DANH GIA KHA NANG CHIU HAN VA MOT SO CHI TIEU H6A SINH CUA CAC DONG NGO CHUYEN GEN ZMDREB2A GIAI DOAN TRUdC TRO
Nguyen Dde Trpng', Doan Thi Bich Thao', Nguyen Xuan Thdng' TOM T A T
Danh gia kha nang chiu ban vd mgt so chi lieu hda sinh eua 3 dong ngd D3, D14 vd D2I chuyen gen chju han ZmDREB2A the he T6 vd 3 dong ngd nen tUdng dng C436, C7N va \T52 thdng qua thi nghiem gay ban nhan t^o giai doan trUde trd trong dieu kien nha ludi. Ket qua eho thay trong dieu kien 14 ngay gay ban, cac ehi tieu ve hinh thai Id, thdi gian sinh trUdng (TGST), khoang each tung phan phun rau (ASI), cae dac diem ndng hoc va hdm lugng mgt sd chi tieu hda sinh nhu dam tdng so, proline, hydrateacbon, earotenoid, chlorophyll cua dong chuyen gen cao hon CO y nghia so vdi ddng nen tuong flng. Trong dd nang suat ca the cua cae dong chuyen gen vUgt so vdi ddng nen tUOng flng tfl 25,7 - 33,8%. Kel qua da ehgn Igc dUOc 2 ddng chuyen gen ZmDREB2A la D3 va D21 the hi^n khd nang chiu han hieu qua cao sfl dung lam ngudn vat lieu phuc vu cdng tac ebon tao giong ngo bien ddi gen chiu han.
Tfl khda: Ngd {Zea mays L.), chuyen gen, ZmDREB2A, ehiu ban I. DAT VAN DE
Han hdn dang gay thiet hai nhieu nhat trong sd mot loat mdi de dga cua nganh ndng nghiep cua cdc niifn kinh te dang phdt trien. Chi tinh trong 10 nam gay day, han hdn da lam ton that hang chuc ty' dd la chu yeu khu vUc Chau A (FAO, 2018). Tlieo bao cdo eua Tdng cue Phdng ehdng thien tai, tinh den 3/2020 do dnh hudng eua bien ddi khi hau khien tinh trang ban han, xam nhap man d Ddng bdng sdng Cdu Long (DBSCL) cung nhu tinh hinh thieu nUde, khd h^in d Ddng bdng Bdc Bg, Trung Bg vd Tay Nguyen xuat hi?n sdm tren dien rdng. Dac biet cdc tinh DBSCL nhU Ca Mau, Ben Tre, Kien Giang, Sdc Trang do thdi gian thieu mda keo dai ket hgp vdi vi^c sfl dung vd khai thdc lai nguyen nUdc trong lUu vdc da Idm eho han han va thieu nUdc trd nen tram trgng hdn va ddgc cho la khde liet ehUa tflng cd trong lich sfl Idm thiet hai hdng chuc nghin ha cay mau (Tdng egc Phdng chdng thien tai, 2020).
Cdng tdc ehpn tao gidng edy trdng ehiu ban ndi chung va eay ngd ndi rieng la thach thflc Idn cho cac nhd chgn gidng do bdi chiu han ddgc quy dinh bdi h? thdng da gene. Nghien cUu di truyen phan tfl tinh ehiu han da chi ra rdng, phan flng chdng chiu dieu ki?n han dUge dieu khien bdi mdt vdi gene trong h? thdng cac gene quy dinh tinh chju han. Viec dng dung cdng nghe chuyen gene nham chuyen ede gene dong vai trd dieu khien vdo cay trong dang ddgc the gidi lien hanh nhieu nam qua nhdm tao ra gidng ngd chdng chiu dieu kien han. He thdng gen dieu khien bao gdm cae gen chfla ma di truyen tong hdp nen mgt loai protein ndo dd, ldm nhiem vu boat hda tdng hgp mdt loat protein thfl cap giup tang cfldng tinh chdng chiu cua cay ddi vdi dieu kien ngoai canh nhu
ma hda Protein LE.A (late embryogenesis abundant);
Protein sde nhiet (HSP) hay tdng sU tich luy chat khd dde td trong te ba trong edy Arabidopsis.
Hien nay, 4 hg gen dieu khien dUdc cdc nha nghien edu quan tdm Id: Dreb hoac CBF, MYB, bZIP va Zinc-finger (Umezawa el al, 2006), trong do yeu td phien ma DREB ddng vai trd kich boat bieu hien hdng loat cae gen dap flng vdi dieu kien ban cung nhfl cac yeu to stress khac nhd tang cfldng tinh chju ban va man d Ifla (Oh. Song et al, 2005); ndng va lanh d Arabidopsis (Dubouzet, Sakuma el al, 2003, Mizoi et ai, 2010). d ngd nam 2007, nhdm nghien cflu eua Qiu da phdn lap duge cDNA cua gen chiu ban ZmDREB2A tfl ngd. ZmDREB2A ma hod phdn td protein gdm 318 amino acid chfla vung gan vdi DNA ERP/AP2 dae trUng eho DREB. Thdng qua thi nghiem bieu hien trong Arabidopsis, protein ZmDREB2A da dUOc chflng mmh ed kha nang tang eUdng sflc chdng chiu vdi dieu kien ban cho edy (Qin et al., 2007).
II. VAT LIEU VA PHUONG PHAP NGHIEN CtfU 2.1. Vat lieu nghien cdu
Vat lieu nghien cdu bao gdm 3 ddng ngd chuyen gen chiu han ZmDREB2A (d the he T6) ky hieu D3, D14vdD21. Cae ddng chuyen gen dfldc tao ra thdng qua phuong phap chuyen vector pCAMBIA1300 mang gen chiu han ZmDREB2A thdng qua vi khuan Agrobacterium vdo phdi non 12 ngay tudi cua 3 ddng ngd nen tUdng dng ed kha nang tai sinh cad la C436, C7N vd V152. Cac ddng chuyen gen da dUdc ddnh gid sU cd mat vd bieu hien on dinh cua gen chuyen trong eac ddng ngd bdng cac ky thuat phan tfl.
' Vi^n Nghien cuu Ngd
2.2. PhUdng phap nghien cdu
2.2.1. Ph Udng phdp ddnh gid khd ndng chiu han Phfldng phdp gav han giai doan trUde trd theo CIMMVT (2004) va Cains vd cdng tdc vien (2013) Cae ddng ngd dugc gieo trong xd chfla gia the trong dieu kien nha lUdi. Sau khi edy ngd dugc 3 Id tien hdnh tia bd cdc cay khdng ddng deu chi gifl lai 1 cay/
xd/ddng, mdi ddng 5 xd vdi thi nghiem xd ly ban vd sd dung ddi chflng tddi nUdc day dd. Cdc xd thi nghiem dUde bdn phdn va tudi nUde ddng deu d mdi xd de edy smh trUdng phat trien binh thUdng. Tien hanh ngdng tddi nUde nham gay ban nhan tao ddi vdi cdc ddng ngd dam bao thi nghiem han duge tinh d thdi diem trUdc trd khoang 1 tuan. Cae ddng ngd Idm ddi chdng (khdng xfl ly ban) van tddi nfldc day du. Sau 14 ngdy gay ban ede xd thi nghiem tfldi nflde trd lai de edy phdt trien binh thudng cho den khi thu hoach. Cdc chi tieu theo ddi vd danh gid theo phuong phdp cua CIMMYT (2004).
2.2.2. PhUdngphdp ddnh gid chi tieu hoa sinh Theo phUdng phap chuan ve phdn tich ham lugng dam tdng sd. ham lugng ehldrophyll, phan tich NSC cua Kjeldahn (1883) cd cai tien; Ohniehi and Horie (1999).
2.2.3. PhUdng phdp xii ly so lieu
So lieu ve ede ehi tieu theo ddi dugc ddnh gid.
thu thap, xd ly thdng ke bdng ehdcrng trinh Excel va phan mem IRRISTAT 5.0.
2.3. Thdi gian va dia diem nghien cdu Cdc thi nghiem ddgc thdc hi?n trong \-y He Thu 2019 trong dieu kien nhd lUdi tai \'ien Nghien cdu Ngd - Dan Phdgng. Hd Ngi.
in. KET QUA VA THAO LUAN
3.1. Kha nang chiu ban cua cac dong chuyen gen vd ddng nen trong dieu hien han giai doan trddc tr6 3.1.1. Miic do cugn Id
Ket qua ddnh gia mflc dd eugn Id trong thdi gian xd ly han eho thay: Sau 7 ngay gay ban, cdc ddng ngd sinh trddng cham lai, Id bat dau chuyen td mau xanh dam sang xanh nhat. Cae ddng nen cd hien tugng cugn d mep la (diem 2). Sau 10 ngdy gay ban. d cdc ddng chuyen gen bdt dau la ed bieu hien cugn lai (diem 2) nhUng d mdc do nh? hdn ede ddng nen (diem 3). Sau 14 ngay, tat ca ede ddng tham gia thi nghiem deu heo vd cd mdc do eugn Id diem 5. Tuy nhien trong thi nghiem chung tdi ghi nhdn ede ddng nen deu cd bieu hien chay la d lang la dUdi trong khi cdc ddng chuyen gen chi bi ehdy phan dinh, mep Id.
Ddng/ Ddng D3 C436 nen
D14 C7N nen
D21
\T52nen CV(%) LSDoo.
Bdng 1. Muc dg eugn la, thdi gian sinh trUdng va mgt sd dac diem hinh thai eua eac dong ngd tham gia thi nghiem trong dieu ki?n han Dp cudn Id sau (diem)
•gdy 1 2 1 2 1
10 ngdy 2 3 2
,
14 ngdy 5 5 5 5 5
AS!
5 6 4 6 3
T G S T (ngay)
115 113 125 124 123 120
Cao cay (cm) 99,2 91,4 108,2 94,2 104,0
91,4 4,9 7,6
C a o bap (cm) 69,4 64,0 70,3 65.9 72,8 63,9 5,0 4,8 3.1.2. Khodng cdch tung phdn phun rdu vd TGST
Ket qua bdng 1 cho thay ban giai doan trUde trd dnh hudng rd ret den sinh trUdng phdt trien cua cay ngd. 6 cdc ddng chuyen gen va ddng nen khoang each thdi gian tung phan va phun rau deu bi keo ddi hdn so vdi d dieu kien thudng. Tuy nhien cac ddng chuyen gen it bi anh hddng the hien khodng cdch tung phan phun rau ngan hdn cac ddng nen td 1 - 2 ngdy. Ddng chuyen gen D21 ed khoang each ngdn nhat (3 ngdy) so vdi ddng nen tddng flng V152
(5 ngay) tiep den la 2 ddng chuyen gen D14 va D3.
Theo eac nha khoa hgc mdt ngudn vat lieu chiu han tdt thi khoang each tung phap-phun rau cang nhd vd bieu hien ngay khi cay bi ban (Bolafios and Edmeadres, 1996).
Thdi gian sinh trUdng cua cdc ddng vd ddng nen trong dieu kien gay ban the hien khde nhau va ngan hon so vdi dieu kien khdng gay han. Cac ddng nen ed thdi gian tfl gieo den chin sinh ly ngdn hdn 1 - 3 ngay so vdi cac ddng chuyen gen. Ddng nen C436 vd
TGD chi Khoa hex: Cong none ;\ _ - ddng chuyen gen D3 ed thdi gian sinh irudng ngdn
nhat lan lugt 113 ngay va 115 ngdy Ddng nen C7X va ddng chuyen gen D14 cd thdi gian sinh trUdng dai nhat tUdng flng 124 ngdy vd 125 ngay trong dieu ki?n han.
3.1.3. Chieu cao cdy, chieu cao dong bdp Trong dieu kien xfl ly han, khd nang sinh trUdng phdt tnen cua cae ddng chuyen gen D3, D14 va D21 vd cac ddng nen tUdng flng C436, C7N vd ddng VI52 deu bj dnh hUdng. Tuy nhien cac ddng nen chiu anh hudng manh hdn so vdi cae ddng chuyen gen trong ciing dieu kien han the hien chieu eao eay va chieu cao ddng bap eua eac ddng nen thap hon eac ddng chuyen gen. Ddng nen C7N cd chieu cao edy, chieu cao ddng bap Id 94,2 cm vd 65,9 cm tUdng dng d ddng chuyen gen D14 la 108 em va 70,3 em. TUdng tfl dddng nen C436 la 91,4 em vd 64 em so vdi ddng chuyen gen D3 Id 99,2 em va 69,4 cm. Trong qua trinh sinh trddng, phdt trien cua cay trdng, nUde can eho mgi boat dgng sdng cua te bao. Khi klidng dugc cung cap ndde day du ho^t ddng trao ddi chat eua te bao b\ anh hfldng, qud trinh phdn chia te bdo dien ra cham lai. Do dd sinh trfldng phat trien cua cay
trong dieu kien han se cham lai so vdi khi khdng xay rahan.
3.2. Cac yeu to eau thanh nang suat va nang suat ca the cua ddng
Cdc yeu td cau thanh ndng suat gdm chieu ddi bap, dudng kinh bdp. sd hdng hat, so hat/hang, ty le khdi lUdng hat/bdp, khdi lUgng 1000 hat vd ndng suat ca the eua eac ddng ddgc trinh bay d bang 2.
Ket qua thi nghiem cho thay. sau thdi gian 14 ngay xfl ly ban giai doan trfldc trd, ede ddng chuyen gen ZmDREB2A deu cd chi sd cao hdn so vdi ddng nen tddng flng.
3.2.1. Chieu ddi bdp vd dUdngktnh bdp Trong dieu kien ban, eac ddng chuyen gen D3, D14 vaD21 deu ed chieu ddi bdp vd dfldng kinh bap cao hdn eac ddng nen tfldng flng. Ddng chuyen gen D21 cd chieu dai bdp va dudng kinh bdp lan lUgt dat 13,6 cm vd 3,6 cm cao hdn cdc ddng cdn lai vd eao hdn so vdi ddng nen V152 (11,0 em va 2,9 em).
Tuong tu ddng chuyen gen D3 vdi ddng nen C436 la 9,9 em; 2,7 cm vd 8,8 cm; 2,4 cm vd ddng chuyen gen D14 vdi ddng nen C7N la 11,2 em; 2,4 em vd 9,4 em; 2,0 cm.
Bang 2, Mgt sd yeu to cau thanh nang suat va nang suat ea the cua cac ddng ngd tham gia thi nghiem Irong dieu kien ban TT D 6 n g
1 D3 2 1 C436 nen 3 D14 4 C7N nen 5 D21 6 V152 nen
CV(%) LSD,,,
Dai bap 9,9 8,8 11,2 9,4 13,6 11,0 7,2 1,12
D K b a p 2,7 2 4 2,4 2,0 3.6 2.9 5,0 0,19
So hang hat 10,2 9,2 8,8 7,4 8,8 7,6 9,9 1,27
So hat/
hang 18,6 16,5 19,9 16,8 24,9 20,0 7,4 2,1
Tyle hat/bap (%)
52,5 46,5 52,1 44,0 59,6 48,0
PIOOO hat (gr) 219,3 201,8 200,1 191,5 219,1 209,3 4,5 14,08
Nang suat (g/CT)
41,2 30,6 35,0 23,8 48,0 31,8 4,0 4,36
3.2.2. So hdng hat vd so hqt/hdng
Ngoai chi tieu chieu dai bap va dfldng kinh bap thi ehi tieu ve sd hdng hat vd sd hat/hdng cung ddng gdp dang ke vdo ndng suat cua ddng. Sd hang hat la tinh trang phu thude ehfl yeu vdo kieu gen va kha on dinh khi ddng cd do thuan cao. Trong khi sd hat/hdng phu thugc vao dieu kien mdi trfldng va dinh dddng. Qua ket qud danh gia dflgc trinh bay d bang 2 cho thay sd hang hat vd sd hat/hdng cua ede ddng chuyen gen deu cao hdn so vdi ddng nen tfldng dng trong dieu kien xfl ly han. Sd hang hat va sd hat/hang cua ddng chuyen gen D3 so vdi ddng
nen C436 tfldng dng dat 10,2; 18,6 va 9,2; 16,5. Ddng chuyen gen D14 va ddng nen C7N dat tfldng dng la 8,8; 19,9 vd 7,4; 16,8. Dieu nay cd the giai thich khi stress ban xay ra giai doan trfldc trd lam anh hfldng Idn den qud trinh thu phan, thu tinh va kha nang ket hat d tat ea cac ddng ehuven gen va ddng nen. Stress han Idm giam dang ke sd Ifldng hat/hang dae biet a nhflng ddng nen. Giai doan trd cd ede ddng ngd ehiu han ed khoang cdch tung phan, phun rau (ASI) ngan hdn ddng nen, qua trinh thu phan thu tinh thuan Igi hdn lam tang sd hat/hang.
3.2.3. Ty le hat/bdp vd khoi lUdng 1000 hat Han ldm tj- le hat/bdp giam mdt each ddng ke.
ty' le hat/bap cua cae ddng chuyen gen dao ddng tfl 52,1 - 59,6%, trong dd cdc dong nen dat 44.0 - 48,0%, ty le thap nhat la ddng nen C7X. Khdi Ifldng 1000 hat cua cae ddng chuyen gen vd ddng nen trong dieu kien han dao dgng td 191,5 - 219,3 g. Trong dd ddng chuyen gen D3 vd D21 cd khdi lUdng 1000 hat eao nhat, thap nhat la ddng D14.
3.2.4. Ndngsudt cd the cua cdc dong ngo Trong dieu kien han, nang suat eua ddng chuyen gen va ddng nen khde biet ed y nghia vdi do tin cay 95%. Ddng chuyen gen D21 dat ndng suat ca the 48,0 g cao hdn 16,2 g (tddng dng 33,8%) so vdi ddng nen V152 (31,8 g). TUdng tU ddng chuyen gen D14 dat ndng suat ca the 35,0 g cao hon 32% so vdi ddng nen C7N (23,8 g); va ddng D3 cho nang suat cao hdn 25,7% so vdi ddng nen C436.
3.3. Ket qua danh gia mgt so chi tieu hda sinh cac ddng chuyen gen vd ddng nen trong dieu hien han giai doan trddc trd.
Td ket qua bang 3 cho thay sau 14 ngdy gay han nhdn tao trong dieu kien nha ludi, ham lUdng eac chi sd hda sinh trong ede ddng chuyen gen tang eao
hon d ede ddng nen. Ddng D3 eho ham lUdng d?m tdng sd eao nhat dat 6,12 g/100 g mau tdng 1,49 g so vdi ddng nen C436 (4,36 g) va the hien sU chenh lenh eao hdn so vdi 2 ddng cdn lai. Tiep den la ddng D14 vdi ddng nen C7N vd ddng D21 vdi ddng \ ' 152.
TUdng tu ham lUdng lipid trong cae ddng chuyen gen deu eao hdn ddng nen tUdng dng trong cung dieu kien han. Ham lUdng lipil eua cdc ddng dao ddng tfl 1,62 -1,78 g/100 g chat mau trong khi ham lugng lipid d cdc ddng nen deu thap hdn dao dgng tdO,Sl - 1,15 g.
Ket qud bang 3 cung cho thay ham Idgng proline va hdm Idgng cabonhydrat trong cdc ddng nen khdng chuyen gen deu thap hdn d ede ddng ngd chuyen gen tUdng flng. Hdm lugng proline va hdm Idgng cabonhydrat d ddng C436 cho sfl chenh leeh gifla cay chuyen gen vd edy ddng nen cao hdn so vdi 2 ddng cdn lai. Trong nhieu nghien edu da ehi ra cd sU tich luy dang ke amino acid nhU proline khi eay ngd gap phdi dieu kien ban va ham lugng proline gidm khi dfldc tUdi nUdc trd lai (Mansour, 2000).
TUdng tu, cac dang carbonhydrate khdng cau true hay dudng tan ddgc tang cUdng trong cac te bdo khi gap dieu kien ban va ddng vai trd quan trgng trong nee dieu hda tham thau va bdo ve eau true te bao.
Bang 3. Ham lugng mgt so ehi lieu hda sinh cua eac ddng chuyen gen va dong nen sau thi nghiem xfl ly han nhan tao
Dam tong , Carotenoid Proline CabonHydrat Chlorophyll Ham lUgng 1
^
3 4 5 6
D3 C436 nen
D14 C7N nen
D21
\ 152 nen s o ( g )
6.12 4.63 4.50 4,09 4,05 3,86
1,64 0,86 1,62 1,15 1.78 0,81
(mg) 7,0 6,55 6,56 6,10 6 4 9 6,28
jmg) 4,3 2,9 4,1 2,8 3,6 2,3
Jmg)
7,38 5,78 7,56 6,30 7,63 6,21
(mg) 11,9 6,6 9,8 6 , 4 12,7 7,4
niidc (g) 62,55 51,12 64,14 49,42 61,68 50,50 Sau 14 ngav gdy ban, hdm Idgng earotenoid dao
ddng 6,49 - 7,0 mg trong eac ddng va the hien eao hdn 0,21 - 0,46mg so vdi cdc ddng nen tUdng dng 3.2 - 7.0''ii. Trong dd, ham Idgng chlorophyll trong la eua hai ddng D3 va D21 dat cao nhat lUdng flng 11,9 va 12,7 mg vd deu cao hdn 5,3 mg so vdi 2 ddng nen tUong dUdng 44.5 va 41,79o. Ddng D14 cd ham lugng chlorophyll dat 9,8 mg va cao hdn 3,4 mg tUdng flng 34,7% so voi ddng nen C7N (6,4 mg). TUdng tU, hdm lugng nUde trong cdc ddng chuyen gen deu cao hdn td 18,1 - 22,9% so vdi ddng nen tUdng flng sau 14 ngay gay han. Ket qud nay eung tUdng tU nghien cflu eua Gubis va cdng tac vien (2007) cho thay, cdc
cdy thude la chuyen gen P5CSF129A tdng cudng tdng hdp sac td carotenoid va cd chlorophyll cho t>' le cao hdn lan lugt la 63% va 52% so vdi eac cay khdng chuyen gen cd ty le thap hon lan lugt la 36%
vd 31% sau 12 ngdy xfl ly ban.
IV KET LUAN VA DE NGHI 4.1. Ket luan
Ket qua danh gid khd nang chiu han cua 3 ddng ngd chuyen gen ZmDREB2A the he T6 thdng qua thi nghiem gay han nhdn tao giai doan trUdc trd cho thay cdc ddng chuyen gen thich dng vdi stress han tdt hdn ddng nen the hien cac ehi tieu ve hinh
•CD cniKhoa noc Cong nghe 'id'-g ng^^!eD i :etNam - So -Jj) IW2020
thai Id, dde d i e m n d n g h g c vd cac yeu td cau t h d n h nang suat d e u cao h d n d d n g n e n . N a n g suat cd t h e cua cac d d n g v u g t so vdi d d n g n e n t u o n g d n g t d 25,7 - 33,8%. Sau 14 ngdy t r o n g dieu k i e n h a n , h d m lugng mgt sd ehi tieu h d a sinh n h u h d m l u g n g d a m tong sd, lipid, proUne, h y d r a t e a c b o n k h d n g eau t r u e , carotenoid hay h d m lUdng chlorophyll t r o n g Id eac dong chuyen gen d e u tdng vd cao h d n ed y n g h i a .so vdi d d n g n e n . Ket q u a da c h g n Igc dUdc 2 d d n g chuyen gen ZmDREB2A la D 3 vd D 2 1 t h e hien k h a nang chiu h a n cao h d n so vdi ede d d n g n e n k h d n g chuyen gen.
4.2. D e n g h i
Hai d d n g chuyen gen D 3 vd D 2 1 can tiep tuc danh gid cdc chi tieu d a p flng dfl eac tieu chi d d n g ky khdo nghiem ddi vdi d d n g ngd bien doi gen va sfl di^ng lam n g u d n vdt lieu p h u c vu c d n g tdc c h g n tao giong ngd bien ddi gen ehiu h a n .
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Evaluation of drought tolerance and biochemical components of ZmDREB2A transgenic maize lines at pre-poUination stage
Nguyen Due Trong, Doan Thi Bich Tliao, Nguyen Xuan Thang Abstract
Three ZmDREB2A transgenic maize lines, including D 3 , D14 and D21 of T6 generation and three non-lransgenic maize inbred lines C436, C7N and V152, respectively were evaluated for drought tolerance and some biochemical indicators at pre-pollination stage in greenhouse condition. The results showed thai under 14 day watered stress condition, the leaf characteristics, groivth duration, anthesis-silking interval, morphological traits, and some biochemical components such as total nitrogen, proline, hydrateacbon, carotenoid, chlorophyll of all three transgenic lines showed higher significant difference as compared to non-transgenic inbred ones, in which, the grain individual plant yield of transgenic lines was increased by 25.7 - 33.8% in comparison with the correlative non-lransgenie inbred. Two out of three transgenic lines (D3 and D21) with high drought tolerant effect are selected as potential materials for breeding of drought tolerant C M maize varieties.
Keywords: Maize (Zea may L.), gene transformation, ZmDREB2A. drought tolerance Ngdy n h d n bdi: 10/4/2020
Ngay p h a n bi?n: 2 5 / 4 / 2 0 2 0
NgUdi p h d n bien: TS. VUdng Huy M i n h Ngdy duyet d a n g : 2 9 / 4 / 2 0 2 0
