdhaqanka stomatal, wadarta chlorophyll, saamiga Fv / Fm ee labada genotypes bariiska ganacsiga ah ee lagu soo rogay isku-darka kulaylka (heerkulka habeenka iyo maalinta sare), heerkulka canopy iyo heerka biyaha qaraabo) iyo doorsoomayaasha biochemical (malondialdehyde (MDA)) iyo prolinic acid). Tijaabooyinka koowaad iyo labaad ayaa la sameeyay iyadoo la adeegsanayo dhirta laba genotypes bariiska Federrose 67 ("F67") iyo Federrose 2000 ("F2000"), siday u kala horreeyaan. Labada tijaabo ayaa si wada jir ah loo falanqeeyay sidii tijaabo taxane ah. Daawaynta la aasaasay waxay ahayd sida soo socota: xakamaynta buuxda (AC) (dhirta bariiska oo lagu beeray heerkul aad u fiican (heerkulka maalinta / habeenka 30/25 ° C)), xakamaynta cadaadiska kulaylka (SC) 30°C)], iyo dhirta bariiska ayaa la adkeeyay waxaana lagu buufiyay nidaamiyayaal koritaanka dhirta (stress+AUX, stress+BR, stress+CK ama stress+GA) laba jeer (5 maalmood ka hor iyo 5 maalmood kadib diiqada kulaylka). Ku buufinta SA waxay kordhisay wadarta chlorophyll ee labada noocba (miisaanka cusub ee dhirta bariiska "F67" iyo "F2000" waxay ahayd 3.25 iyo 3.65 mg/g, siday u kala horreeyaan) marka la barbar dhigo dhirta SC (miisaanka cusub ee "F67" dhirta wuxuu ahaa 2.36 iyo 2.56 mg 0" bariis "2 F) g-1. CK waxa kale oo ay guud ahaan wanaajisay dhaqdhaqaaqa caloosha ee bariiska “F2000” dhirta (499.25 vs. 150.60 mmol m-2 s) marka la barbar dhigo xakamaynta cadaadiska kulaylka. Cadaadiska kulaylka, heerkulka taajka dhirta ayaa hoos u dhacaya 2-3 °C, iyo waxa ku jira MDA ee dhirta ayaa hoos u dhacaya. Tusaha dulqaadka qaraabada ayaa muujinaya in codsiga foliar ee CK (97.69%) iyo BR (60.73%) ay kaa caawin karaan yaraynta dhibaatada kulaylka la isku daray. stress inta badan ee dhirta bariiska F2000. Gabagabadii, buufinta foliar ee BR ama CK waxaa loo tixgelin karaa inay tahay istaraatiijiyad beeraley ah si loo yareeyo saameynta xun ee isku-darka xaaladaha kuleylka kulaylka ee habdhaqanka jireed ee dhirta bariiska.
Bariiska (Oryza sativa) waxaa iska leh qoyska Poaceae waana mid ka mid ah badarka ugu badan ee la beero aduunka oo ay la socoto galley iyo qamadi (Bajaj and Mohanty, 2005). Aagga lagu beero bariiska waa 617,934 hektar, wax soo saarka qaranka 2020-kii wuxuu ahaa 2,937,840 tan oo leh celcelis ahaan waxsoosaar dhan 5.02 tan/ha (Federarroz (Federación Nacional de Arroceros), 2021).
Kulaylka caalamiga ah wuxuu saameeyaa dalagyada bariiska, taasoo keentay noocyada kala duwan ee walaaca abiotic sida heerkulka sare iyo xilliyada abaarta. Isbeddelka cimiladu waxay keenaysaa in heerkulka caalamiga ahi kor u kaco; Heerkulka ayaa la saadaalinayaa inuu kor u kaco 1.0-3.7 ° C qarniga 21aad, taas oo kordhin karta inta jeer iyo xoojinta cadaadiska kulaylka. Kordhinta heerkulka deegaanka ayaa saameeyay bariiska, taasoo keentay in wax-soo-saarka dalagga uu hoos u dhaco 6-7% . Dhanka kale, isbeddelka cimiladu waxay sidoo kale keentaa xaalado deegaan oo aan wanaagsanayn dalagyada, sida xilliyada abaarta daran ama kulaylka sare ee gobollada kulaylaha iyo kuwa hoose. Intaa waxaa dheer, dhacdooyinka kala duwanaanshiyaha sida El Niño waxay u horseedi kartaa diiqada kulaylka waxayna ka sii dari kartaa dhaawaca dalagga ee gobollada kulaylaha qaarkood. Kolombiya, heerkulka meelaha bariiska laga soo saaro ayaa la saadaalinayaa inuu kordho 2-2.5°C sanadka 2050, taasoo hoos u dhigaysa wax soo saarka bariiska oo saamaynaysa isu socodka badeecadaha suuqyada iyo silsiladaha sahayda.
Inta badan dalagyada bariiska waxaa lagu beeraa meelaha heerkulku ku dhow yahay heerka ugu wanaagsan ee koritaanka dalagga (Shah et al., 2011). Waxaa la soo sheegay in celceliska ugu fiican habeen iyo maalinba heerkulkakoritaanka iyo horumarka bariiskaguud ahaan waa 28°C iyo 22°C, siday u kala horreeyaan (Kilasi et al., 2018; Calderón-Páez et al., 2021). Heerkulka ka sarreeya marinnadan waxay sababi karaan xilliyo dhexdhexaad ah ama diiqad kulul oo kulul inta lagu jiro marxaladaha xasaasiga ah ee horumarinta bariiska (calaamadaha, suuxinta, ubaxa, iyo buuxinta hadhuudhka), taas oo si xun u saameeya wax-soo-saarka badarka. Hoos-u-dhacan wax-soo-saarka ayaa inta badan sabab u ah muddada dheer ee cadaadiska kulaylka, kaas oo saameeya physiology dhirta. Sababtoo ah isdhexgalka arrimo kala duwan, sida muddada cadaadiska iyo heerkulka ugu sarreeya ee la gaarey, diiqada kulaylku waxay sababi kartaa waxyeello kala duwan oo aan laga beddeli karin dheef-shiid kiimikaadka iyo horumarinta dhirta.
Cadaadiska kulaylku wuxuu saameeyaa hababka kala duwan ee jireed iyo biochemical ee dhirta. Photosynthesis caleentu waa mid ka mid ah hababka ugu nugul kulaylka dhirta bariiska, maadaama heerka photosynthesis uu hoos u dhaco 50% marka heerkulku maalin kasta dhaafo 35°C. Jawaabaha jireed ee dhirta bariiska way kala duwan yihiin iyadoo ku xiran nooca diiqada kulaylka. Tusaale ahaan, heerarka photosynthetic iyo dhaqdhaqaaqa stomatal waa la joojiyaa marka dhirta ay la kulmaan heerkul sare oo maalintii ah (33-40 ° C) ama heerkul sarreeya maalintii iyo habeenkii (35-40 ° C maalintii, 28-30 ° C). C macnaheedu waa habeen) (Lü et al., 2013; Fahad et al., 2016; Chaturvedi et al., 2017). Heerkulka habeenkii oo sarreeya (30°C) wuxuu keenaa xannibaad dhexdhexaad ah ee sawir-qaadista laakiin wuxuu kordhiyaa neefsiga habeenkii (Fahad et al., 2016; Alvarado-Sanabria et al., 2017). Iyadoo aan loo eegin xilliga walaaca, diiqada kulaylku waxay sidoo kale saamaysaa nuxurka chlorophyll caleen, saamiga chlorophyll variable fluorescence ilaa ugu badnaan chlorophyll fluorescence (Fv/Fm), iyo firfircoonida Rubisco ee dhirta bariiska (Cao et al. 2009; Yin et al. 2010). ) Sanchez Reynoso iyo al., 2014).
Isbeddellada kiimikaadku waa dhinac kale oo la qabsiga dhirta ee diiqada kulaylka (Wahid et al., 2007). Maadada Proline waxa loo isticmaalay tilmaame bayoolaji ah oo walaaca dhirta (Ahmed and Hassan 2011). Proline waxay door muhiim ah ka ciyaartaa dheef-shiid kiimikaadka dhirta sida ay u shaqeyso sida kaarboon ama ilo nitrogen iyo sida xuubka xajinta ee xaaladaha heerkulka sare (Sánchez-Reinoso et al., 2014). Heerkulka sare wuxuu kaloo saameeyaa xasilloonida xuubka iyada oo loo marayo lipid peroxidation, taasoo horseedaysa samaynta malondialdehyde (MDA) (Wahid et al., 2007). Sidaa darteed, maadada MDA ayaa sidoo kale loo isticmaalay si loo fahmo qaab-dhismeedka qaabdhismeedka xuubka unugyada ee cadaadiska kulaylka (Cao et al., 2009; Chavez-Arias et al., 2018). Ugu dambeyntii, isku-darka kuleylka kuleylka [37/30 ° C (maalin/habeen)] waxay kordhisay boqolleyda daadinta korantada iyo maadada malondialdehyde ee bariiska (Liu et al., 2013).
Isticmaalka xakamaynta koritaanka dhirta (GRs) ayaa la qiimeeyay si loo yareeyo saameynta xun ee cadaadiska kulaylka, maaddaama walxahan ay si firfircoon ugu lug leeyihiin jawaabaha dhirta ama hababka difaaca jireed ee ka soo horjeeda walaaca (Peleg iyo Blumwald, 2011; Yin et al. et al., 2011; Ahmed et al., 2015). Codsiga dibadda ee kheyraadka hidde-sidaha ayaa saameyn togan ku yeeshay dulqaadka kuleylka kulaylka ee dalagyada kala duwan. Cilmi-baaristu waxay muujisay in phytohormones sida gibberellins (GA), cytokinins (CK), auxins (AUX) ama brassinosteroids (BR) ay horseedaan koror ku yimaada doorsoomayaasha jireed iyo kiimikaad ee kala duwan (Peleg iyo Blumwald, 2011; Yin et al. Ren, 2011; Mitler et al., 2001 et al., 200). Kolombiya, adeegsiga kheyraadka hidde-sidaha iyo saamaynta ay ku leedahay dalagyada bariiska si buuxda looma fahmin lana darsin. Si kastaba ha ahaatee, daraasad hore ayaa muujisay in buufinta foliar ee BR ay hagaajin karto dulqaadka bariiska iyada oo la hagaajinayo sifooyinka gaaska sarrifka, chlorophyll ama proline ee caleemaha seedling bariiska (Quintero-Calderón et al., 2021).
Cytokinins waxay dhexdhexaadiyaan jawaabaha dhirta ee cadaadiska abiotic, oo ay ku jiraan diiqada kulaylka (Ha et al., 2012). Intaa waxaa dheer, waxaa la soo sheegay in codsiga dibadda ee CK uu yarayn karo dhaawaca kulaylka. Tusaale ahaan, codsiga exogenous ee zeatin waxay kordhisay heerka photosynthetic, chlorophyll a iyo b content, iyo hufnaanta gaadiidka elektaroonigga ah ee gurguurta bentgrass (Agrotis estolonifera) inta lagu jiro diiqada kulaylka (Xu iyo Huang, 2009; Jespersen iyo Huang, 2015). Codsiga dibadda ee zeatin wuxuu sidoo kale wanaajin karaa dhaqdhaqaaqa antioxidant, kor u qaadida isku-dhafka borotiinnada kala duwan, wuxuu yareeyaa waxyeelada noocyada oksijiinta firfircoon (ROS) iyo wax soo saarka malondialdehyde (MDA) ee unugyada dhirta (Chernyadyev, 2009; Yang et al., 2009). , 2016; Kumar iyo al., 2020).
Isticmaalka gibberellic acid ayaa sidoo kale muujisay jawaab celin wanaagsan oo ku saabsan diiqada kuleylka. Daraasaduhu waxay muujiyeen in GA biosynthesis ay dhexdhexaadiso waddooyinka dheef-shiid kiimikaadka kala duwan waxayna kordhisaa dulqaadka xaaladaha heerkulka sare (Alonso-Ramirez et al. 2009; Khan et al. 2020). Abdel-Nabi iyo al. (2020) waxay ogaatay in buufinta foliar ee exogenous GA (25 ama 50 mg * L) ay kordhin karto heerka photosynthetic iyo waxqabadka antioxidant ee dhirta orange-ku-cadawsan marka la barbar dhigo dhirta la xakameeyo. Waxa kale oo la arkay in codsiga ka baxsan ee HA ay kordhiso qoyaanka qaraabada ah, chlorophyll iyo carotenoid ka kooban oo ay yarayso peroxidation lipid ee timirta timirta (Phoenix dactylifera) ee cadaadiska kulaylka (Khan et al., 2020). Auxin sidoo kale waxay door muhiim ah ka ciyaartaa nidaaminta jawaabaha koritaanka la qabsiga ee xaaladaha heerkulka sare (Sun et al., 2012; Wang et al., 2016). Nidaamiyaha korriinka wuxuu u shaqeeyaa sida calaamadaynta biochemical ee habab kala duwan sida proline synthesis ama hoos u dhaca cadaadiska abiotic (Ali et al. 2007). Intaa waxaa dheer, AUX waxay sidoo kale kor u qaadaysaa dhaqdhaqaaqa antioxidant, taas oo horseedaysa hoos u dhaca MDA ee dhirta sababtoo ah hoos u dhaca peroxidation lipid (Bielach et al., 2017). Sergeev iyo al. (2018) ayaa lagu arkay in dhirta digirta (Pisum sativum) ee cadaadiska kulaylka, waxa ku jira proline - dimethylaminoethoxycarbonylmethyl) naphthylchloromethyl ether (TA-14) ayaa kordhay. Isla tijaabadii, waxay sidoo kale arkeen heerarka hoose ee MDA ee dhirta la daweeyay marka la barbar dhigo dhirta aan lagu dawayn AUX.
Brassinosteroids waa nooc kale oo xakameynaya korriinka oo loo isticmaalo in lagu yareeyo saameynta kuleylka kulaylka. Ogweno et al. (2008) ayaa sheegay in buufinta BR exogenous ay kordhisay heerka sawir-qaadista saafiga ah, dhaqdhaqaaqa stomatal iyo heerka ugu sarreeya ee Rubisco karboksylation ee yaanyada (Solanum lycopersicum) ee dhirta cadaadiska kuleylka 8 maalmood. Ku buufinta foliar ee epibrassinosteroids waxay kordhin kartaa qiyaasta saafiga ah ee sawir-qaadista ee dhirta qajaarka (Cucumis sativus) ee hoos yimaada cadaadiska kulaylka (Yu et al., 2004). Intaa waxaa dheer, codsiga qalaad ee BR wuxuu dib u dhigayaa hoos u dhaca chlorophyll wuxuuna kordhiyaa waxtarka isticmaalka biyaha iyo wax-soosaarka tirada ugu badan ee PSII photochemistry ee dhirta ku hoos jira diiqada kuleylka (Holá et al., 2010; Toussagunpanit et al., 2015).
Isbeddelka cimilada iyo kala duwanaanshiyaha awgeed, dalagyada bariiska ayaa la kulma xilliyo heerkul maalinle ah oo sarreeya (Lesk et al., 2016; Garcés, 2020; Federaroz (Federación Nacional de Arroceros), 2021). In phenotyping dhirta, isticmaalka phytonutrients ama biostimulants ayaa loo bartay sidii istaraatijiyad ah si loo yareeyo cadaadiska kulaylka ee meelaha bariiska-koritaanka (Alvarado-Sanabria et al., 2017; Calderón-Páez et al., 2021; Quintero-Calderón et al., 2021). Intaa waxaa dheer, isticmaalka biochemical iyo doorsoomayaal jireed (heerkulka caleen, habdhaqanka stomatal, chlorophyll fluorescence xuduudaha, chlorophyll iyo qaraabada biyaha, malondialdehyde iyo proline synthesis) waa qalab lagu kalsoonaan karo oo lagu baadho dhirta bariiska ee cadaadiska kulaylka gudaha iyo caalamiga ah (Sánchez -Reynoso et al., 2014; Si kastaba ha ahaatee, cilmi-baaris ku saabsan isticmaalka buufinta foliar phytohormonal ee bariiska ee heerka maxalliga ah ayaa weli ah naadir. cabbirada iyo nuxurka biyaha qaraabo ah) iyo saamaynta biochemical ee codsiga foliar ee afar nidaamiya koritaanka dhirta (AUX, CK, GA iyo BR). (Photosynthetic pigments, malondialdehyde iyo proline ka kooban yahay) Kala duwanaansho laba nooc oo bariis ganacsi ah oo ay la kulmeen cadaadis kulayl oo isku jira (heerkulka maalinta/habeenka oo sarreeya).
Daraasaddan, laba tijaabo oo madaxbannaan ayaa la sameeyay. Genotypes Federrose 67 (F67: genotype oo lagu sameeyay heerkul sare tobankii sano ee la soo dhaafay) iyo Federrose 2000 (F2000: genotype oo la sameeyay tobankii sano ee la soo dhaafay ee qarnigii 20aad oo muujinaya iska caabbinta fayraska caleen cad) ayaa la isticmaalay markii ugu horeysay. abuur. iyo tijaabadii labaad, siday u kala horreeyaan. Labada genotypes waxaa si weyn u beera beeralayda Colombia. Seeraha waxaa lagu beeray saxaarad 10-L ah (dhererka 39.6 cm, ballac 28.8 cm, joog 16.8 cm) oo ka kooban carro ciid ciid leh oo leh 2% walxo organic. Shan iniin hore u biqlay ayaa lagu beeray saxaarad kasta. Pallets-yada waxaa la dhigay aqalka dhirta lagu koriyo ee Kulliyada Sayniska Beeraha ee Jaamacadda Qaranka ee Kolombiya, Kambaska Bogotá (43°50′56″ N, 74°04′051″ W), oo joog 2556 m ka sarraysa heerka badda (asl). m.) waxaana la fuliyay Oktoobar ilaa Diseembar 2019. Hal tijaabo (Federroz 67) iyo tijaabo labaad (Federroz 2000) isla xilli ciyaareedka 2020.
Xaaladaha deegaanka ee aqalka dhirta lagu koriyo inta lagu jiro xilli kasta oo beerto waa sida soo socota: maalintii iyo habeenkii heerkulka 30/25 ° C, qoyaan qaraabo 60 ~ 80%, photoperiod dabiiciga ah 12 saacadood (photosynthetically firfircoon shucaaca 1500 µmol (photons) m-2 s-). 1 duhurnimo). Dhirta ayaa la bacrimiyey iyadoo loo eegayo nuxurka cunsur kasta 20 maalmood kadib soo bixitaanka abuurka (DAE), sida uu qabo Sánchez-Reinoso et al. (2019): 670 mg nitrogen halkii geed, 110 mg fosfooraska halkii geed, 350 mg potassium halkii geed, 68 mg calcium halkii geed, 20 mg magnesium warshad kasta, 20 mg baaruud halkii geed, 17 mg silicon halkii geed. Dhirtu waxay ka kooban tahay 10 mg boron geedkiiba, 17 mg naxaas geedkiiba, iyo 44 mg zinc geedkiiba. Dhirta bariiska ayaa lagu dayactiray ilaa 47 DAE tijaabo kasta markii ay gaadheen heerka phenological V5 muddadan. Daraasadihii hore waxay muujiyeen in marxaladdan phenological ay tahay wakhti ku habboon in lagu sameeyo daraasadaha cadaadiska kulaylka ee bariiska (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2017).
Tijaabo kasta, laba codsi oo kala duwan oo maamulaha koritaanka caleen ayaa la sameeyay. Qaybta ugu horreysa ee buufinta foliar phytohormone ayaa lagu dabaqay 5 maalmood ka hor daaweynta kuleylka kuleylka (42 DAE) si loogu diyaariyo dhirta walaaca deegaanka. Buufin labaad oo foliar ah ayaa markaa la siiyay 5 maalmood ka dib markii dhirtu ay soo gaadhay xaaladaha walaaca (52 DAE). Afar phytohormones ayaa la isticmaalay iyo sifooyinka walxo kasta oo firfircoon oo lagu buufiyo daraasaddan waxay ku taxan yihiin Shaxda Dheeraadka ah 1. Xakamaynta xakamaynta koritaanka caleenta ee loo isticmaalo waxay ahaayeen sida soo socota: (i) Auxin (1-naphthylacetic acid: NAA) oo ah 5 × 10-5 M (ii) 5 × 10-5 M gibberelin; GA3); (iii) Cytokinin (trans-zeatin) 1 × 10-5 M (iv) Brassinosteroids [Spirostan-6-one, 3,5-dihydroxy-, (3b,5a,25R)] 5 × 10-5; M. Isku-dhafkan ayaa la doortay sababtoo ah waxay keenaan jawaab celin wanaagsan waxayna kordhiyaan caabbinta dhirta ee cadaadiska kuleylka (Zahir et al., 2001; Wen et al., 2010; El-Bassiony et al., 2012; Salehifar et al., 2017). Dhirta bariiska iyada oo aan lahayn buufinta xakamaynta koritaanka dhirta ayaa lagu daaweeyay biyo nadiif ah oo keliya. Dhammaan dhirta bariiska ayaa lagu buufiyay gacan-buufiye. Codso 20 ml H2O geedka si aad u qoyso dusha sare iyo hoose ee caleemaha. Dhammaan buufinta foliar waxay isticmaaleen adjuvant-ka beeraha (Agrotin, Bayer CropScience, Colombia) 0.1% (v/v). Masaafada u dhaxaysa dheriga iyo buufiyehu waa 30 cm.
Daaweynta cadaadiska kulaylka ayaa la maamulay 5 maalmood ka dib buufinta foliar ee ugu horreysa (47 DAE) ee tijaabo kasta. Dhirta bariiska ayaa laga wareejiyay aqalka dhirta lagu koriyo ilaa 294 L qol korriin (MLR-351H, Sanyo, IL, USA) si loo dhiso stress kulaylka ama loo ilaaliyo xaaladaha deegaanka la mid ah (47 DAE). Daawaynta isku-dhafka kulaylka waxaa lagu sameeyay iyadoo qolka la dhigayo heerkulka maalinta/habeenka soo socda: heerkulka maalintii oo sarreeya [40°C 5 saacadood (laga bilaabo 11:00 ilaa 16:00)] iyo xilliga habeenkii [30°C ilaa 5 saacadood]. 8 maalmood oo isku xigta (laga bilaabo 19:00 ilaa 24:00). Heerkulka cadaadiska iyo wakhtiga soo-gaadhista ayaa lagu doortay iyadoo lagu saleynayo daraasadihii hore (Sánchez-Reynoso et al. 2014; Alvarado-Sanabría et al. 2017). Dhanka kale, koox ka mid ah dhirta loo wareejiyay qolka koritaanka ayaa lagu hayaa aqalka dhirta lagu koriyo isla heerkulka (30 ° C maalintii / 25 ° C habeenkii) 8 maalmood oo isku xigta.
Dhammaadka tijaabada, kooxaha daaweynta ee soo socda ayaa la helay: (i) xaaladda heerkulka koritaanka + codsiga biyaha nadiifka ah [xakamaynta saxda ah (AC)], (ii) xaalada kulaylka kuleylka + codsiga biyaha diirran brassinosteroid (BR) Lifaaqa. Kooxahan daawaynta waxa loo isticmaalay laba genotypes (F67 iyo F2000). Dhammaan daawaynta waxaa lagu fuliyay naqshad gebi ahaanba randomized oo leh shan nuqul, mid kasta oo ka kooban hal geed. Geed kasta waxaa loo isticmaalay in lagu akhriyo doorsoomayaasha lagu go'aamiyay dhamaadka tijaabada. Tijaabadu waxay socotay 55 DAE.
Dhaqdhaqaaqa caloosha (gs) waxaa lagu cabiray iyadoo la adeegsanayo porosometer la qaadi karo (SC-1, METER Group Inc., USA) oo u dhaxaysa 0 ilaa 1000 mmol m-2 s-1, oo leh muunad qolka muunad ah oo ah 6.35 mm. Cabbiraadaha waxaa lagu qaadaa iyadoo lagu dhejiyo baaritaanka stomameter caleen baaluq leh oo birta ugu weyn ee geedka si buuxda loo ballaariyay. Daawaynta kasta, akhrinta gs ayaa lagu qaaday saddex caleemood oo geed kasta ah inta u dhaxaysa 11:00 iyo 16:00 iyo celcelis ahaan.
RWC waxaa loo go'aamiyay iyadoo loo eegayo habka lagu sifeeyay Ghoulam et al. (2002). Xaashida si buuxda loo balaariyay ee loo isticmaalay in lagu go'aamiyo g ayaa sidoo kale loo isticmaalay in lagu cabbiro RWC. Miisaanka cusub (FW) ayaa la go'aamiyay isla markiiba goosashada ka dib iyadoo la isticmaalayo cabbir dhijitaal ah. Caleemaha ayaa markaa lagu riday weel caag ah oo biyo ka buuxaan waxaana looga tagay mugdi heerkulka qolka (22°C) ilaa 48 saacadood. Ka dibna ku miisaam miisaan dhijitaal ah oo qor miisaanka la balaariyay (TW). Caleemaha bararsan waxaa lagu qallajiyey foornada 75 ° C 48 saacadood waxaana la duubay miisaankooda qallalan (DW).
Waxyaabaha klorophyll-ka qaraabada ah waxaa lagu go'aamiyay iyadoo la isticmaalayo mitirka chlorophyll (atLeafmeter, FT Green LLC, USA) oo lagu muujiyay cutubyada atLeaf (Dey et al., 2016). Akhriska waxtarka ugu badnaan ee PSII (saamiga Fv/Fm) ayaa la duubay iyadoo la isticmaalayo kicinta joogtada ah ee chlorophyll fluorimeter (Handy PEA, Hansatech Instruments, UK). Caleemaha waxay ahaayeen kuwo la qabsaday mugdi iyadoo la adeegsanayo isku-xidhka caleenta 20 daqiiqo kahor cabbirada Fv/Fm (Restrepo-Diaz iyo Garces-Varon, 2013). Ka dib markii caleemaha mugdiga ku jiraan, saldhigga (F0) iyo fluorescence ugu badnaan (Fm) ayaa la cabbiray. Laga soo bilaabo xogtan, fluorescence variable (Fv = Fm - F0), saamiga fluorescence variable ilaa fluorescence ugu badan (Fv/Fm), soosaarka tirada ugu badan ee PSII photochemistry (Fv/F0) iyo saamiga Fm/F0 ayaa la xisaabiyay (Baker, 2008; Lee et al., 2017). chlorophyll qaraabo ah iyo chlorophyll fluorescence akhrintiisa ayaa lagu qaaday caleemo isku mid ah oo loo isticmaalo cabbirada gs.
Qiyaastii 800 mg oo caleen miisaan cusub ah ayaa la soo ururiyay sida doorsoomayaasha biochemical. Shaybaarada caleenta ayaa markaa isku mid lagu daray nitrogen dareere waxaana lagu kaydiyay falanqayn dheeraad ah. Habka spectrometric ee loo isticmaalo in lagu qiyaaso chlorophyll a, b iyo carotenoid ka kooban yahay waxay ku salaysan tahay habka iyo isla'egyada uu qeexay Wellburn (1994). Shaybaarada unugyada caleenta (30 mg) ayaa la ururiyay oo la isku daray 3 ml oo ah 80% acetone. Shaybaarada ayaa markaa la saaray (qaabka 420101, Becton Dickinson Diagnostics Care Primary, USA) at 5000 rpm 10 daqiiqo si looga saaro qaybaha. Ka sarreeya waxa lagu qasi jiray mugga ugu dambeeya ee 6 ml iyadoo lagu daray 80% acetone (Sims iyo Gamon, 2002). Waxa ku jira chlorophyll waxaa lagu go'aamiyay 663 (chlorophyll a) iyo 646 (chlorophyll b) nm, iyo carotenoids 470 nm iyadoo la isticmaalayo spectrophotometer (Spectronic BioMate 3 UV-vis, Thermo, USA).
Habka thiobarbituric acid (TBA) ee lagu sharaxay Hodges et al. (1999) ayaa loo isticmaalay si loo qiimeeyo xuubka lipid peroxidation (MDA). Qiyaastii 0.3 g oo unug caleen ah ayaa sidoo kale isku mid ah nitrogen dareere ah. Tijaabooyinku waxa lagu qiyaasaa 5000 rpm, nuugista waxa lagu cabiray spectrophotometer 440, 532 iyo 600 nm. Ugu dambeyntii, uruurinta MDA waxaa la xisaabiyay iyadoo la adeegsanayo isku-dheellitirka baabi'inta (157 Mml-1).
Mawduuca proline ee dhammaan daawaynta ayaa lagu go'aamiyay iyadoo la adeegsanayo habka lagu qeexay Bates et al. (1973). Ku dar 10 ml oo ah 3% xal aqueous ah oo ah sulfosalicylic acid muunada la kaydiyay oo ka miir warqada Whatman filter (No. 2). Kadib 2 ml oo sifayntan ah ayaa lagu falceliyay 2 ml oo ah ninhydric acid iyo 2 ml oo ah glacial acetic acid. Isku darka waxaa lagu dhejiyay qubeyska biyaha 90 ° C 1 saac. Jooji falcelinta adiga oo barafka ku dhex shubaya. Si xoog leh u rux tuubada adigoo isticmaalaya ruxruxiyaha vortexka oo ku milan xalka ka soo baxay 4 ml oo toluene ah. Aqrinta nuugista waxaa lagu go'aamiyay 520 nm iyadoo la adeegsanayo spectrophotometer la mid ah oo loo adeegsaday qiyaasida pigments sawir-qaadista (Spectronic BioMate 3 UV-Vis, Thermo, Madison, WI, USA).
Habka uu tilmaamay Gerhards et al. (2016) si loo xisaabiyo heerkulka saqafka iyo CSI. Sawirada kulaylka waxaa lagu qaaday kamarad FLIR 2 (FLIR Systems Inc., Boston, MA, USA) oo sax ah ± 2 ° C dhamaadka xilliga walaaca. Dhig dusha cad gadaashiisa geedka si sawir looga qaado. Mar labaad, laba warshadood ayaa loo tixgeliyey moodallo tixraac ah. Dhirta ayaa la dhigay meel cad; mid waxaa lagu dahaadhay adjuvant-beeraha (Agrotin, Bayer CropScience, Bogotá, Colombia) si ay u ekaato furitaanka dhammaan stomata [qaabka qoyan (Twet)], kan kalena wuxuu ahaa caleen aan lahayn wax codsi ah [Qalab qalalan (Tdry)] (Castro -Duque et al., 2020). Masaafada u dhaxaysa kamarada iyo dheriga inta lagu jiro duubista waxay ahayd 1 m.
Tusmada dulqaadka qaraabada waxaa loo xisaabiyay si aan toos ahayn iyadoo la adeegsanayo stomatal conductance (gs) ee dhirta la daweeyay marka la barbar dhigo dhirta la xakameeyo (dhirta aan lahayn daawaynta walaaca iyo habeeyayaasha korriinka ee la dabaqay) si loo go'aamiyo dulqaadka genotypes-ka la daweeyay ee lagu qiimeeyay daraasaddan. RTI waxaa lagu helay iyadoo la adeegsanayo isla'egta laga soo xigtay Chavez-Arias et al. (2020).
Tijaabo kasta, dhammaan doorsoomayaasha jireed ee aan kor ku soo sheegnay ayaa la go'aamiyay waxaana lagu duubay 55 DAE iyadoo la adeegsanayo caleemo si buuxda loo ballaariyay oo laga soo ururiyay dusha sare. Intaa waxaa dheer, cabbirada ayaa lagu sameeyay qolka koritaanka si looga fogaado in la beddelo xaaladaha deegaanka ee dhirta ay ku koraan.
Xogta laga helay tijaabadii koowaad iyo labaad ayaa si wada jir ah loo falanqeeyay sidii tijaabo taxane ah. Koox kasta oo tijaabo ah waxay ka koobnayd 5 dhir, geed walbana wuxuu ka dhigay unug tijaabo ah. Falanqaynta kala duwanaanshaha (ANOVA) ayaa la sameeyay (P ≤ 0.05). Markii la ogaaday kala duwanaansho weyn, Tukey's post hoc test comparative test waxaa loo adeegsaday P ≤ 0.05. Isticmaal shaqada arcsine si aad u bedesho qiimaha boqolleyda. Xogta ayaa lagu falanqeeyay iyada oo la adeegsanayo barnaamijka Statisix v 9.0 (Software Analytical, Tallahassee, FL, USA) waxaana la qorsheeyay iyadoo la adeegsanayo SigmaPlot (nooca 10.0; Systat Software, San Jose, CA, USA). Falanqaynta qaybta ugu muhiimsan waxaa lagu fuliyay iyada oo la adeegsanayo barnaamijka InfoStat 2016 (Software Analysis, Jaamacadda Qaranka ee Cordoba, Argentina) si loo aqoonsado xeeriyeyaasha ugu wanaagsan ee koritaanka dhirta ee daraasadda.
Shaxda 1 waxay soo koobaysaa ANOVA oo muujinaysa tijaabooyinka, daawaynta kala duwan, iyo is dhexgalka ay la leeyihiin midabada sawir-qaadista caleenta (chlorophyll a, b, total, iyo carotenoids), malondialdehyde (MDA) iyo proline content, iyo stomatal conducting. Saamaynta gs, nuxurka biyaha ku jira. (RWC), maadada chlorophyll, cabirrada chlorophyll alpha fluorescence, heerkulka taajka (PCT) (°C), index stress dalagga (CSI) iyo tusmada dulqaadka qaraabada ee dhirta bariiska ee 55 DAE.
Jadwalka 1
Farqiga (P≤0.01) ee caleen photosynthetic midabka isdhexgalka, qaraabo chlorophyll content (Atleaf readings), iyo alfa-chlorophyll fluorescence xuduudaha u dhexeeya tijaabo iyo daawaynta ayaa lagu muujiyay shaxda 2. Maalinta sare heerkulka iyo habeenkii kordhay wadarta chlorophyll iyo carotenoid ka kooban yahay. Geedo bariiska oo aan lahayn buufin foliar ah ee phytohormones (2.36 mg g-1 "F67" iyo 2.56 mg g-1 ee "F2000") marka la barbardhigo dhirta lagu koray xaaladaha heerkulka ugu fiican (2.67 mg g -1)) waxay muujisay wadarta guud ee chlorophyll. Labada tijaabo, "F67" waxay ahayd 2.80 mg g-1 iyo "F2000" waxay ahayd 2.80 mg g-1. Intaa waxaa dheer, geedo bariiska oo lagu daaweeyay isku-darka AUX iyo GA ee buufinta kuleylka kuleylka ayaa sidoo kale muujiyay hoos u dhac ku yimaada maadada chlorophyll ee labadaba genotypes (AUX = 1.96 mg g-1 iyo GA = 1.45 mg g-1 for "F67"; AUX = 1.96 mg g-1 iyo GA = 1.45 mg g-AUX) g-1 iyo GA = 1.43 mg g-1 (loogu talagalay “F2000″) xaaladaha diiqada kulaylka. Xaaladaha kuleylka kuleylka, daaweynta foliar ee BR waxay keentay koror yar oo ku yimid doorsoomahan labada nooc ee genotypes. Ugu dambeyntii, buufinta CK foliar waxay muujisay qiimaha midabaynta sawirada ugu sarreeya ee dhammaan daawaynta (AUX, GA, BR, SC iyo AC) ee genotypes F67 (3.24 mg g-1) iyo F2000 (3.65 mg g-1). Nuxurka qaraabada ah ee chlorophyll (cutubka Atleaf) ayaa sidoo kale hoos loo dhigay cadaadiska kulaylka oo la isku daray. Qiimaha ugu sarreeya ayaa sidoo kale lagu duubay dhirta lagu buufiyo CC labadaba genotypes (41.66 ee "F67" iyo 49.30 ee "F2000"). Saamiyada Fv iyo Fv/Fm waxay muujiyeen farqi weyn oo u dhexeeya daawaynta iyo beeralayda (Shaxda 2). Guud ahaan, doorsoomayaashan ka mid ah, cultivar F67 aad bay ugu nuglayd diiqada kulaylka marka loo eego cultivar F2000. Saamiyada Fv iyo Fv/Fm ayaa aad u dhibtooday tijaabada labaad. Geedo cadaadis leh oo 'F2000' ah oo aan lagu buufin phytohormones ayaa lahaa qiimayaasha Fv ee ugu hooseeya (2120.15) iyo Fv/Fm saamiga (0.59), laakiin buufinta foliar ee CK waxay gacan ka gaysatay soo celinta qiimayaashan (Fv: 2591, 89, Fv/Fm ratio: 0.73). , helitaanka akhrin la mid ah kuwa lagu duubay dhirta "F2000" ee ku koray xaaladaha heerkulka ugu wanaagsan (Fv: 2955.35, Fv / Fm ratio: 0.73: 0.72). Ma jirin farqi weyn oo u dhexeeya fluorescence hore (F0), fluorescence ugu badnaan (Fm), dhalidda tirada ugu badan ee kiimikaad ee PSII (Fv/F0) iyo saamiga Fm/F0. Ugu dambeyntii, BR waxay muujisay isbeddel la mid ah sida lagu arkay CK (Fv 2545.06, Fv/Fm ratio 0.73).
Shaxda 2. Saamaynta cadaadiska kulaylka ee la isku daray (40°/30°C maalin/habeenkii) caleenta midabada sawir-qaadista [wadarta chlorophyll (Chl Total), chlorophyll a (Chl a), chlorophyll b (Chl b) iyo carotenoids Cx+c] saamaynta ], qaraabo chlorophyll content (Atliphyll fluremeters) (F0), fluorescence ugu badnaan (Fm), fluorescence variable (Fv), waxtarka ugu badnaan ee PSII (Fv/Fm), soosaarka ugu badnaan ee kiimikaad ee PSII (Fv/F0) iyo Fm/F0 ee dhirta labada genotypes bariiska [Federrose 67 (F67) iyo Federrose 2000 kadib) 5000DDA.
Mawduuca biyaha ee qaraabada ah (RWC) ee dhirta bariiska ee si kala duwan loo daweeyay ayaa muujiyay kala duwanaansho (P ≤ 0.05) ee isdhexgalka u dhexeeya daaweynta tijaabada iyo foliar (Jaantus. 1A). Marka lagu daweeyo SA, qiimayaasha ugu hooseeya ayaa loo diiwaangeliyay labadaba genotypes (74.01% F67 iyo 76.6% ee F2000). Xaaladaha kuleylka kuleylka, RWC ee dhirta bariiska ee labada genotypes ee lagu daweeyay phytohormones kala duwan ayaa si aad ah u kordhay. Guud ahaan, codsiyada foliar ee CK, GA, AUX, ama BR waxay kordhiyeen RWC ilaa qiyamka la mid ah kuwa dhirta oo lagu koray xaaladaha ugu wanaagsan inta lagu jiro tijaabada. Xakamaynta dhammaystiran iyo dhirta la buufiyay ee foliar waxay diiwaan geliyeen qiimayaal ku dhow 83% labada genotypes. Dhanka kale, gs ayaa sidoo kale muujisay kala duwanaansho weyn (P ≤ 0.01) ee isdhexgalka tijaabada-daawaynta (Jaantus 1B). Warshadda kontoroolka saxda ah (AC) waxay sidoo kale diiwaangelisay qiimaha ugu sarreeya ee genotype kasta (440.65 mmol m-2s-1 ee F67 iyo 511.02 mmol m-2s-1 ee F2000). Dhirta bariiska ee lagu soo daray isku-darka kulaylka oo kali ah ayaa muujisay qiimaha gs ugu hooseeya ee labada genotypes (150.60 mmol m-2s-1 ee F67 iyo 171.32 mmol m-2s-1 ee F2000). Daawaynta foliar ee dhammaan maamulayaasha koritaanka dhirta ayaa sidoo kale kordhay g. Dhirta bariiska ee F2000 ee lagu buufiyay CC, saamaynta foliar ku buufinta phytohormones ayaa ahayd mid aad u cad. Kooxdan dhirtu ma muujin wax farqi ah marka loo eego dhirta xakamaynta buuxda (AC 511.02 iyo CC 499.25 mmol m-2s-1).
Jaantus 1. Saamaynta isku-dhafka kulaylka (40 ° / 30 ° C maalintii / habeenkii) ee biyaha qaraabada ah (RWC) (A), stomatal conductance (gs) (B), malondialdehyde (MDA) wax soo saarka (C), iyo proline content. (D) ee dhirta laba genotypes bariiska (F67 iyo F2000) ee 55 maalmood ka dib soo bixitaanka (DAE). Daawaynta lagu qiimeeyay genotype kasta waxaa ka mid ahaa: xakamaynta buuxda (AC), xakamaynta kulaylka cadaadiska (SC), diiqada kulaylka + auxin (AUX), diiqada kulaylka + gibberellin (GA), diiqada kulaylka + mitogen cell (CK), iyo kulaylka + brassinosteroid. (BR). Tiir kastaa wuxuu u taagan yahay celceliska ± qaladka caadiga ah ee shan dhibcood (n = 5). Tiirarka ay raacaan xarfo kala duwan waxay muujinayaan kala duwanaansho weyn oo tirakoob ahaan marka loo eego imtixaanka Tukey (P ≤ 0.05). Waraaqaha leh calaamad siman ayaa tilmaamaya in celcelisku aanu ahayn mid xisaab ahaan muhiim ah (≤ 0.05).
MDA (P ≤ 0.01) iyo proline (P ≤ 0.01) waxa ku jira ayaa sidoo kale muujiyay farqi weyn oo u dhexeeya isdhexgalka u dhexeeya tijaabada iyo daaweynta phytohormone (Jaantus. 1C, D). Kordhinta peroxidation lipid ayaa lagu arkay daawaynta SC ee labada genotypes (Jaantuska 1C), si kastaba ha ahaatee dhirta lagu daweeyay buufinta korriinka caleenta waxay muujisay hoos u dhaca peroxidation lipid ee labada genotypes; Guud ahaan, isticmaalka phytohormones (CA, AUC, BR ama GA) waxay keenaysaa hoos u dhac ku yimaada peroxidation lipid (macluumaadka MDA). Wax farqi ah lagama helin dhirta AC ee labada genotypes iyo dhirta hoos yimaada cadaadiska kulaylka oo lagu buufiyo phytohormones (qiimaha FW ee dhirta "F67" waxay u dhexeeyaan 4.38-6.77 µmol g-1, iyo FW "F2000" dhirta "qiimaha la arkay waxay u dhexeeyaan 2.84 µ4-1 µmol . Isku-dhafka proline ee dhirta "F67" waxay ka hooseysay dhirta "F2000" ee hoos yimaada cadaadiska isku dhafan, taas oo keentay kororka wax soo saarka proline ee dhirta bariiska kuleylku, labadaba, waxaa la arkay in maamulka hormoonnadan ay si weyn u kordhiyeen maadada amino acid ee dhirta F2000 (AUX iyo BR waxay ahaayeen 30.44-18 g). 1G).
Saamaynta buufinta korriinka dhirta foliar iyo isku-darka kulaylka isku-dhafka ah ee heerkulka dabada dhirta iyo tusaha dulqaadka qaraabada (RTI) ayaa lagu muujiyey jaantusyada 2A iyo B. Labada genotypes, heerkulka daaqsinka ee dhirta AC wuxuu ku dhowaa 27 ° C, iyo tan dhirta SC waxay ku dhowdahay 28 ° C. leh Waxa kale oo la arkay in daawaynta foliar ee CK iyo BR ay keentay hoos u dhac 2-3°C ah ee heerkulka saqafka marka la barbar dhigo dhirta SC (Jaantuska 2A). RTI waxay soo bandhigtay dabeecad la mid ah doorsoomayaal kale oo jireed, oo muujinaya kala duwanaansho weyn (P ≤ 0.01) ee isdhexgalka u dhexeeya tijaabada iyo daaweynta (Jaantus 2B). Dhirta SC waxay muujisay dulqaadka dhirta hoose ee labada genotypes (34.18% iyo 33.52% ee "F67" iyo "F2000" dhirta bariiska, siday u kala horreeyaan). Quudinta foliar ee phytohormones waxay wanaajisaa RTI dhirta ay la kulmaan diiqad heerkul sare ah. Saamayntan ayaa si aad ah loogu caddeeyey dhirta "F2000" ee lagu buufiyay CC, taas oo RTI ay ahayd 97.69. Dhanka kale, kala duwanaansho la taaban karo ayaa lagu arkay kaliya tusaha cadaadiska wax-soo-saarka (CSI) ee dhirta bariiska ee hoos yimaada xaaladaha buufinta foliar factor (P ≤ 0.01) (Jaantus. 2B). Dhirta bariiska oo kaliya ee lagu soo dejiyay diiqada kulaylka adag ayaa muujisay qiimaha tusaha cadaadiska ugu sarreeya (0.816). Marka dhirta bariiska lagu buufiyo phytohormones kala duwan, tusaha walbahaarka ayaa ka hooseeya (qiyamka 0.6 ilaa 0.67). Ugu dambeyntii, geedka bariiska ee lagu beeray xaaladaha ugu wanaagsan wuxuu lahaa qiime dhan 0.138.
Jaantus 2. Saamaynta isku-darka kulaylka (40 ° / 30 ° C maalin / habeen) heerkulka saqafka (A), index dulqaad qaraabo (RTI) (B), iyo index stress dalagga (CSI) (C) ee labada nooc ee dhirta. Genotypes bariiska ganacsiga (F67 iyo F2000) ayaa lagu sameeyay daawayn kulayl oo kala duwan. Daawaynta lagu qiimeeyay genotype kasta waxaa ka mid ahaa: xakamaynta buuxda (AC), xakamaynta kulaylka cadaadiska (SC), diiqada kulaylka + auxin (AUX), diiqada kulaylka + gibberellin (GA), diiqada kulaylka + mitogen cell (CK), iyo kulaylka + brassinosteroid. (BR). Cadaadiska kulaylka ee isku dhafan waxay ku lug leedahay soo bandhigida dhirta bariiska heerkulka sare ee maalin/habeenkii (40°/30°C maalin/habeenkii). Tiir kastaa wuxuu u taagan yahay celceliska ± qaladka caadiga ah ee shan dhibcood (n = 5). Tiirarka ay raacaan xarfo kala duwan waxay muujinayaan kala duwanaansho weyn oo tirakoob ahaan marka loo eego imtixaanka Tukey (P ≤ 0.05). Waraaqaha leh calaamad siman ayaa tilmaamaya in celcelisku aanu ahayn mid xisaab ahaan muhiim ah (≤ 0.05).
Falanqaynta qaybaha ugu muhiimsan (PCA) ayaa shaaca ka qaaday in doorsoomayaasha lagu qiimeeyay 55 DAE ay sharaxeen 66.1% jawaabaha jireed iyo kiimikaad ee dhirta bariiska kuleylku cadaadisay oo lagu daweeyay buufinta nidaamiyaha koritaanka (Jaantus. 3). Vectors waxay u taagan yihiin doorsoomayaasha iyo dhibcuhu waxay matalaan nidaamiyayaasha koritaanka dhirta (GRs). Qaybaha gs, maadada chlorophyll, hufnaanta ugu badan ee tirada PSII (Fv/Fm) iyo cabbiraadaha biochemical (TChl, MDA iyo proline) waxay ku jiraan xaglo u dhow asalka, taasoo muujinaysa xidhiidh sare oo ka dhexeeya habdhaqanka jireed ee dhirta iyo iyaga. doorsoome. Hal koox (V) waxaa ku jiray geedo bariiska oo lagu beeray heerkulka ugu wanaagsan (AT) iyo dhirta F2000 ee lagu daweeyay CK iyo BA. Isla mar ahaantaana, inta badan dhirta lagu daweeyay GR waxay samaysteen koox gooni ah (IV), daawaynta GA ee F2000 waxay sameeyeen koox gaar ah (II). Taas bedelkeeda, geedo bariiska kuleylku cadaadisay (kooxaha I iyo III) iyada oo aan wax buufin foliar ah ee phytohormones (labada genotypes ahaayeen SC) waxay ku yaalliin aag ka soo horjeeda kooxda V, taas oo muujinaysa saameynta diiqada kulaylka ee physiology dhirta. .
Jaantuska 3. Falanqaynta taariikh nololeedka ee saamaynta isku-dhafka kulaylka (40°/30°C maalin/habeen) ee dhirta labada genotypes bariiska (F67 iyo F2000) ee 55 maalmood ka dib soo bixitaanka (DAE). Soo gaabinta: AC F67, xakamaynta buuxda F67; SC F67, xakamaynta cadaadiska kulaylka F67; AUX F67, diiqada kulaylka + auxin F67; GA F67, diiqada kulaylka + gibberellin F67; CK F67, diiqada kulaylka + qaybta unugyada BR F67, diiqada kulaylka + brassinosteroid. F67; AC F2000, xakamaynta buuxda F2000; SC F2000, Xakamaynta Cadaadiska Kulaylka F2000; AUX F2000, diiqada kulaylka + auxin F2000; GA F2000, diiqada kulaylka + gibberellin F2000; CK F2000, diiqada kulaylka + cytokinin, BR F2000, diiqada kulaylka + steroid naxaasta; F2000.
Kala duwanaanshiyaha sida maadada chlorophyll, habdhaqanka stomatal, Fv / FM ratio, CSI, MDA, RTI iyo proline content waxay kaa caawin kartaa fahamka la qabsiga genotypes bariiska waxayna qiimeeyaan saameynta xeeladaha agronomic ee hoos yimaada cadaadiska kulaylka (Sarsu et al., 2018; Quintero-Calderon et al., 2021). Ujeedada tijaabadani waxay ahayd in la qiimeeyo saamaynta codsiga afar nidaamiye oo korriin ah oo ku saabsan xuduudaha jireed iyo kiimikaad ee geedo bariiska oo hoos yimaada xaaladaha kuleylka adag. Tijaabada seedling waa hab fudud oo degdeg ah oo loogu talagalay qiimaynta isku mar ah ee dhirta bariiska iyada oo ku xidhan xajmiga ama xaaladda kaabayaasha la heli karo (Sarsu et al. 2018). Natiijooyinka daraasaddan ayaa muujisay in isku-darka kuleylka kulaylku uu keeno jawaabaha jireed iyo kiimiko ee kala duwan ee labada genotype ee bariiska, taas oo muujinaysa habka la qabsiga. Natiijooyinkani waxay sidoo kale muujinayaan in buufinta korriinka foliar-ka (badanaa cytokinins iyo brassinosteroids) ay ka caawiyaan bariiska inuu la qabsado diiqada kulaylka adag maadaama ay doorbidku inta badan saamayso gs, RWC, Fv/Fm ratio, midabada sawir-qaadista iyo waxyaabaha ka kooban proline.
Codsiga maamulayaasha korriinka waxay gacan ka geysataa hagaajinta xaaladda biyaha ee dhirta bariiska ee ku jira diiqada kulaylka, taas oo laga yaabo inay la xiriirto diiqad sare iyo heerkulka hoose ee geedka. Daraasadani waxay muujisay in ka mid ah dhirta "F2000" (genotype u nugul), dhirta bariiska ee lagu daweeyay ugu horrayn CK ama BR waxay lahaayeen qiyamka gs sare iyo qiimaha PCT hoose marka loo eego dhirta lagu daaweeyay SC. Daraasadihii hore ayaa sidoo kale muujiyay in gs iyo PCT ay yihiin tilmaamayaasha jireed ee saxda ah kuwaas oo go'aamin kara jawaabta la qabsiga ee dhirta bariiska iyo saameynta xeeladaha agronomic ee cadaadiska kulaylka (Restrepo-Diaz iyo Garces-Varon, 2013; Sarsu et al., 2018; Quintero). -Carr DeLong et al., 2021). Caleenta CK ama BR waxay kor u qaadaysaa g walaaca sababtoo ah hormoonnadan dhirta ayaa kor u qaadi kara furitaanka stomatal iyada oo loo marayo isdhexgalka synthetic ee unugyada kale ee calaamadaha sida ABA (kor u qaada xiritaanka stomatal ee cadaadiska abiotic) (Macková et al., 2013; Zhou et al., 2013). 2013). ). , 2014). Furitaanka caloosha waxay kor u qaadaa qaboojinta caleen waxayna caawisaa dhimista heerkulka saqafka (Sonjaroon et al., 2018; Quintero-Calderón et al., 2021). Sababahaan dartood, heerkulka saqafka dhirta bariiska ee lagu buufiyay CK ama BR ayaa laga yaabaa inuu hoos u dhaco marka la isku daro kulaylka.
Cadaadiska heerkulka sare wuxuu yarayn karaa nuxurka midabka sawir-qaadista ee caleemaha (Chen et al., 2017; Ahammed et al., 2018). Daraasaddan, markii dhirta bariiska ay ku jirtay diiqad kulayl oo aan lagu buufin wax nidaamiya koritaanka dhirta, midabada sawir-qaadista waxay u janjeersadeen inay hoos u dhigaan labada genotypes (Shaxda 2). Feng iyo al. (2013) ayaa sidoo kale sheegay hoos u dhac weyn oo ku yimid maadada chlorophyll ee caleemaha laba genotypes sarreen ah oo soo gaadhay diiqada kulaylka. Soo-gaadhista heerkulka sare waxay badanaa keentaa hoos u dhaca maadada chlorophyll, taas oo laga yaabo inay sabab u tahay hoos u dhaca chlorophyll biosynthesis, hoos u dhaca midabka, ama saameyntooda isku dhafan ee cadaadiska kulaylka (Fahad et al., 2017). Si kastaba ha ahaatee, dhirta bariiska ee lagu daweeyay inta badan CK iyo BA waxay kordhisay uruurinta midabada sawir-qaadista ee caleenta ee cadaadiska kulaylka. Natiijooyin la mid ah ayaa sidoo kale laga soo sheegay Jespersen and Huang (2015) iyo Suchsagunpanit et al. (2015), kuwaas oo arkay korodhka chlorophyll caleen ka dib codsiga zeatin iyo hormoonnada epibrassinosteroid ee bentgrass iyo bariiska, siday u kala horreeyaan. Sharaxaad macquul ah oo ku saabsan sababta CK iyo BR ay kor ugu qaadaan korodhka chlorophyll caleen ee hoos yimaada cadaadiska kulaylka ee isku dhafan ayaa ah in CK laga yaabo inay wanaajiso bilawga soo-gelinta joogtada ah ee dhiirrigeliyeyaasha (sida dhiirrigeliyaha firfircoonida firfircoonida (SAG12) ama dhiirrigeliyaha HSP18) oo ay hoos u dhigto lumitaanka chlorophyll ee caleemaha. , dib u dhigista sniinta caleen waxayna kordhisaa caabbinta dhirta kulaylka (Liu et al., 2020). BR waxay ilaalin kartaa chlorophyll caleen waxayna kordhin kartaa chlorophyll caleen iyada oo dhaqaajisa ama kicinaysa isku dhafka enzymes ee ku lug leh chlorophyll biosynthesis ee xaaladaha walaaca (Sharma et al., 2017; Siddiqui et al., 2018). Ugu dambeyntii, laba phytohormones (CK iyo BR) waxay sidoo kale kor u qaadaan muujinta borotiinnada shoogga kulaylka waxayna hagaajiyaan hababka kala duwan ee la qabsiga dheef-shiid kiimikaadka, sida kororka chlorophyll biosynthesis (Sharma et al., 2017; Liu et al., 2020).
Chlorophyll a xuduudaha fluorescence waxay bixiyaan hab degdeg ah oo aan burburin taasoo qiimeyn karta dulqaadka dhirta ama la qabsiga xaaladaha walaaca abiotic (Chaerle et al. 2007; Kalaji et al. 2017). Halbeegyada sida saamiga Fv/Fm ayaa loo adeegsaday tilmaamayaasha la qabsiga dhirta xaaladaha walaaca (Alvarado-Sanabria et al. 2017; Chavez-Arias et al. 2020). Daraasaddan, dhirta SC waxay muujisay qiimaha ugu hooseeya ee doorsoomiyahan, oo u badan dhirta bariiska "F2000". Yin iyo al. (2010) waxa kale oo ay ogaatay in saamiga Fv/Fm ee caleemaha bariiska beerashada ugu sarreeya ay si weyn hoos ugu dhacday heerkul ka sarreeya 35°C. Sida laga soo xigtay Feng et al. (2013), saamiga hoose ee Fv / Fm ee hoos yimaada diiqada kulaylku waxay muujinaysaa in heerka tamarta kicinta ee qabashada iyo beddelka xarunta falcelinta PSII ay hoos u dhacday, taas oo muujinaysa in xarunta falcelinta PSII ay ku kala jajabto cadaadiska kulaylka. U fiirsashadani waxay noo ogolaanaysaa in aan ku soo gabagabeyno in khalkhalka ku jira qalabka sawir-qaadista ay aad ugu dhawaaqaan noocyo xasaasi ah (Fedearroz 2000) marka loo eego noocyada u adkaysta (Fedearroz 67).
Isticmaalka CK ama BR guud ahaan waxay xoojisay waxqabadka PSII ee xaaladaha adag ee kulaylka. Natiijooyin la mid ah ayaa waxaa helay Suchsagunpanit et al. (2015), kuwaas oo arkay in codsiga BR uu kordhiyay waxtarka PSII ee cadaadiska kulaylka ee bariiska. Kumar iyo al. (2020) waxa kale oo ay ogaatay in dhirta chickpea ee lagu daweeyay CK (6-benzyladenine) oo ay la kulantay diiqada kulaylku ay kordhisay saamiga Fv/Fm, iyada oo la soo gabagabeeyay codsiga foliar ee CK iyada oo la kicinayo wareegga midabka zeaxanthin ee kor u qaaday waxqabadka PSII. Intaa waxaa dheer, buufin caleen BR jecel yahay PSII photosynthesis hoos xaaladaha stress isku dhafan, taas oo muujinaysa in codsiga phytohormone this keentay in la dhimay ee tamarta excitation ee anteenooyinka PSII iyo kor u raasamaal ee borotiinada shoogga kulaylka yar ee chloroplasts (Ogweno et al. 2008; Kothari iyo Lachowitz). , 2021).
MDA iyo waxa ku jira proline inta badan way kordhaan marka dhirta ay ku hoos jiraan walaaca abiotic marka la barbar dhigo dhirta ku koray xaaladaha ugu wanaagsan (Alvarado-Sanabria et al. 2017). Daraasadihii hore ayaa sidoo kale muujiyay in MDA iyo heerarka proline ay yihiin tilmaamayaasha biochemical kuwaas oo loo isticmaali karo in lagu fahmo habka la qabsiga ama saameynta habdhaqanka beeraha ee bariiska maalintii ama habeenkii heerkulka sare (Alvarado-Sanabria et al., 2017; Quintero-Calderón et al. ., 2021). Daraasadahani waxay sidoo kale muujiyeen in MDA iyo waxyaabaha ku jira proline ay u janjeeraan inay ka sarreeyaan dhirta bariiska ee soo gaadha heerkul sarreeya habeenkii ama maalinta, siday u kala horreeyaan. Si kastaba ha ahaatee, buufinta foliar ee CK iyo BR waxay gacan ka geysatay hoos u dhaca MDA iyo korodhka heerarka proline, inta badan genotype dulqaadka (Federroz 67). Buufinta CK waxay kor u qaadi kartaa xad-dhaafka ah ee cytokinin oxidase / dehydrogenase, taas oo kordhinaysa waxa ku jira xeryahooda ilaalinta sida betaine iyo proline (Liu et al., 2020). BR waxay kor u qaadaa soo saarista osmoprotectants sida betaine, sonkorta, iyo asiidhyada amino (oo ay ku jiraan proline bilaashka ah), ilaalinta dheelitirka osmotic-ka gacanta ee hoos yimaada xaalado badan oo deegaan oo xun (Kothari iyo Lachowiec, 2021).
Tusaha walbahaarka dalagga (CSI) iyo tusaha dulqaadka qaraabada (RTI) ayaa loo isticmaalaa si loo go'aamiyo in daawaynta la qiimaynayo ay ka caawinayso yaraynta culaysyada kala duwan (abiotic iyo biotic) oo ay saamayn togan ku leeyihiin physiology dhirta (Castro-Duque et al., 2020; Chavez-Arias et al., 2020). Qiimayaasha CSI waxay u dhaxayn karaan 0 ilaa 1, oo ka dhigan xaaladaha aan walbahaarka iyo walbahaarka, siday u kala horreeyaan (Lee et al., 2010). Qiimaha CSI ee dhirta kuleylka-cadaadiyey (SC) waxay u dhaxaysaa 0.8 ilaa 0.9 (Jaantuska 2B), taas oo muujinaysa in dhirta bariiska ay si xun u saamaysay walaaca isku dhafan. Si kastaba ha ahaatee, buufinta foliar ee BC (0.6) ama CK (0.6) waxay inta badan keentay hoos u dhac ku yimaadda tilmaameha xaaladaha walaaca abiotic marka la barbar dhigo dhirta bariiska SC. Dhirta F2000, RTI waxay muujisay koror sare marka la isticmaalayo CA (97.69%) iyo BC (60.73%) marka la barbar dhigo SA (33.52%), taas oo muujinaysa in maamulayaasha koritaanka dhirta ay sidoo kale gacan ka geystaan hagaajinta jawaabta bariiska ee dulqaadka halabuurka. Kulayl xad dhaaf ah Tilmaan-bixiyahan ayaa la soo jeediyay si loo maareeyo xaaladaha walaaca ee noocyada kala duwan. Daraasad uu sameeyay Lee et al. (2010) waxay muujisay in CSI ee laba nooc oo cudbi ah oo hoos yimaada cadaadiska biyaha dhexdhexaadka ah waxay ahaayeen qiyaastii 0.85, halka qiimaha CSI ee noocyada si fiican loo waraabiyay ay u dhexeeyaan 0.4 ilaa 0.6, iyada oo lagu soo gabagabeynayo in tusmadani ay tahay calaamad muujinaysa la qabsiga biyaha ee noocyada. xaalado walaac leh. Intaa waxaa dheer, Chavez-Arias et al. (2020) waxay qiimeeyeen waxtarka elicitors synthetic sida istaraatijiyad maaraynta walbahaarka oo dhamaystiran ee dhirta C. elegans waxayna ogaadeen in dhirta lagu buufiyo xeryahooda ay muujiyeen RTI sare (65%). Iyadoo lagu saleynayo kuwa kore, CK iyo BR waxaa loo tixgalin karaa inay yihiin xeelado beeraley ah oo looga golleeyahay in lagu kordhiyo dulqaadka bariiska ee culeyska kulaylka adag, maadaama nidaamyadan korriinka dhirta ay keenaan jawaabaha biochemical iyo physiological togan.
Dhowrkii sano ee la soo dhaafay, cilmi baarista bariiska ee Kolombiya waxay diiradda saartay qiimeynta genotypes u dulqaadan kara heerkulka maalinta ama habeenkii iyadoo la adeegsanayo sifooyinka jireed ama kiimikaad (Sánchez-Reinoso et al., 2014; Alvarado-Sanabria et al., 2021). Si kastaba ha noqotee, dhowrkii sano ee la soo dhaafay, falanqaynta tignoolajiyada wax ku oolka ah, dhaqaalaha iyo faa'iidada leh ayaa noqotay mid sii kordheysa oo muhiim ah in la soo jeediyo maaraynta dalagyada isku dhafan si loo wanaajiyo saameynta xilliyada adag ee diiqada kulaylka ee dalka (Calderón-Páez et al., 2021; Quintero-Clderon et al., 2021). Sidaa darteed, jawaabaha jireed iyo kiimikaad ee dhirta bariiska ee cadaadiska kuleylka adag (40 ° C maalintii / 30 ° C habeenkii) lagu arkay daraasaddan waxay soo jeedinaysaa in buufinta foliar ee CK ama BR ay noqon karto hab ku habboon maareynta dalagga si loo yareeyo saameynta xun. Saamaynta xilliyada cadaadiska kulaylka dhexdhexaadka ah. Daawayntani waxay wanaajisay dulqaadka labada genotypes bariiska (CSI hoose iyo RTI sare), taasoo muujinaysa isbeddelka guud ee jawaabaha jireed ee dhirta iyo bayooloji ee hoos yimaada cadaadiska kulaylka ee isku dhafan. Jawaabta ugu weyn ee dhirta bariiska waxay ahayd hoos u dhac ku yimid nuxurka GC, wadarta chlorophyll, chlorophylls α iyo β iyo carotenoids. Intaa waxaa dheer, dhirtu waxay la ildaran tahay burburka PSII (hoos u dhaca cabbirrada fluorescence chlorophyll sida saamiga Fv/Fm) iyo kororka peroxidation lipid. Dhanka kale, markii bariiska lagu daweeyay CK iyo BR, saamaynta xun ayaa la yareeyay waxaana kordhay maadada proline-ka (Jaantus. 4).
Jaantuska 4. Qaabka fikradda ah ee saamaynta kulaylka isku dhafan iyo xakamaynta koritaanka dhirta foliar ku buufin dhirta bariiska. Fallaadhaha casaanka iyo buluuga ah waxay muujinayaan saamaynta taban ama togan ee isdhexgalka ka dhexeeya cadaadiska kulaylka iyo codsiga foliar ee BR (brassinosteroid) iyo CK (cytokinin) ee jawaabaha jireed iyo kiimikaad, siday u kala horreeyaan. gs: habdhaqanka caloosha; Wadarta Chl: wadarta guud ee kolorofilka; Chl α: chlorophyll β ka kooban yahay; Cx+c: maadada carotenoid;
Marka la soo koobo, jawaabaha nafsiyeed iyo kiimikaad ee daraasaddan ayaa tilmaamaya in Fedearroz 2000 dhirta bariiska ay aad ugu nugul yihiin muddada kuleylka adag ee Fedearroz 67 dhirta bariiska. Dhammaan xeeriyeyaasha korriinka ee lagu qiimeeyay daraasaddan (auxins, gibberellins, cytokinins, ama brassinosteroids) waxay muujiyeen xoogaa hoos u dhigista cadaadiska kulaylka. Si kastaba ha ahaatee, cytokinin iyo brassinosteroids waxay keeneen la qabsiga dhirta ka wanaagsan maadaama labada maamulayaasha koritaanka dhirta ay kordhiyeen maadada chlorophyll, cabbirada fluorescence alfa-chlorophyll, gs iyo RWC marka la barbar dhigo dhirta bariiska iyada oo aan wax codsi ah la helin, iyo sidoo kale hoos u dhac ku yimid nuxurka MDA iyo heerkulka korka. Marka la soo koobo, waxaan ku soo gabagabeyneynaa in isticmaalka nidaamiyeyaasha koritaanka dhirta (cytokinins iyo brassinosteroids) ay tahay qalab waxtar leh oo lagu maareeyo xaaladaha walaaca ee dalagyada bariiska oo ay keento diiqada kulul ee daran xilliyada heerkulka sare.
Qalabka asalka ah ee lagu soo bandhigay daraasadda ayaa lagu soo daray maqaalka, su'aalo dheeraad ah ayaa lagu hagaajin karaa qoraaga u dhigma.
Waqtiga boostada: Agoosto-08-2024