TS.

NGIJYEN MINH CHON

L\;!, {
(: ,t- 4 61

GI.AO TRINH

;\
CFIAT DXETI HOA SINH

ru secg DAr HQC cAN THo

?005
 NIUC LUC

N'ii dunu T
t-ttls _

l,t)'i rrrri' diu

Nlgc lyc

Olrtrottg 1. Lrr.o'c sri lrghi0lr r:ri.u vii cl,rc lill:ii ni6rrr r,0 chit diiu lrt)a sirrlr
Inr'<irrg Ilru'qc v{t
l.l. l.rrtrc srr rrulriirr c(rtr I

l.l.l. Auxin
I

I
L 1.2. (litrbcrcllin ((iA)
l. I.3. Cl,tol,.irrin ......
5
l.1.4. Atrscisic acicl (AtlA)
6
6
l.l.(r. llrlssinostcroitl (illf ) .. 7
l.1.7. Sirlicl larc (.tA)
7
l. 1,8. .lasnronatc (SA)
8
| .2. Cric lilriri niCrn crr lrrirr r,ii tlrrrirt ngir. ...
tr .l.
1r t . \':.. I 11.
r cu eilu dol | ^. ,.r I
\'()l n),.)l clliil rlicju lrira sirrlr lrrrrit:-g tt
1.2.2. Cric l<llii rriCrrr vti tlrrrit ngiL... o
1,2.2.1 .ll.rrrrrrrrc tlirrc v.r (l'l,rrt lr.r'rrro'c. pll't()ll()r'*.rrc)
. .......... ()
| .2.2.2. Cliit sirrh trtriirrg tlrtrc vit (lrlant
Lrrrirvih sutrtancc) ()
1.2.2'1. Chrit tli0rr ltoi sirrh trLro'g tlrrrc vlt (l)la.l er..rvtlr rcgrrlutor,
Ir(il{).. l0
1.2.2.4. chat irc clrc vi chat lijrrr itri,ir si,rt, tru.ing (rnhibitor
vir rctarclirrrr) .. l0
(.lttnrrrg 2. l)lrrrrr'rrg plr:i;r tr.ich, lhrrrrh lgc r,ii xric rlirrh chit sirrir (ru'iilrg
thuc viit
2. . l)ltrttyrrs phtill li, tr[clr ,.
il
1

2. l. | . l)hrLong phrip lilrul'fch tlilr ....

ll
1l
J. r.l.
^
|
t,.\' tIrclr
' ':lriirrg
ll
rlurrg rnoi l')
.t r r r , .r I i
.-.1.!.|. L nrtil tI Il tOtl l)
2.1 .2.2. l .y lr'ich
t2
2.2. 'l'inlr loc diclr tr.ich ..
t3
2.3. l)inh lrrrrn_g chilr sinlr trutirrg thu.c vlit
.la | .r' , i. l4
1..r. t. Snttt trilc.llshi['n] (l]ioassa1,)
l4
1.3.l. L sirrlr tlr'rc rrgltiCrrr irrrxirr ,
l5
2.3.1.2. Sinlr tlic rrglrii.rn gihber.e llirr
... , i l-5
,:.J. |.J. Srnlt tt'ac rrglriCrrr c1,tol.;.irrin
'r -t 1 | t' t ! l6
t.).t.+. billlr Irilc nglrrCnr abscisic acicl ...
l(r
I 5 Sinlr tric nghi,irn crlr,r'lcnc
i1 r Irl.(r./ .t.Slnll, Iriilc l7
: ,,^
f.J. ltglriCrn britssin(,stct1)id .......
1't't rl r. ! l8
L.J,1-. I l{)il lV tf'lC lllltL-nl
'\ -t. I r\r ,, r.^ , i, l8
j tlrirt trrfrr ctr,it.sirrlr
i ; l)rrrlr lrtr.trrrl e tlrr'lcrrctfrr(ing tlrirc riit trarrg siic l,l klrr)i l)1r0 .,,
l t8
:.J.:.1.
I ) r ) rrr | | : tli
tlr.rlt-tlicrrlrurr stttlr [r'u(rnlttlrrrevil brirrglll'l.r'....,......
;l;:::lrtll lrtj.'rr
t.J.J.4. Srrrtr tliic rrgltii'nr nrierr dich lroc ...
|lJ
l9
2.1.3. Xiic clinh cu0i cirrr_u
t0
tl
2.4. Kdt luan l9
('lttnrtrg J. Cjiu tl.ric h6a lroc, sirrlr t6ng hg'p vri :inh lru'ri'ng sinh l;i
cria
ciic rrh<inr chiit di6u hoa sinh tru'6'rrg thu'c viit al
LI
3.1. Arrxirr 2l
3.l.l. Sirrh tOng htrp auxin
l. | .1. (':rc rrrrrin plrri biCrr 23
-3, | .3. Nlirirrg iinh hrrtirrg sinlr ly 25
3. 1 .4. Str ltlr/irr ltr'rv auxin . 27
3.2. (iibbcrcllirr ((iu\) 28
3.2.1. Sirrh tirrrg lrtr;.r gibbeLcllin 29
3.2.2. Nlrirrrg iirrlr huring sinh ly cua gibbcrcllirr 34
3.J. Cr,tokiniu ..... ,
35
3.3.1. Sinh tdng hop cytokinin 36
3.3.2. Nhfng tinh hr"r<rng sinh ly cua cytokinirr 36
3.4. Abscisic ircicl ... 3ti
3.4.1 . Sinh tr>ng lro-p abscisic acid 38
3.4.2. Srr bit lior.rt cLia abscisic acicl 39
3..1.3. Nlrirng iinh huong sinh ly cua abscisic acid 39
40
3.5.1. Sirrh tirr.r-u lrop ethylcne 4l
1.5.2. Str l<ich thich tdng ho-p ethylene cua Auxin 42
3.5.3. SLL irin sinh cthylcne do stress A1
+-)
3.5.,1. Nhirng lirrlr lrtrtilrs sirrlr ly cul ctlrylcrrc .t -)
l.(;. llrassinostcroid (Blf) . 46
:
J.6. l. l'hatr lour va eau tr'[rc lroa hoc 46
3.(r.2. Sinh ttirrg lrtrp blassinostcloid 47
3.6.3. Nlrirng drrh hr,Ldng sinh ly ctia br.assinosteroicl 5l
3.6.1.l. Arrh lrtrr'rrrs cuit lll{ lCu stL sinh tr.tLting nghiOng -5t
1.6.3.2. Anh hLrrrns ctia lll{ lOn srr vuon dai .... 52
3.(r.3.3 llll ciirr tlriet clro su phdt trien binh tlrLrdng cua thuc vAt .... 52
,: clriu vrii di0u
-Srr cliong
'1.6.3.4. ki€n l<hirc nghict.cua m6i r.rtrtrng, tirrrr
li.lriine siu bCnh vir tinh chong chiu vcri thu6c co 53
3.6.3.-5. l(iclr thich sLl sitrh t6ng hc-rp ethvleue 54
3.6.3.(t.l(lrir nirng [rng dung ciia brassirrosteroid 55
3.7. Salicylatc (SA) 56
.1.7.I. Sirrlr torrg lrop sllieylic ircitl ... .....,.. -56
1.7.2. Anh hrLt)rrg sinh lj 51
.l.tl. .lasnronirtc (.lA) -5 ti
3.8. L Sirrh torrg lrtrp, clruyen lroir vi'r vlin chuyCn
-jasnrtlnaLc -s8
3.[i.2. NhiLng iin.h huring sinh ly cua jasmonate 59
3.9. ('iic chirt cliCrr lrira sinh tfuous klrirc ..,.... 60

Chtnntg 4. Vai trd ciia ch6t cli6u hoa sinh tru'ri'ng trong siulr tru'ri'ng v:i
phril triirr rril llru'c vil (rl
4.1. Dii:u khiin srL niy nrim cua h6t va su lrhdt tri€n ctia ciiy con 6l
4.1.1. Arrh huriu-r cr'ra gibbcrcllin vd abscisic acicl 62
4. 1.2. Anli hurrn_g ctia cytokirrin oz
||l
 4.1.3. Anit hu'tin-e cr.ia c'thylcne
4.1.4. Arrlr lrtring e Lra nhirrrg chit lihic 62
..i l-.r r' |
()J
4.2. Sr,r thlnh liip ru r)itr utnn -.ttI cllllll qlilnl
,1.3. MiOlr tlarrg 65
.i..1. Arrlr lrrrrirrr cila 66
clrat cli0u hixr sinh trrrriult lCn clLrri tr.irrlr li'io hrrii......... 6(t
('lttrtntg J. \/ai trr) cria clrit di6u
hdl sirrlr tru'ring lCn c:ic qrr:i lrirrh sinlr
, siirr cri:r llrg.c vit
(rll
5.1. 'l'nl ll()u ....
< | | A.-r | - : ; 68
r)rr()t,re c.il rrrrrrg ^.
:.', l.i ir. ,r,1,:' lcrr ru rrr0i tr.rro'g li,'srr Iluit r.r.icrr sirrh s.rr ... 68
l. (Juitnu li\' (plrolrrPcriorlisrir) (r.3
< I r a ir
.).1.1.1. Irr -l
lll\t lrurr (Ve r.rrir lizulitrrr).
a r ,1 ., : 69
J. I .1,. .\l.l ltl(.)llg tiu ltoil ...
<') | | .l 69
--.i../\nrl^ tru(rns cuil clrlil tliCtr lrixr sirrlr [r.rR]nr lirrsrr lrLorrg nrirrr lroir, l<iclr
tlriclr I t'r rrc ch0 tr6 hoa . . .... _. . 10

] ],i, !,:lliil'i t.,#l I i:j l'iil u l;'::l :,Iiil,;, lli* ljx ; l; :i :;

;,
:; 10
]]l l:j,l',":tltirrriisu.l)triirr,.i0,i.t,,,,,,tr,utrrlirc,t,;;;..:...'...'...........
.r.-r.1. Lt);lt (tlLiu lrr)il sirrlr tr.urirrgYii srr tlrd lrielrr
7\)
gitii tirrlr.... ll
5.4. Srr nurs .. ..
'1
r'. .
: ,r r (rtilt ' i |
.)..r. | . l)nitIl ll(.)c cull su rlltlg ..
72
5.21.2. Sirrlr l1i cua str rullg ..
12
Arrlr lrrrrirrg crirr .rritrr r.rri..1-1 gc11 r'i rrrri'irrrr'c,rr0,ri,,rr,i,,:n,,*
i).4.t.t.
-l I i /\l lr liu(lr_q
cr'ra clrat diCtr hixr sirrlr tru,drrg l0rr str rung
7?.
........ -t't
I)
5.5.Sirrlr|,ctiastrclaLrtr.iii.sitl|rtr.trtilrg.p|uittriCir'cl,ilir,ariirr-etrrii
/1+
5.-5. I. Sirrlr 11 cti;r srr. tlirr tl.lii . ... 1,1
aa1
.r.l.l. /\ltlt ltll()tlg e ltil clrAt tli,,1tr lrir sirrlr tr.ttrirrr. rcrr
r:.. slllll
..:..r tl u(tllg
,
-r vil l)llal
tr.ic' cLia lrr)i tr.i
5.5..1. liir Iirrrlr lroir'avir lriii harrr lroir chlit ........ 74
75
.r.-r..r. )!l ctlln cLti.l Irat ...,
NgIrr srr rlurg rrhi ....
i 5. Ciay
I5.5.6.
lil su. t.\lng trii .... T'
76
('hr,trg 6. \':ri rrr) ciia crr6( tliiu trt)n sirrrr rr'.ti'rrg r0n qu:i trirrh rilrarrg
. hg.p cii:r tirrrc vAt ....
(r.1. L'lrat ciin sitrlr tr.u,tlng
(r. l. l. Nhiirrg
.. .......................... 17
clriit ilc ch6 sirrlr r0ng lrop gitrbcrcllin
(r. l.l.l. Nhrrng ltt.rp 77
chit onir.rnr . . ..... .... 11
6. L L2. l'1,r'irrritlirrc
(r. L l.l. 'll'iirzolc 77
6.1.I.4. Nlriurg chiir lilrac 7n
6.1.2. Nhutrs clrlit ciin sirrlr lrutirrg lihirns ilc t9.
chC sinh tOng hqr gibbcr.cllin .. 80
6. 1.2. l. Mor'lthirctirr
6.|.2.2. |)ikcgulac 80
8l
ctlr1,lcnc
til
6.1 .2.5.1)I1 xuir ,:rr' 8l
ir cctarrr itic
82
6.1.2.6. l)irr xuat cira ircicl bdo.
82
(r.2. lJng clurrg cun chiit ciin sinh tr.u.ring
82
6.3. l\l0i licn tlran gii'a chiit sinlr triltiue cay tLong Lr.orrg quii trirrh cluang ...
lrrrp vl\ s.tr phan.chia cria chiit cl0ng hria 83
6.4. L'ric vin clC vC phdng tr.ir cir dai 84
6.4. | . l)huort,u phiip plrr)ng tr.iL cd .. . ..
84
(r.4. | . | . Ngirrt rrgrra, phitng Lr.Ll vil uho cri .....
ri4
6.4.1.2. Qrran l! co dai . 84
61 1 .(;llti rhigu v0 phorig rrir cr),tri:rng hoa chrir 84
6.,1 .2.1.
'l lrtrdc cri cti
liic dulrg gi6rrg uhrr IAA ti.5
,l
(t.4.2.2. Ntriurg clriir irc clre tfing hop gibbere llin .... 85
(r.4.3. S1r iLc crlrC c;ui'r trirrli sinh t6ng hgrp, quang lrgp vA
lr<l lrdll 85
.i
(;.4.,1 ,l. NltiLrrr: chtir ch0 lro lidp (MAA, dirtoseb, brorrroxynil)
/ . ^ ^ L'lrat
6.,1 .1.'2.
.'' : fre chci: irc
rlrrarrg lro-p . ..
ti5
85
,f ) '' l\rl-:..--
(r.4.-t.J.
/-
(r | l, , ,it { il
4.1.1. Nlluntl
Nhurr-e cllil(
chli( tlc
uc eelrc
lrc quii
u1li trinh silrh ltllrp
tdrre lrrrrr
hop 86
,1
(r.4.4. ('rirrg rrtrllrC <li lrrrr,Crt ,:
vi lirrlt klriirrrt tlrrrrr. cir ri tlrr,rt. r,it har. (.r() ....... fi6
-l'ui
li6u lh:rrrr ltlriio 88
 Lo'i IVIti'Diiu

itirt trinlt " Chril f,lii u I !od Sinh Tt'ttrrttg 'l'lttrc VQt " lir tttdtr lpc giti'i tltitu vi
G
lich su' ngltien cilu, phtroug phrip ly tric'lt, cau tao hoa hoc, .sittlt tong lnp, vai trd
sittlt lnc vd co'cht ttic cllrng ctia chdt diQtt lrcti sinh Inrr)ng thu.c vdt. lvl1rt hoc nity
c'fitrg giri'i tlti0u vi klrci ndng ti'ng dung ctia c.itc chh diiu ltoit.sirilt !rtriug tntng san
.ttttil ttitrg trsltit;1t. Nt) c'tittg lit rttirr lt4tt' c'trrtg t'ti1t tthii'rtg liiItt tlti't, c.titt tlriit r.lto
ttltiittg tt.gt\tth sinlt 11, llt4rt'vdt, klrca lnc <.tiy tringvit sinh hoc plrutt tri,.

Gititt tt'inlt ntiv tltrutc'viit ,1,;1fiuc'vu clto nlru c'rit,t dito tutt cri uhdn ngtittlt
t'6ng rtgltt.sitth ltoc', lttt, rtltiAn trit tti nltitrg ngvd'i trgltien cti'Ltvi tlttrt,vtit cliu ctj the
llrrtttt kltttrt tltntt. Nii tlrttr,q c'ltrrrnru lt'irtlt ttir.tt g,iritt I't) sttttq ttltirtt! liiirt tlti.c'rritt tltiit
t'lto .sittlr viin lu2c'xottg trritrr tltil. ltui citc rtgtirtlt tt6nt ltoc, trrittu trot t,ci sinlt lrlc.
Sittlt t'i0tt c'oo lu)c'thtpc ttgdnlt ttdtrg hoc t'it sinh lx.tc diu c'<i tli tltcttn ftltdo tirio
lrirrlr ttit.v.

Dti1, l; ttitr biAn soan dtir tidn vi volt klrcng th,i trdnlt klfii thiiu r6t. 'l-dc giti
xitt cltdn thirrlt nlrcin tlhil'ng tl6ng grip c'tio tloc giri iti ttirt ttii hdtt sutr dtroc bi srutu
Iroin tltiett lro'tt. xin cltdtr thdrrlt c'am oLr pho gitio su.tiitt sl L0 vdn Itr.d, lien si
llu.inh Tlttr llor), tiitt si iVgtrin lJuo Titiut, !ltuc sT Larn l,,lgoc, Plrtro'ttp1 r,it tltctc'sT LA
Vutt lli tlti t'i nltiitt t1 kiett ding gtip tlttl bio trong viQc hi0n sottn t,d clrt'trlt stia gido
tri rt lt ttut'.

('itt tlt<t, rtgit.t, )! tltitrg I ) rriutt )00.1

Nguy€n trlittlt Clto'tt

 ('lttnttrg I
L,IJ'QC SU'NGI lltrN CU'tJ VA CAC t(r-tr\r Nl[1M Vtj
CI.IAI'DtEU IIOA SINI-I ]'ITU'ONC 'fI-IUC VN 'I'

('lrit diitr ltoii sirtlt tlr-Ltrrrg vrii nltltng rrting d1i clrc thip dir cir klrri nirng cli6u hda
nhic\u lrrrh vr,rc sirrlr trrtrirrg vr\ phiit tri0rr cria tlrr,r'c valt til uiiy ntiirn d6n lio lrriri vi ch6t.
Ar,rxin lti nhilnr clriit cliiu hori sinh truong dau tien di ihruc phirt hien. Ngi\y nay, sdu
nh6m clrlit di0u hor\ sinh tltrong thpc vit cld clrrgc cdng nhdn. BCn canh iuxin cdn c6
gibbclcllin, cylol<inin, atrscisic acid. ethylenc vir [rlass inostcloicl. Gin diiy. salicilate vi'r
.i:tslttottitle citttl', 1l;11',,, tlttt.rc rcttt ttlttt ttlrt'irr11 ttltrirn tttr'ii trirr e lr:il rliiu lroi:;irrlr llrtrirrl,, vir
rrlrtirrg rrglriCrt cittt cir biitt vC s.irrli lro:i. sirrlr ly va sirrlr hoc 1llr0n lticring dat <lrruc,rrlriCu
tlirinli Lpu. Moi liCrr ht cua chAt di0u hoir sinh tlu'crng vdi sy'niy nrAnr. sp phdt tri6n cua
i .-
ciiy con, sU tao rC, rniCn tlilng. srL phat dpc. sy chiu. sp' liio hori. sy tro lroir, su' l'ung, su'
diu trrii, su lllriit tri0rr ctur triii, srr runs tlii non, sr.r chin, srr l<ich (hiclr rurrg tliii. su lao
ctl. (lttlttle ltr.rp vir plrorrt tltr cr) clui clir clrrtrc [riCt. Lliic [r'ng ih,rrrg chlit di0u hor\ sirrh
lltrrirtg ltrlttg n0ttg IrghiQll vu vir;c tlrurrrrg nriri lrriir ching ciirrg cli'i trtr thi'rnh hiCn tlrlrc,

l. l. Lrrot sri' ngh iOn cri u

l.l.l. Auxirr
Vicc phtit liiCn la ar.rxin dii <h.rgc l)alrvin (1880) khao sat tr'€n
luqng quang lriQn
lrrrtlng dorrg. i)n-e tha;'ngc1n clidp ti6u.hrLong v6 ph(a co duh sang vir cho ring anh sing
dir liiclr thich ngorr rliCp liCrr lrurirrg r,C plril rlo. llirng rrlriCrr thi rrglriinr tlrtrr gilirr dirng
.- :
nrdt uirp clrc chop dicp ti0u ha1, cdt no di thi diQp ti€u khdng con hu6ng ve 6nh sdng
n ira.

Salliorvski (ltltt5) clii plrrit hi"^n indolc-3-acc(ic acicl trong nroi truing l0u nrcn.
Mii rlCn nhi0u uirur r'€ sau chiit niy cirng clii clrroc tiu thAy trong nrd tltLc viit. Ngiy
rrirl'. clrit niv rlrtr.rc biCt rrlrrr li clrit <liCrr lrilr sinlr llrtrirrl,,(luiln lr'(.)r)t'llrrrirc rrlrrinr lrrrxirr.
rr(r currg eir lic!rr tprirrr clcrr rrhiirr clua tlinlr sirrli ly tlong ciiy. ltothcrt (ltttt4) dii khiurg
. .(
clinh lai vii tidll trrc circ thi nghi0rn cua l)alu'iu cho thAy liurg tin hiQu cluang huong
rkrng gay ra srr nqhiirrg rlutrc ltiCrrr so(it tt()ng tC biiu nlrrr nr0 cr'ra di0ll liitr.

. liitting (1907) cl?i rrryc luo, ng anh hu<rng cua vet cdt nrQt plrir len cliQp ti0u cria
y'crr rrrirch (lvenu) trong nr6i tnr(rng biio lroi tlo iinr dd nhirrrg vit cit b0 urirt khong bi
lilio. h.it rluii cho thil' khdng cri irnlr hucrrrg cua nhirng cit ben l0n t6c dd ph6t triirr
"6t
vii str clalr u'ug cr)a thc.ddng irnh sang b.lt chip cl€n rrhimg vi tri cit so viri lrrLorrg cua
iinlr siirrB.. lritling clro lirug chat l<ich llriclr tlrrr;c van chuyCn cpra clrat s0ng r,ir rli chtryCn
rlLrtnh vCl, ctit. Olrg cirrg suy d-oan rirng stL cliip [urg tiic dQng iinh sang clrrrlrg tirrlr clir
xay ra brii vi arrlr siing dii sirll xep chiOu lllran crrc trong nhinrg t0 biio cua chrip cli0p tiOu
vii clrit liiclr thiclr tlutrc cli clruyCrr tit nlrriug ti lrio criir clrtip cli0p ti0tr chrr.rc clri,ju sirng
rutit llhil dCn ''( nhirng tC' bao u phdn trong tr-ri phia drriti.'l'hirt lthong nray nrtin, uhirng
-

cpran sirt cua Ong da kh6ng chinh xac bcri vi rnQt vich ngirn su van chuyOu dd klrong
lrao siiv tlrtrrc lrinh thr\nh.

(;hh) ltiDlt( lnitIrttt ll,\i \rttlt lrtntn),. Ihti Irir

 ,Trr l,rl/r irlr r.( 14^AK
.ud di,(11, t,ilu. Srr rrgfiiirg
S'tr l.q,ltir,rtg
t't ru d iilt Kh6ng vrii vAl ctit t'i tfu ditp tiiu
tiiu nghit)ttg t'titt !tlt It t stittR l tll l'(l L'ul V(
diilt riiu bi pltio rri i

Durx,in (1880) I;itting (1907)

Kltr)rrg cti :;v

nghiitr: t'tttt 'rArdrw
rlillt rititt klri Srr rrg/rir;rr1.1 r'rrrl
DUf N ntt(u tliri l,r litltr lilri
trtiirtg gc'lutitt
ltltitt tt)i (llft.tc'(l(ll ,ql I l(l
duth vi t
ltlr rit gtic'
t'uu I itlt I it;u

lJo.l'sart-,latrsan ( 191 -l)

Atth stirrg 'tiittrg ttii 'li'ottg t6i

I.rong toi
Klfittg cri ,ttt.tinlt o
trtrury c d di[p
Srntgltiling ,\*.sit r
titu klti chltp
lt'lt()tlg I llLtttK
cua digp tiAu
dvtt ltit'lt rr'ti cttu di01t t i0tt
vQ lthitt klrc)ng
tlit clfip klt,i clttip hi
dtLrtc' t'dl ri'i L'ul r(rl (ll(oL'
ditl lri lrti

I'tul(1918) Sotlirtg (1925)

I littlt I . L Lttttc sti' rtglttttt c'itu t(; r ,\r,, lt'in tltll) ttlu lrt (it L'lti,l)

(irio liinh t h<it Drt t ltoi Snth littottg I hrtt llit 7

 lloyscn-.lcuscn (1913) cho lirrg chit kiclr thich gliy la brii rinh siitrg cri lhd dtrgc
cli clrtrl 0n xuy0rr cpra v{t thC l<hdrr,e co su's6ng. Orrg cli ch(rng rninh cli0u rriry bing chclr
ciit chtip digp ti0u lvena vir chCrn rndt nridng gelalin giirn clt6p vl lllrAu goc. Klri phiin
chrilt clrrrrc chiiu sirng, phin dtr<ri kill gelatiir di cong cli. Kh0ng nhr.L thi rrghi€m cua
Ititting. ting cirng cirt nhilng v€t o uhirng rnirt khdc nhau cria diQp ti€u virr clr€rn ntQt
nric{ng ruica vt\o clC tao vriclr ngirn. Khi v6ch ngiur dugc clirt plria khdng dttgc.chi6rr sirrrg
ctra cliQp tiOu thi l<liCrng c<i hi€n trrong cong.'l'uy nhi€n khi n6 (ltL<rc dirt.r,'C phia siing
cr,ra cliCp ticu s1r cong xiiy la. kdt qua cho thiy liurg tin hiQu clii clugc trtry0n xudrtg clua
phia tlong toi cua cliQp tieu vd cl6 kich thfclr su sinh tluoug coug.

l'aul (1918) di klrang tlinlt lui rrhring phrit hiCn c[ra l]o;'scn-.lcrtsctt ri cho tirlg
co nrfit ch|t cri khri nirng hoa tarr clii san sinlt ra tlorrg di6p ti€u vii didu khicln s1L plr6t
lliCrr cir:r tlitlgr ticlrr ,lr'",,r, . N.1tr cit clrtip tlitrp tirrtr r a tld clrrip ri1 rrglriCrtg tttol lr.rrt e tto lr0
rrtiL cit trrrrg tui tlri clicp tii'tr sC cottg vc plria l<lrOng cti chtip.

Sodirrg (1915) (lli tri0n l<lrai cong tlinh cLra l'iral ['ing chclr dirng thi nghiQnr sirllr
Irtrtirrg llrturg rlrrlr tlerrr sr.t vrLrrrr rlii cirir diill tiitr.,l lr'rrrr ltoltg tr)i. Nertr tlierp lir"rrr bi cirt
rrri, tlri srr vu(ln rlai sC tli girinr. l(hi ctirt dinh cliQp ti0u trtr lai thi sr,t sirrh tltrritrg tltirrtg
tuhu ban tlatr tluo-c phtrc l16i.
-'n
'_ri'
lll
( tit
fr
chip Ddr chrip ltn kltir
I
D(jt ktk;i agIt
uyiar l-1 git) lt'o l<ti diep tio|,
plntc ltii str sinh truorrg

tlinh | .2. 'l'hi ciro I,Vettt ( 192(t) cho tfui.t' cri ttrr.it c'ltdt ltrsri hoc ti' cl6p cliip tiAu
trg,ltigtrt
bi cdt kich thic'h.srr plrdr triAn tro'lqi cila diQp tiAu bi cat mdt cltdp trong toi

Wcut (192(r) dit tlru rltrtrc rnot chit lroa lroc hoat clQrtg til clrtip cligp tidtr Avena
biuig circh tlirt nhilng ch(rp diclp ti€u uiry,tr'0n khdi agar'. Sau mQt th<ri gian, bo nhlLng
cltrip dieAp ti0u. vi cirt agar la ttng khoi nho. Ong dir thiy rang kh6i agar nay chira durg
clrit lrorr lurr tu rlinlr chri;r drrrrc.it,1,-i l,i.lr tlrich str;rhiit triirr tr'<.r li.ri ctra tliCp ti0tr l<lri
clii dur,ic tlut tlcrr rrlrurrg llrirrr ctrt datr.
\.)\
tt \
tt \
tt I

^^3,,
,)\y'"
,,.t
'''.
t:t'
\n
"ri t,

tl

Ilinh 1.3. Sinh trdc ngltient ti)u Avena ctla Went (1928)

(inio Itinhl hLtt l)rirt Iloi.\nlt Irwttty Il:lac Iit

 Wcrrr (l()2ll) ciing tlti 1ilrr'rt triCrr rtr$t plrrrrrrrl,, grluigr rlt. tlirrlr lrrqrrrg clrit diirr holr
, i1lt ttrr(yltg lri|.'rr tliCtr trurrg rnirr. t)rrg tlriil tirnr.l cir stt lir-1tt ,;trrtrr g.iiilt stI eollg cirl cliilt
tiCu r'<ri lLrt-yrrq cira clrit sinh tlur'rrrg tlrlrc vit lroat ddng. NlriLrrg phrit lriCn ctra Wcttt clii
l,.icI tlriclt rruurlr rrrer viic nglri0n c[Lu clrit siulr tltrrinp thuc r'1t. 1'iL tlri rlglriQrlr str cotl!]
cin /r,errrr.'irrdolc Acctic Acid (lAA) dI chLrrc phlit hi'^rr. l)ay'li tlt0t I)ll1lt hierr t'it qtran
llong diiuh tlou su khtii diu cr.ra ngiinh ltlroa hoc vi chit cliiu tron sirth tluting thuc vat.

Kiigl va llirlgcn-Srrritlr (lt).ll1tlil bit tlau vtii .l.l gallorr rrtLtic (iCrt cirlt rrgtttri.
'lr:ii clrra rurlt loat frrLtic llranh l9c, r'rii ri(c tlrt lroat tirrh sirrlt lrqrc *",.t niii bricrc bang
criclr tlirng thir rrghi0rn vi srr nglrieng cria lucrrrr. lltttic tltanh loc c.rr0i cirng cira lrqr dri
tfiu dr.Ltlc 4() rrrg lrr.rp clrit <lrroc goi lii auxirr l] (auxcnolcic acitl). llortg rlich tlich rrtlry
cti clr(ra lrcrerrxtttxirr vi rrgriy rray chrqc g9i lii inclolc-3-acctic acitl. I)ir1'chirrh lii chilt
clutrc Salliorvski phiit hi0n viro nirrrr 1885.

Nirnt 1934, l(0gl vri llaagcn-S.nrith di'i phan lip lnn til rtrcrt [ria vri'l-hirtrautl
cirrrg tlii plrrrn l/ip lnn ti viec nudi cAy lllrizolttts sttirtas viitl nirrn 1935. lVltii tl0rt rriilu
1942. lltilgcrr-sinith vi\ nlriir.r ngLrdi l<lrric.cirng clir pharr lip tltrrrc IAA titrh lilriit til n0i
plrrii nhir crn lrOt bip. l)iirr uil cho tlri1, 1i',u IAA dir tltttrc tittt thiv ti tltttc vat lrio cao.
Vlicgclthrrr t (1966) tluir lir lrarrg ch[rug ring Aurin A vti l] lilt0rtg llltrii li siin;rltitrtt ctra
tIrLc viit trr rrlriCn.'l'ul nhi0rr. IAA tli cluuc phiin li;l tlirr nrrlt s0 ltLtlrtg ltin loii tlllLc vit
vii xLri'it lritrr rnoi n(ri trolrg thyc vit iric cao. Auxin ctra Wcnt c:ir tlt0 Ll IAA. ttrl' rrhiCrr
nlrirng clrat kiclt tlrielr Uric co thi cti trorrg rrlrir'rr.g rrglriCrt cttu siL lilrrruiclr rirn lilriri dirr
tl0n s\rrlup frng ui tiic dirng cira iirrh sang. cri thC lii dart xttit cuil lAA.'l'lruiit ltgil'lltlxill
c6 nguon gric trr ti0rrg lly L1p. iluxeirr cri nghia li\ "glorr."' (tttoc. sinlt lrtLcrng). l(i1gl,
llaagcn-Srritlt vii Wcnt rlir de rrglri su durrg tlruit ngir nriy'cl0 clhrrlr diiu str plrirt hiclr nrOt
clrjit ,1i," hir..t cir lilui rrirrg liiclr thic,lr srL sinh tltr<!rrg cottg cirt tligp tiitr yin tlrach /t,ctttt'

l. 1.2. (libbcrcllilr (OA)

'l'ir'liu ngrfiri n0llg, clin Nhiit llrin dir thiiy'hiCn tucrng c:iry lira cao si-rnr lron binh
tlrrLirrrg. llc., nghi lirug cki la str sinh tr-rrung t6t va se co.rtrQt nriur biri tlrtt.'l'try nhien, lthi
Ir.r ntua clcrr thi nhring cay nAy 1,r'o rr€n lons th<in-e. bitt thtr, hOt ldp.'l'lray vi ln6t mira
btli thu.4O'Xr uiing suat clir b! rrrut tli hiing nirur do tri0u chirng nt\y. llCrrh nriy'di-r dLLrrc
ngr.firi na)nr,, rllln Nhilr lllin goi nhiitr tCn rh,r'a thco triCtr clriLrtg tltrlttt siit tltroc, viti tCrl
tlrtirrg tlLrnll li bakarrac (nra ngr.r). nhonilc (nrri lihirng), ):r'ci (lllIt). sL)rllctltlirc (rlrt1 rni
firn)...I'hrurt ugi quclt thuoc clutrc clirng k\ nra bal<anac. () Vigt Nant. tlicu chflng niy
cirng rit tli tlriv tr lira nitrt.

Vao uirnr 189u. lloli li\ ngrroi diu tiOn clio Ling lrernh l]llilrtac giy ra biri s.rr'xinl
nlriili cua nrQt loiii ninr thuOc chi Fusariunr.(tloli, lti98). Sau'atll (1912) cho rirng.slr
v1o1 drii cua l<'rng li'r clo chiit liiclr thich tit so.i niint. Kurosau'a (1926) chiurg rninh tiutg
u[inh chit cluoc tiCt la b<ri ninr l]al<anac giy ra str.r'uun diii. (lii nrQt loal trarth luill v0
viec dinh <lauh uanr llalianac vi nguoi ta c6 thd thay Iro tr nhfrrg tlarrg lthic nhatr. Vin
di nil,v cla.lutrc giiii tori r,rio lliint lgJl khi Wollcnrvcbcl clat (Cn giai drrrln bit toiirr (vd
tirrlr) frrsarirrilt ntonili.fornre (Slrclclon). vi giai tloatt ltoi\tt totin (hii'u linh) Gibberella
litjikrtnti (liirrv.) Wr. lrrl,nhiCrr str tlrlrrrlr loc clril sirtlt rir clo ttittr Ilitliitttltc [ri tlit ttgai
(; 0lti h(hr;t I
bdi str lricrr rlicn criu rnot chiit u'c chc sirrlr tlrrting la lirsllic:rcirl (-5-rr-rrrrtrlpicolirric
acid). Virrl nirrrt 1935. Yitlrttta.cl:i plriin lup niCrt chil rlirng (irrlr llri cri 1111.1 tirrh lir tlich
l<rc rrttli ttttirrtq tlratrlr trirrre narr (rihht,r'clltt./itlikttroi. Clrit rriry' clii liiclr llriclr srr sirrh
tltttlttg klri cltLtrc iip tlLrng viio r'6 rna lua r,ii rltLcrc goi lr) gibbcrcllirr A. l)at, lir lin rtirr
tiCrr lhtrlt rrgir'eilrbcrcllirr cltLoc tlirng llotrg'lirnlr plriip l<hoa lroc. \'allull vi'r Srrruil<i
(l93ti).tlri tlrirrrlt ctlrrg lt()tlc \ i,-:c tirrlr tlrC lrtli gitrbcrcllin A vi\ plihhclcllirr ll. 'luy nlriCn
dtt clriCrr tlarrlt. trsltiCn c[ru ve gibbcrcllin dii [ri xclI loi. Vio tltiP ni9111 l9-50. ciic rrlri\
liltttit ltoc Arrlr. l\,1)- r i'r Nlrrit lliin rli'i cri rrlrirng rrglricln cLrrr siu h,lr r,-i rlirc lirrlr rliirr ltira
sittlt llrttitr-r-l ctrl !libbclcllic :rcirl. Ciric gihbcrcllic trolLr nlrilrrg rliclt trielt nirrr tlii rlrrt-yc
clirtlr <lirttli ri\ clrring cirrrg rlii tlrtcrc plriit lriCrr ri thrrc viit br'ic cao. Viro rrirrrr 195.1, rrlrirrrg
rrhti rrghiClr c[LLr rrsttiri Anh (l]rian vri cdrrg thc vicn. 1954)clir nluin tlriv nhi'ng dirc tinh
cliCrr lrorr sittlr tttttrtrs cua eil)l)clclic ircid til clich tlich nirrr Gibbelcllo /itjikttroi. Yirt
rri-rrn 1955. rrlrLlrrq nlti kltoit lroc I\4n (la nlrarr clit-rrr tlrroc chat rni ho qoi la gilrtrclcllirr A
vi gilrlrcrcllirr X tir tlit:lr tr iclr rriii llrriirril llurrrlr trirrrg,. tti'lt ( rrlrltr't r'lltr lirlilirrr.or (llg
vio rrirrrr l()-55. rrlri'irrg rrlri l<ltol lroc Nlral llrin tli tlri-r rinq qilrhcr.cllirr z\ clririr bir ltrrlr
clrat plriirr biil tlrrr.rc goi lir (in r. (in.,. (iAq. Ngiv rrirr'. rlibtrcrcllic X. gibbcrcllic rrcirl vir
( iA, tlrrr.it' lriel li cirrll l1t.'qr r'lril. li:r.llc.r (l().:(rl tli
Illil lrit:rr rir rrlrrrrn,. 1r,.,;, \.lr;il trr()n1.1
ItL vtii eitrbclcllic lcicl tlortg tltlLc viit triic c,ao. Ngi'r),rrrn,. ei[)Lrclcllin dii tlrrrrc tinr thly
tlong rtltictr thttc vnt bac cao.'l'aliahashi vi) cdng tiic vicu (1951). dl-i phat lriCn (iA,1 tir
ttirn (iiltbet'ella /itlikut'oi. lVlaclVlillarr vii -l'al<ahaslri (196S) rli-r clel n-qhi ciiclr goi t0n
gibbclcllin A1-A, lrit chip nguorr gdc cia chung. Clch rrii'r'iu cr)n tlirrrg rl6n rrgiiv na1,
clro lilroang l,l(r {ribbcrcllin dii chroc phiit hie n.

l. 1.3. (lvtol<irr irr

. llabcllirrrrlt ( lgll) thi1, r'iurg chit l<hu0ch trirr cua lnd libc cti lilrir rrirrrg kiclr thiclr
t0 lriro tirrtgsittlt trrrrte rtrir cu lilroai ti1'. (iiirr lrrrn 3(] rrirrn sltr Virn Ovclbccl< vi c0ng tiic
l'i0n (l()4 l) dir tkil r'irrg siirr plriinr ttr nhiCn tinr thay lrong rrtrric tliur (nOi nlrii kinu) c<i
f,.f li rrirrru. hitlr tlriclr srr llrnt sinlr lci lrlro lrorrl plrtii rrorr crrir ci tltlc <lrrrrc l)ttltu.tr. yun

Ovcr'trccl,' tit critts tric r ierr 1lt).14) cring plriil lriCrr trong rliclr lriclr llrtl crrir grlrt)i ci clt)c
tltLtrc 1)rtlitlrr. ttartt tttcrt. tttittt l[rir rrri vi [rtll hach Ititrir (Alurorrdtrrecrl) liich thich su'
lrlrarr chia tc blitl lrong rrLrtli .ir pht\i cli rlt\e' cluoc Dattta.l)iirr niy cJro tlray rrlrtiug
t'lril rri.1,rliPlrirrlrotrlrrr', lrorrr',rrlrii'rrlrr;ri l..illrir;rcirrr1,, lrirrlri:rurlllrlrerl;rrrrll.r.irorr:irrr
194-5..llrtrlorrsli.i vl Slirroq rlii Plriit lriJrr ri-rrre rrlrijne li bi\o rnir nrirclr cltrrl rrlrir.rrg clriit
liiclr tlrich stt phan clril tC biro ciiy,thrrrlc [i. lVl illcl lri ctirrgl lr'rc viCn (l9-55) (lai c.lrg l,t)
li stt tlrirrrh llip ri tlilrlr tllrrrh liirr.'1 in (h-lirllirr'1'lirrrrirroprrlin) llru cltrtrc tir l)NA cuir titrh
tl'irrrg cir tt iclt tlur)rrg.thi'rnlr lral'1il l)Nn cLra tinh tlung cir tliclt rlLr'<rc lrip thrrrrlr tlrrrrg.
llo tlr'i clirt tCrr litr'p chr:it rri\'lir I(iuctin b<ri vi nti cri l<lrii nii'rg l<iclr (ltich srr pluirr clria tC
lriro((.'rl,r/'ir(',\r\)lr(rrgrrrirliri tlrrrrir:lli. lllrll ri,..,lrrr:rrir:ritlrr(1,)55)rrlr:lrr llr;ir l,irrt'tilr
,.'rr llrr' rtttr.te slirr sirrlt l,lrrr.q ericlt llr:rrrlr trrrrru rrrtil lrorr 1r,.i1., ctur ltlerrirre ri lirr lrrrrl
itlcoltol. I)ictr Irirr clro thiv rirrg l<irrctirr cti Ilrc rlrrtrc lao thiirrh tir rrhilnq slirr Plriirl Iluirr
la l)NA. Nl illcr (19(rl)tln xric clirrlr rlrloc srin plrirrr giong rrhrr liirrctin Irr nhiclrr tlorrg
lriip. stilr ;rhattt na-r'sau tki clrLoc r2.oi lii zeatin. Su plriit iri.*,r','a clrrit rril ciirrg;rhiii ke ,lei
vtti lto ctnt l.clltirnt rrr i\l iller t 1963). I(i tiL lilri plriit lri.:rr r:r zeirtin. rrrtlt sri lr;r.rr1s l(Ji11
c1'toliinirr lihiic ctrrrg clir cltttrc phitt hiin i i\ clro tlrrir rrir plriirr bti lting trorrg thtrc vat bac
c ao.

( ;kit, II inlt I lt,it I tn'u Il,ti \tnlt It rtnt]' llnrL lrit

l .4. Abscisic rrcitl (A llA)
1

Litr lit ('rrrrts ( 196 l) tli phlin lip rrrtrr chlit ttr (lul bdng chirr vi tlriiy r'irrg nri l<ich
tlr[clr srL ltrrrg, crroug lti b6ng. Criu trirc cuu ltcrp clrit niy tlutrc gr,ri ll\ abscisin I dti klrting.
rltrr.rc xtic rlirrlr. Narrr l9(r3, uhorrr cua Adclicott ir Nl1i di-r lthiin lap llldt chiit trr quli lrorrg
ttt'n vrt tlril tJrrrg ttu c[rug gi;' nr s1r ntng crrorrg. li btrrrg. Ilo di tliic Liulr ltrrt rrlrrirn niry
tlrirnh tiLng lrlrarr. cho thay rirng n6 lt\ rnot hrrp clriL chiLrr lS-carbon vi dii goi lA abscisiu
ll. llirtr ttlttr rlirttt thir'i vrii blio crio vt ubscisirr ll, rrlr<irrr crur W.arcirrg <i Arrlr ciing rlii
plrarr lap drrr.ve rrrril clrirt [r'c clr0 tir lri cua cay lruC (trirclr) trorrg rliCu liigrr rrgiy rrgrin. l(lri
lrrip clriil rriy tlLrtrc iip drrug l0rr ciiy huC rron, n(r tlir uc ch0 sr,r'sinh tltrirng c,ua clt6i
rr1'.on. l)iiLr ni1' {,'i rlin dOn kCr luin ning: lltrp clriit ni}, k\ thc nhiin giiy nr nlien traltg
vli lrcrp chit clruu lir.tlic tinh uay dutrc gtli liitlorrnirr. Vrio rlinr 196-5, nhting nlri nglti0n
cri'u cria phong thi ngliignr Addicott dri cl0 xuit catr tt'irc hori lrgc cua abscisin ll. M0t llin
nitu hiu rrhrL tl0rrg tlrtri. Ii6t tltrii cua Wulciug cQngl tlic viri nlrii'rrg nha nghiCrt c:ilu cuu
ctlrru {r'slrcll tr Arrlr tll'i clro tltliy linil riolrtrirt vi abscisirr ll ll\ cirng nlot ll(.tp clrat. l)i
tltrn qiiirr hria.rilrr dC. r,0 tlanh phrip. nlrirrrg rrlll tlrtLc vlit hoc lr.r\ng.rliu trorrg linh vttc
rrglriCrr c'iLu rr,\r dii dOns j'cklt tCrr abscisic acirl vi [rrio ciio y'kii'n [i't lrriiu ctia lro ir hili
rrglri quilc tC lilrr tlrir sirrr r'0 clrat cliCu lrol'r sinlr tltrtirrg <) ()tturvlr viro niurr 1967. Ngriy
lurr)', tlluilt rrgn ubscisin l, atrscisin ll vi\ dolrtrirt clii rli vuo tlu0n kirrg. l'ir lilri prltht hiQn.
Alln dn cho tlril'tirrh pluin [rti rtirrg llone tlruc vit bic cao vll cri rthilng iiuh htrttrtg
r'0rrg ltin, tlr0rrr viro drl ln tiuh hrtorrg lCn tirrh nricln (trrrtg vii s1t rr,trtg.

1.1.5. lillrylcur
Nlrrirrl, itrrg tltrrrg llrtrc liin clrir clllvlcnc.li bit tlirr lit rrrrr lrtri rrlrrtnlt rrgrriri co
Ai Llip, ho tlir vaclr rrlrirrrg vit. tllu()ng le\rr tliri dC liich thiclr str chirr. Nlrrirrg ngrriti
'lrtrng lloa rli rlot lnL<lrg lr'iinr lrong bLr0rrg kin cld gia ting str clrin cua lC. Ngtxii ViQt
Nrrnr t irrr1,. rl;r rlrrr;, lilrrji llrrrr lrrr' l,,lriri rrllrrrl,. ,lt' l:rlrr .lrrrr lr:ri r';rr. (iir;rrrlrrr (llt('l)
rrlrirr llrir,y lutrg. lrtrr tlrrriil ru lil iirrh siing tli'n clrLirrrg cl0t tu tlrarr rlii gay,ta s1t ryttg stiru
cirl lli gin tigltron slirrg. Ngrriti ta cirrrg llril' lrtri tir rirrlr slirtg tlc'rr cirng, rli giv m srr ciitt
.ui. t',',' rt'rr rir rrr sirrlr tlrrtrrrt nglng [)iit tlrrrirrrg erilr clrili. Nrirrr l')()1, rrlri lilrtrl lroc
Nga NcljLrborv clro liing cthl'lcnrr la tlrt\rrh l)lliur lroilt tinh ctia lr<r-i iinh sirrrg clcn. l(hi
ctlrl'lc.rrc tlir giil rl rrrot rligr irng b6 lra nlru lren vtii nhlnrg cliy diLr llir [.an lrl vring. KCt
rlLrir niv tlr-r rlrLtrc dirng cli l)lriit tl'i0n sinlr lrirc rrghi0nr diu tiCn vC cthl'lcnc cluir tlCn hiCn
Iutrnq lirrr girrnr srt vutrrr dili thiin cr'ra nrl. r.r.ia lirrrr,, str' plrit triCrr [rCrr. r'ij lticlr thiclr s1t
rrglriCrrg lroac sinh [r'ur)rr-e ngarrg tfoug s1t cliip ring vrii tlong lrrc. l)ourbt (1917) dli phat
lric.rr ra ltirrg i:tlrllcnc kicli thich su rUng.. Niinr 1923. l)cnrrv clir giinh clLLuc bi'rng siurg
clrC birrrg pltuorru plriip rlirng siin plriirn tlOt chiy clC l<ich tlriclr srr clrin cr-ra canr vir c1r.rit.
Nirrrr 192.t, l)crtrrv chirrrg rninlr ling l'tlrl lcrrc c:lriuh lii tlrirrrh phirr hoat tinh tlong
nlrr-Lrrs srin pluirrr dtit chii1,gir1"r'a sLr chin. Nirnr.l9l0. Cor.rins cho rirng tLrii c[ing plr6ltg
tlriclr la lrtri I'ich llriclr srt clrirr.'l'r'orrg biitl ciio lrirrrg rriCrr vtii tlir N0ng NglriCp.lauraica.
Couitrs clii cting tlri riug triri carn dri chin srrur lion lrinh tliLrtrng khi chLr-rc tr'lr chung r,<ii
clrrrtli. Nlili tlin 20 ttitttt sau, lilri nay nrtri rlLL<.rc Carrc (l9l.l) clt[Lng nrinh li lihi
cthi'lcnc. lrtlt,r,lcnc dutTc tltitc virt t0ng hqrp vir cti licn rlturrr trii toc dd cria (ptil ttiull chin,
lv40t tlriri giittt uglitt suu, Crockcl vi cting tic viin (1935) ri vitlrr llo)cc'l'lrornpson rLi
e lrti'ng. rrritrlt luttgl cllrvle ttc lit ltot trrrrtre gii.1' r'lr srt clrirr. ctlr;,le rrc e tirrri lroal rl0rrg rrhrt

chirt diCu hoir sirrh tnLring tlorrs nhir'rru c(r (luilu thtrc viit. Giii thuy'ct triy clir clutro uhieu

r )tt)rt lt itth t lttir lt .tu ll,'ti \nlt lr ttrttu llnt< llt

rrlrli rlglliCrr cfitt trltg hQ, Ilialc vr) cQng tiic vien (1954) clii <lirrrg rrhii'rrp. plrrrrrrrp. ph:ip (rric
rtqlrir-'trt tlriclr lrtrp ttlrtLrtg l.,ltrirru rrlriry ctirrr viri ctlr;,lcrre vir tllr tirrr tlrlrl r;rrrg triii ciiy tlir
klrtirtg stirt xrrit cltr lur'rng ctlrylcuc Ir'rrric klri chiu tlC tiro ni.rr s1r chin. Cirng trinh niiy
cirrrg cllrt gili tlruy'Ct rirrrg ctlrylcnc iirrh lruting lCn str chin cr.rit Lliii. Vi'ro nirnr l9-59. sU
lthiit hirlrr ctltvlcttc birug plrr.Lone phiill stic liy khi trong tlrtrc viit rlrroc plriit tri€n clir giirp
xhc clirtlr ctlrylcnc r'<ti nrir'c dd nhd hon gan urol tli0u liin so vrii nhirng llhLrong plrirp
l<lrric clarrg tdn tai. Viri k!'thLrat thich hup niiy. nhlLng c6ng trinh rrghiin ui cthylcne
dii ngril'rtrQt nhiCu vli ethylcne dir th.roc cdng nhiin li chit cliiiu hoa sinh "uu truring tlrrrc vlit
crj nhiCr,r iirrh hLLong lCn thlrc vit tir su nA1, nrinr cua hQt cl0n sq. liro htia va clti't cira cAy.

l .l .(r. llr:rssirr<lstcroitl (l)ll)

edrrg b6 dirrr tiCrr vi Iihti nirrtg cil rrt-ri cr.ia rnilt rrlionr clr.'1t diiu lr<)a sirrh tru'<rng
thtrc viiL rrrtii tli'i xuit birn t,i)o lrirnr l96ll o Nhirt lliin.'l'i'' 4-10 ltg lir trLtri cr,ru ciy isunoki
(Distilittttr t'ace to.\uttt Sicb. l;t Zucc.),3 nhtiur cil lroat tinh li\ Distilirun.fhc:tor At (751
pg). A2 (50 pLg) vi l) (23(r;tg) dn tlugc plriirr liip. Ilrr nhrirrr rriry rlc\u clro hoat tinh rtranh
lrtrtr IAA tt.otrg sittlt tlic Itghirlnr vC srr rrqhicrrg cua lii. l(ra (ricc lanrirra inclirrlti<ln tcst,
larttitta.ioirtt tcst (l..l l')).'l'Lry rrhiCrr, vi qitli han cira s0 ltrqrng Lrich tluoc dir hltting ch<r
plicll dinlr rlanh clr,Ltrc tiLIrg h<,lp clrrit vao th<ri cli€rn nay. Vno nirnr l970, "llrassins" clfr
dtr<.tc ciic rrhir l<lroa lruc M! tlich drLcrc tir pharr hoa ca-y ciii cliiu (l\ros;;ic:ct ttttpu.s 1,,)
ciing cho lroal tinlt sinh hoc liit nranh. Ilrassins gAy sr,r vrrrvn cliii cua lring tlru' lrai cua
cay clitr leltr clcn 155 rttnr ti lii'tr ltrrrrrg l0 1tc/ cay. rrrii lli o rloi clriutg clrr cri ll nrrn.
Vi,-:c ltlrin l6p vir clinh clanh no glrp rAt nhi€u ltho khan. ll'urlc hit rrguiri ta rrghi riurg no
la J)-glycosiclc cLra chit bclo. Nirnr 1974. rnot chuong trinh clirc biit c[ra 136 N6ng
NglriCp M! dir l<lroi clc)ng. 227 kg phan hoa cai cliu tlru rlrtoc tir circ td crng cli clu<-yc siL
tl.trrrg cho ticLt trinh. llirng phurrng phap phiirr tich tinh th€ hoc vcti tia X cki cho thiy
lirng lr<rp chiit rtrrii nriy'li\ nrot lactone sLcloicl vrii ciu tnrc cltrtre tlirt ten kj brassinolicle
(BL) ((irovc & cdng tiic vi€n, 1979). Dtra tr€n khiim phti nay circ nhii khoa hoc Nhiit
lliirr dii ttilth clarrh tlti lli Distiliunt.fdt:tot'Al k\ hr]n hop cua cilstastcronc vi\ 2tl-
Itorcastastct'()rrc. ll Ir\ h5rr. hop cua blassirroliclc vri 2[i-nolblassirrolitlc. A2 vlrr chr.ra
clinh clanh rlrroc vi\ clurrc dd nelri lar 2-clcoxy-ty pe lrrassinoliclc.

l(C tit lihi l]1, chr.rc phril hi0rr. cric rrlli lilrol hr.rc Nhit lliirr vir ciic rrhil klrou lroc
Itlrrrc tlil Plriit Iti':rr tlri'rrr rrlriirr eltirt tlrrrrlc rrltt'rnr rri1 . r,ic1c l0rrr lrt.'l' cirrgi rli tlrirrlr cirrrg.
'l iL tlo Irlrt'ine
rrglriin c[Lu vi I]1. rri rrlrirnc cluir co liCrr rprirn clir pluit triCrr rit rrhrrrrlr.
Nh(rnr chit cliiu hor\ sinh lrudng uiy cla cluclc xcnr nhu la nhonr thir siiu l<ri til ldii vai
tld sinlr lr ctrit itttxitt. giltbcrellin. cytohirrirr. abscisic acid.''ii etlry,lcne drrr.rc pltat hi0n.
Itl4 ,l,r,lrr,.', lrrrr llurr lrrrtry,rrlrr, rr l,r.rr llrr.r, r..rl I,ir,r l'o1t r 1\ ',.,111, lrr ,1r,. 1, ,l,,rr l1 ,1r,.'1,
hlttxt ltL tit tito. lilt cirrrgrlurrc Littt tlri;'tr orrg nlric\rr Ll0 plriirr I<luie nlxru crn tlrrrc riit rrlrrr
ttri plrarr. lti. hoa, hot.clx)i. ttrutt lit vi'r tlriin. Neat\ nay clir co lrrrrr 40 chit t[rtrc phtrt lrieln
tltuoc nlttirtt rrii1,. ;,',,,,* srl d,i lll- vlr clrstirs(clonc tltLtvc xcln lii (luan tl(.)llg rrlrit vi tinh
plriin bti 16ng vir huat tiuh sinlr lroc rnanh.

l. 1.7. Sllicy,latc (SA)

Nhlrng nguoi ily lap co vi thd dan lloa Kj,tlir phiit lriCn ling ki vi vo ciiy li0u
lri dttcrc b"^nh clatt rtltitc cuc bCl vd nhirng benh sot. Vi\o nurtr l828.,.loliann Iluchner lam
Ono lt inh ('hdr lttrit l loi .\tnh Ii 'lng t htrc I ir
vigc rr Munielr. [)irc lan clAu {iiu philr lip nhiing vft cun salicin lri glrrcosirlc cria salicyl
alcohol vii sllicl'latc trotrg vo cay liiu (Wcis.snrunrr 199 l). l{allaclc l)iria (ltt38) di dirt
tCrr cho thiinlr phin lroat tirrh tlong vt) ciry lilu la salicilic acicl (SA) tiL chir latin "salix",
c<i n*rthia lr'r ciiy li6u. Vrio nirtn 1874, siin phinr tlrrtong rnqri cliu ti0n cr.ra SA dir dutlc tr:in
<r I)ir'c. Vlio rtirrrr ltl9ll. aspirin t0n tltr"rtlrrg ruai cLrir acctylsalicl lic acitl dii drLtlc cdrtg ty
llal'cl gitii tlriQu, C(l uhi€u tiri liQu thaur khao, trorrg 116 rrhirng nhi thgc viit lrrlc <ki dirng
aspirin vi\ salicylic acid [lruy'tlrd clro nhau trortg thi rrghienr.'l'Lry nlriOrr, nOn cltfr j rirng
aspilin kliirng clrrrrc \cnl lir rrr0t siin plriirrr tu nlriCn. N(r co the c(l lrieu qrri hiri vi
acetylsalicylic acitl sin sarrg drLtre cirr.rl,dn thiurh saliclic ircicl tlonq lrC tlr6ng tlung tlich
rrrrrrc. Ngiry rrirl srrlicvlie acid (lLr'(Ic triit tlirr nlriiu Ioii ca1 r'li rltLrrc xcnr rrlru lir rrr0t
chiit clicu hori sinh truring tlruc vat qllan trong.

l. l.ll.,lirsrrrorrllc (,lA)
l)crnr,lc r.ir cQng tiic ri0rr (1962) lan rlitr ti0n dir phiin liip (-).iirsnronic acitl
rrrctlrylcstcl lir.diiu thiCt y0u cua Ju.ttttirtttttt prarrdiflontnr (ltt2 liti). Ngriy nay, .lasntottic
rrcid 1.lA) vir tl0rru plrrin lill th0 crirr rrri ( t) 7-iso-.lA lir rrhilng tlrri rlir-1rr elrinlt cint ttlttitrt
.lusrrronirtc rrric rlir rrrtlt sd lrtrlrrg lrin cirir ircitl bi'o cyclopcrrtirrrc cri tlrurrr liC citr trirc
l<hirc clir duryc xirc clinh. hhrii cltru,.ilsnronic acicl clir drrivc nhin la clo hoirt dQng irc ch0
sinh tltnrng cua rro. Ngriy nay nir cli cho 1[riiy stL phin bO lQng tlong thlrc vit bic cao vi'r
rliirr lli tlrir li klra rrirns cua no trong viec gin ting srr tlrc: lri.:n crra nlriir.rg genc tlrrrc vit
clirc bi0t. trorrg s0 116 co nhiLng dirp irng v<ri srL tOn thu'crng.

1.2. Cric hh:ii ni0rn co'biirr v:'r thuiit ngii'

1.2.L Ytrr cirr d6i vri'i nr6t ctrit Oii,u hrja sirrh tlrlii'ng
Nirnt l959,.lacobs clir drLa fa clrcll ck'uih giii ve ilnh huring cria cic h(ra chit l6n
sinh r,..rt vit cltu f iu clu<rc irp clung d,.ri viii auxin vii Lirrlr quang hrLiine rl6ug
(ltltolotloPisttr). ('iic ltltrrrrrrrl ltlnip rri\1 rirr cirrr rlinrg rlCrr rrgil,rrrrl,tlei tliirrlr gii'r rirrlr
lrrLrirrg cirir rrrtlt clrlt tirng trrrt'rng tlrtrc r,at lclrr cric (luri trinll sirrlr tnrr)'ng vir lllxit tliCrr.
l'l:Sl(iS (l'r'cscttcc/ parallcl vafiatioli. l:xcisiou. Substitution, lsolation, (icrrclality vi\
Spccilrcitr,) li 6 clrir ciii clrtrrc tlirrrg tli'tlirt tCn clro lrhrtrrng lrlriip rrirl,trrrrrrg dtrrrrg vrti 6
tlui luit sau:

(l). tui vli triin tloi tutrng d0nq: Chit lthrio siit phiii hiCn cli0n trorrg sinh
StL tOn
rit: vri cti str biCn dOi trrrrrrg tlOrrg vc sd luqng cLla nr'r vui hoqrt clQng co liin cpran. Su
! ,l' \
r.lcn oor nirY I)llilt utoa llal )/ct-l ^
cau sau:
a. lllrn ltrrlug chirt tiulg trrlong thuc vat plriii durrc do chinh xhc tloug rn6, t0 bi\o
lroi]re rrg.irv cir tr rlutri rnfie lC biro, rrtyi cri ctic pluirr frnq xliy lir.
b. Chit clir plrii dutrc Lirn tlulv tlonp. rrlrii'u loii sinh r'+t.
(2). Srt cat riri: Lir stL thclr riri cLra nr0t co tpran, 1116, lroirc nr0t co cplan tri co klui
Itittts siitt sirrlr lit llril lL)ili lruri clrit nuo rlri. Klri cit roi Lrr) phirn rril thi clie tlua tlinh ct-r
liCrr rpran sC bi rrgilrrg t16.
.( Nl0t hrli lrol chat linh l,hiCt cti thC du'rrc tluri, tlrC cho ruirl bir
(3). StL tlral thc\:
, , L ,:, , .
phiru binh il',ttr"s clii bi cir di vri nir c..-r tlrC plruc lrtli Ili ctrc tluli trirrlr sirrlr tluting vL'r
phirt tlicrr.

(irr.io Itittlt ('hrit l)t t llou .\urlt ltttng lhvt'ldt

 (4). Str c0 lip: 'lhtrc hi0u nrot rluii tlinh c0 lip vtii cric lrc: tlrorru plriirr iLng ltlthc
clCn rn ilc cri th0 chiill nhin dtrcrc vii xirc dinlr drtoc iirrh htrrittg cita Itriit clrat ttLrrrrg l1t nlrtL
trong I rc1,th0n-e.it clutlc c0 lii;r.
.i
(5).'l inh rrirrg rluiit: Clro thliy lirng viQc irp clpng lrtiit chrit rtirl'tlortg nltieu lt'tfitllg
lrrrll li\ t tttrne ltr rtltllu.
(6). lirrlr clrrrvirr biit: llriir cltit ni1 grlrrri tlirc 1r'rttrg.

1.2.2. C:ic ltlrrii nif rrr v:\ lhu$t rrgir'

l,?.2.1 . llorrrronc lhr.r'c vit (l'lln( hurruortc, llhytolronttorrc)
llot tttotta tlrlrt' t,tit li trtrjt sirr pltitrr .sirtlt ln)tt cti(t tndl tr; lriu, lttttic' ttt6l nri tldc
biit gci.t,r'cr trrot ,sqr'tlru.1'dii ltotic rnol lti<:.tlong lrortg rrtot ti bdo lutcic'rtr6 rrdrt.ilrt.trong
ttt6l t'rr tltttrtt. Ilttrntona thrrittg cli clttt-t'irt bOn lrong !ltryt: t,rit tir no'i .trin xrttit ,1in noi
I t( )dl .l()t I,t:.

1.2.2.2. (,-lrlit sirrh 1r'rro'rrg llrry'c viit (l'larrl grrlrvlh strbt:rnce)

'llrrrit rrgir lrolrrrorrc (lrrrc viil rlil rlrrtrc tlirrru rrlriCtr rrirrr. rrlrrtrtg ngil' rrrry' rrgrtiri llt
cti lilru),rrh huting thly, tlrC bing tlruit ngti chit siuh tntting thu'c vit (grllnt gntrvtlt
suhturrcc) hlrl,clrit cliiu lrrxi sinlr tlrrrirrg thlrc vrt (plrrnt glou'th lcgulirlur'). 1'0 ch[rc
:1 I
clu0c tC rrqhir.\rr ciLu vc uhinrg chiiL ruiy drL<rc gcli k\ lli0p lloi Chat Sinlr'l'ruung'l'ltqc
Vat Qrroc 'l'C (lrrtcrrrational l)lant Grorvth Substanccs Associatiorr). lo ch[r'c niy rthont
- ,- rl
ll()tr -t ni. ll nt()t liln.

t'lttit :;irrlt tt'tn)ttg tlu.rc'vcil cltrc.rc' d!nlt ugltkr ttltr" sutt;

A'er).1,rro1,,

. - Ni ltltrii lit rttt)l ltttlt tltit ltitrt lt,.tr',ltrttc .:'itrlt titr14 lt<.t1t 11'1111t: tltryt't'til t'i pltdtt
bi rg,trrg d tltqc: t,cil btit: r:ao.
- Ni 1tlrcii t'it lto<tl littlr.sitilr ltoc cldc biet d trittgd() t'4t('rltLil)
- Ni plili dirrg.voi trit c'irtt hrirt tt'<ttrg vilc diAtt lroi c'ric lti(tt llnn.rg:;itrlt l.i, trrsttl:
ctr tlr[ .tirtg v.iti ttrrit liItr ltttrtrg ttlrit tlittlr ltodt' gti.t'r'u ttlr[i'trg tltu.t'tl6i t'til ttltt.t.t'c'titrt cittr
nt(t tt't;trg sttot tlttti lt'irtlt phcil lriAtr.

l)irrlr rrglriir uuv lrao gonr toln b0 rrlrirrrg clrat sirrlr trrlutrg tlrttc viiL tlii tlur.rc clrilr
nhl.rn rrhtr uuxin. gitrtrclcllin. c1,1sl1inin. abscisic acid, ctlr1,Ict.. llrirssirtostct'oicl,
srrlicr,lltc..]asrrrorurtc vi rrhitng htrp clrit l,.hlic birng ciiclt klai bt) y,.r,, citr vC'r'iin clrttl'Crr.
YCu cau plriii cti sr.t vin ehrrl,Cu tlir rlutrc lrrai bo vi lrlriii ttiCtn clto t'ing clic cltit tirng
tlurrng tlrrrc vit tlic ckltg tl xa nui ch[rug clutrc tong hrrp kh0ng con lrotitt totjn dirng. Vi
clu: Cytohirrirr co tlr0 clrL,.rc sirrlr ra trrrng ri vi cli chui'irr d0n ciic clr0i va liinr clrtlrn s,U'
lroii i'.iii ti tlai'. 'l lrii lr.ri. ctlr;,lcrrc cri tlrC rltLr.vc tli clruyCrr lrlry liich llticlr nlriirrg (lrl1, tliri
rrqirv rrrri rrt'r tlrrt.ic. tt)rrg lrt-,-p. l)irrlr rrglria rrrrii rrirr.eirtt trt'ri lClt stt tttrrttt lric errlr liil c:i
nhir'ng cliat rlii [riCt vrii rrhilng chit l<hirc chrtl rlrtrrc ph:it hiOn. C'ting n0n ltru y rirrrg.
ltoluronc rlotrg vit rr(ri chr,rng la nhiin-e h<1p cluit rrltrr ptotcitt cir ttr.rttt-t Itltrrrg ltrn. lliri
lli, chiit siuh tnLring (hrrc vlt co trgug ltLo;ng ph/in tLL nhr). Nri.chi currg cip lllC)t tillh
hiCu tat lroirc ntri- lrirng circh kich thich urdt rr\i su kiCn ttoug t0 biio din tl0n rn0t dirlr
[Lng nao tlti.

( )nh lr irth ( luir l)tt:u ll,i \ulr Jt rttn{ I lwt I rit

1,2.2.3. (.hit rli0u hoii sinlt lrrlil'rrg (lrr1c vit (l)lrrrrt grorvth I'cgrrlalor', I'(Jt{)
'l'huat ngiL clriit di0u hoi'r sinlr tluting thrLc virt (l)lant glorvtlr t'cgulator. P(;l{) de
( ,.1
tlrruc dirng riit nhi0u bui circ c6ng ty ndng clrrtrc cl0 chi circ chait cliCu hoi't sirth trrrdllg
t0lg hqp. t)iuh rrghia cua Vau Ovclbrcck vri c6ng tiic vi0n (19-54) vin ct)tt rlLroc dirrtg
d0n ngiy nay'.

(.'ltcit dictr ltoa ,sirtlt trur rg llnrc vtit lu ttlrilng Irop clrcit ltfi'tt ctt khcic v6'i tthfi'ng
': d lnll
L'ltLtl (lIt(ing, r\)t rrtdt harn luong nhd kic:lt tltich, itc' t!
tlte, t - ,:
ltttttt'bo:;rurg bit kj'ntit
tTtrti trinlr.sirrlt l.t', nao lrong llurc vat .

l)i tiiu ltri vi\ cli lri0u trong ctich ttirug tu'ti6ng Vi0t, thuiit ngir c'hrit cliiu hoa
sinlt trtrottg |.ltu'c t,cit clrrrrc clirng tr<-rng giiio tlinh ni'ry lrao grirn cii IrhlLng chiit tdrtg lrq;r
vii rrlriLrrg clriit sirrlr trtl(ing tlrpc vit cr'r rrgrrt)u girc trr nhiCn dtttlc sitrt sirrh tiL thlrc v{t,

1.2.2.4. Ctrit ctrit hiru clrirrr sirr!r tru'ii'ug (lrrlribitor vir tctart!arr()
ri'c ch6 vri
,l
lhLriL rrgii chit uc chl (lnltihitor') vi chit llnr chrinr sinh trrLting (relat'clatrl.)ltiQn
rurl,clrrLa drrt-c plrin biqt lil. Abscisic acicl vt\ nhrLng chiit irc elrt li.luic tlir irc clrt lrolc
L\nr chanr hly tli hoiin nhlLng clun trinh sinh l. ltoirc sinh htia, tr-ry IrlriCn, vi0c [Lng tlung
chirng di laur clraur rlutr tlinh sinh trtLtrng thi chtrir clrr<;-c irp clung hot\n totin vi'ro thuc
tiin vi nhiiu Iy do. trong do gia ca cirtg la ntot van clc. Ngey nay c<i nliieu hcrp chit
hiru ctr r6rrg hgp duclc dirng trong nong nghiep d0 lrinr chanr srt sinh tlrrung cuit thlrc
viit. Llhit li\ur chinr sinh truring cLia thtrc vit (plant grorvth retlrcltnt) L) rnQt hqp chit
hiiu cc lanr chinr str plrAn chia t0 [rr\o vir su \,Lr'(n tlai te bao tt'otrg nt0 cltoi vir nlrtr vay
nii clieu lroa chiCu cat-r cii1, urti hhong giil' ra su bicn tlr.rngr cira ll vi thiin. Ciy clut-rc xtt
ly vcri nrQt ch6t lirnr charn sinh trrrcrng cir l:i nu'rr-r xanh dcrr tli0n hinh vir stL tr'6 hoa lri
zinlr hiltirrg.. tltLc.tiCll. Srr phirt Lli0n cua uhiing ciiv niiy thi khdng hoiin toan bi rrgirrt ciirt
rrhung xu0rrg dOc hoi ddt ngCrt vii cho la rntit ilang cay cirn c6i lrtrrr.

Mit vrii vi clu ve nhii'rtg chut larr charu sinli tltr<)rrg thr,rc vilt lir:
- C1'cocel: 2-chlorocthyl tlirtrr:thyl-artrur<lltiuut chloride.
- l'irclubrrtlaz.ol: l -(4-chloroplrcul'l) -r1,4 dintcthl,l -2- ( l l l- 1 ,2.4- tliazol- l-1'l)
pcnlnn-3-ol.
- llunzi: t2 I{S. 3lts)- l-(4-clrlorophcn;'l-4-clinrclhy'l-2- 1l ,2,4-tLiazol- l-yl)
pcnten-3-ol.
- l'r'ohcxuclionc-calciunr: Calc itrrn 3 -ox ido-5 -oxo-4-prrrp ionylcyclohcxa-3-
crrccltbuxl lulc.

Nlrie rr clriil lirrrr clriirrr sinlr trr,rtrrrg tlruc lit cti Llc durrg,:horrg lui irrrlr lttLrirrg cua
gibbcrcllirr corr duoc gqi lii antigibbclcllin.

(;tio ttinh ( htit l)ku lloit:inh Inftrfig Ilnt( ldl l0

 Cltuottg 2
PLIU.ONC PI.IAI' 1'I{ICI I, TINFI L9C VA XAC DINI I
-I'I
CI IA'I' SINI I I'I{U'ONC IU'C VA't'

Chit cliiLr lrira sinh (ruung thlrc v4t li0u quau d6n hiu h0t circ cltu tritilt sdltg
cria thr,r'c vit.'lhLrc t6 cho thdy vi;^c iip tlpng ngoqri sinh cirug cit ltltitng irnh huirng
lCn tlrLrc viit r'6 nct. 'l-uy nlrien, kh0ng cle phirt hi0n cliit cli0u hot\ sitth trr,Ltltg trong
,. '.:. u lj tlo l<hic nhau trong do k! thuqrt phin tich clrung vin con
tlr$c virt vui nhie llltuc
tirp. C<i nhieu phlrorrg phup d6 triclr, thanh lgc vir clirrh luqng chit cli.iu lttta sitrh
tnrirng llrtrc r,|t trry'thco nhonr drrcvc khao s/rt.'l'rong cltr<rtrg niry sd gio'i tlrigu kltdi
cluitt nlriing plirLorrg phiill ctr ban tir chuin b! nro cl6rt dinlt tinlt vii clinh luqng chAt
di0Lr lroii sinlr tlrLting.

2.l. l)ltuu'rtg phrip ly triclr

2. l. l. I'lruo'rrg ph:ip khtr0ch l:itt

'o'.to'l

lvl6 cltLoc dirrrg clro sinh tlirc rrghie nr

I
-'z=-:--.r
6::::l)
N4art
\*.#
cirl cila nrtl drLoc dirt trOrr
dia agar chtla clrat clr6rrg oxy h(ra
I

aaa
aaa

CIrAt di0u hoir sinh truong thtrc viit

klru€ch tiin vdo trong rn6i tru'dng bao ho?r int

, !. '"
I[iltlt 2. ] l ritlt cltul ^,i
di€tt lrci sittlt It'rrirtg tlt4rc vcjt bdtrg plw'o'ttg phdp khuy€ch tdrt
( jrio I t'iuh ( lttir l)tiu Ilrrr .\mh li urntg I lrrrc I it II
 l)hutrng phirp ndy chi c6 tltc ap dLrng cho rrhfr'ng ur6 ngLry0u ct)n tuoi hoirc
,l
nlrtirtrr lllrin cul ciil'. Mri truric lri't tlu,rc lrlttnt.t, viro clurrg tlielr
;iirtt. ,,;, (., -uclltirr 30"C tli giir
- ,
l)cr ttgatt stt but Iroat ctia nhiurg chAt di0u hoii sinlr truting tl0n bC nrirt vCl. ciit,
lrtrtylittc 111'111'1;xytolttcn (llltl') \l-2rro) duo. c trt)rr d0Lr vtitt Irons irsar vii co tirc dupg
,i ,i
klri nr6 duoc nhfrrrg vao trr-rng a_ear. Iv10 duo-c clit lOn
ttltLt rttirt elxiL chon-e oxy hoir
klrOi agar vti duoc giti tlorrg buong anr tu vai phirt din vdi gitr tuy tlrr,rQc vdo loai nrri
str cltrng.

Sau ttr0t thiri gian, kh6i agar co th6 dLLcrc clung cho nhu ciiu sinh tlic nghiOur.
I)ay l:'r Irtit plru'<rttg phhp rit tOt cl6 u'crc lugng rn6i cluarr hg ve nr[rc dir cia chdt dii:u
Ittrit sinlt (rtLtirts Lrong nhD'ng plrin khic nhau cua cay. Cin chir y lirrg qui'r trinh niiy
clri clro
' tlriil
'l rrr0i (lLriut llC cuir r)llfng chat hifrr tliin brri vi rrlriLug clrlt kiclr llrich ha1,
rlc clrC ii [ri rrrirt vCt ciit c(r lhiS gay trri ngai tlCn sirrh Llic rrghi('nr tlarrg ri1-r clung. l)iiy
la rnOt plrurrng phiip don giiin va hhOng thci irp clpng cho viic tlinh tinh hay clinh
Ittortg ttr0t cliclt lttlirr lriro vi clrit diCrr lrira sirrh tlrnirrg tlru tlrroc vtii urr'rt lrrlrrrg circ
trlrti.

2.1.2. Ly trich llirng durrg rudi

2.1.2.1.Ctruin bi ruiu
ViCc tlirng goi rnO carr [lriin sau khi lhu rniu thi rit cluan trong d€ t6i thidu
lroii l<lroi rnitr va han clr0 rluqrc sp tlrat thoat clrrit clidu hoa sirrh trrLong lrong cluri
tt'ittlt triclt. C,itt ltLtr y rirng. nhD'ng hop chAt niry c(r troug thyc viit r,<ri hiirn lucrng rdt
rtlrri. tlo tlit, bit lij'nrot su thit Llroitt ntio do plidu huy thuiic v0 cnzyrnc hoirc srr
| : , I l. ,l , , A ,7
cl)tt)an '1. tltut llti gittit ciic clrit
tltir tlCu cri tlrC tllirr dirr rrhirrrg kct *1rli l;lr0rr-u clrirrlr xiic,
Cliclr lriCu qtrii rrhiit, dd giu nriu sau Iihi thu la ngay lip tuc cl6rrg rulu trrug uitrogcn
Iong. ttrip lartlr vi'r li\nr kho r'Oi trir d -80"C hoirc thip lrtrn cludi nhiLng tlii:u kien khan,
tlrirr rr;r1'si lr,.rrr clrri tiri rll ciic virn tlC lriit tl'i'ic xiiy ra.

2.1.2.2. [,y lrich

| ':
('ti nlttritt pltutltg plriill durrg rlC triclr chat cliCu lroii sinlr tlrrtirrg tlrlLc virt, cirr
cltott ;thtrotte plrihp tlrich hop c:lro cli0u kierr nglriCn cfu cua urinlr. Co nhiiu Ioai
dtrrtt tutri cir. Ihe1,;.
dutrc clung d€ tlich chdt diCu hod sinh tru6'rrg thuc vlit uhrr
nrctlrirnol hoic cthanol co thC dutrc dirrrg cli tLich lnn' nlln' Cn, Bl{, cytokiuin,
SA vii .lA; acctonc c6 thi cluoc tJirng dci tlich lnA' AtlA va GA; isopr.opanol hoic
clorolbrur ctl th0 clugc dLrrrg cle trich Illt; hOn hup lliclcski g0rrr nicllranol/
chlot'olorrn/ 90'% Ibrnric acici/ rruoc (12:5:l'.2vlt,) hoiic pcrchlolic acicl c6 th6 duuc
,. (lc
(lullll ,: tne, ll e [(rli ln ll.
,
v I

Otio I rit t ( hit l)n;t, I loi ,\nlt Ii uong l ltut I r.il

t)
 l'lran lorr durrg dich trich duirc clirng thdng thuong cho tdt ca cdc clrdt di0u
hoa sinh trrrtiug LhLrc vaL la 80% rnethanol cdng theur nr0t chdt ctr5ng oxy hod nhu
BII-l-. Mdu rluo-c nghidn trong mOt lugng'du dung rn6i lanh rdi c6 tli0 dugc pha
,. I
tltim viri luuu ndug dqi cao theo rnQt lugng ti6u chuAn dC hiQu chinh sp thit thorit
thinh linh trong clua trinh triclt vd nhfl'ng buoc ldrn sqch ti6p theo. Miu sau d6 dugc
loc vri nr6 clugc srl dprrg d6 tr'lch trti l4i d6 thu duqrc sin phArrr nhidu nh6t. Nhin
. : ,:.
chrurg. so lan (riclr tu! thr.rQc viro rnd vd nh0ng y€u tCr khric, nruc dich nhirur thu
dugc san phAnr t6i rJa. tlrOng thuong clud trinh trich c6 tlr6 lAp lai 3-4 lAn. Dich tr(ch
cluo. c nOn trQn lin vcri nhau va cd dgc bing cach cho b6c lro-i trong chlin kh6ng ltolc

lanr kh6 hoirc pha trong dr.rng cliclr d€ phin doan hoic loc xcn k0 vrii circ rluh trinh
li\rrr lilrir. Sirrr lilri loai lrtr rltttrg ttriri lt[r'rr crv tir tlielr trielr. rrr0t lttr.rrrg tlirrrg lii crr;r
:,
nlrtnq rni-rrrlr vun liin t.huirng gilp trong pha nu<rc vir Lrong sp phdtt dotltt tuang tinlt
acicl ti6p thco. Nhilng clrAt rin niy se hoir tan trong pha lt0u co, clo cl<i sE li\rn trhi6m
; ,'
nriiu voi nhirng htrp chit khirc, lldng cich li2c hoirc ly tinr d0 loai b6 circ lrtirnh vutt
rrin nriy, tlrlng lucr-ng khd cua mAu c6 thd giirn di cluqc d6n 80%. Nen chir y ring satr
:.
cluii tlinh tlich, nriiu con llt thO vr\ clrua nlrtug chiit kich thich cing rtltu cltit uc chO
vir nhfi'ng chit khdc. 'l-rong nhi0u truong hgp, dich trich clugc co th6 khong co ho4t
: ,.
tirrh trong sinh tric rrghidnr vi hoat dQng cua chAt di0u hoa sinh trrrirng c6 thO bi chc
phri lroirn todn boi nh[ng hqp chdt co li0n c1uun.

2.2. 'l'inlr dich triclr

lr2c
Buuc tltanh lgc tliu Lidrr sau quri trinh tricli nr0 thyc virt [hr,rong tiCn hurrh la
phiur doau. ,J
.
blng dung m6i. Sr,r phdn doqur bf,ng dung mdi gOn sg phdn doqtt gila
rnol pha nut-rc va urot cluug m6i hiru ctr kh6ng trQn lln (dung mOi cl4c biQt dugc
dirng se clurrc xac dinh btli clrdt cliiu hrlir sinh tluting thr,Lo vit rlang durr-c triclr, nt6
clu'gc srr c|,urg vii nhiLrrg y6u t6 khfcl. Co nhieu su lihlic triCt vi dung rnOi ch<l ttltitug
cluit trinh plrdn doqn, mQt vi clu vd chit didu hod sinh truing thgc vtlt co tiulr acid s0
dugc gitri thi0u d diiy, E6i voi nhirng chdt dieLr hoa sinh truoug co tiuh acid thi ptl
cua pha chua nuoc tu' dich trich th6 n6n dugc didu chinh d6n khoang 2,5 vir drurg
diclr ducrc cl.ro viro trong binh chi6t vOi mQt lugng tuong clrro-ng cua diethyl ether r6i
lfrc cirr th{n. l}inh chi€t sau clS dugc clti y€n vii phrit cl6 c6rc pha tirch nhau. N€u nhfr
lirung ti,ro thanlt clo ' lirc
': clLrii nranh. lrinh chiCt rtCn dirt viio rttity'tL<ip lanh clro cl0rr l<hi
lrhirrr triQt thirrrlr lrai ltlra liCng biQt hoirc clLrrg CaCl2lray Na2SOadi loai nh0 tuung.
,i
M0t plriin cua nhDng chdt di€u hoir sinlr tru<rr.rg thpc vqrt s€ di ttr pha chu'a nuoc d6n
pha chfra <liethyl cthcr. Qua trinh nay co th6 l4p lai tu 3 d6n 4 liin hoirc hon rrta d6
,,1 , l. ,:. ,,:
liy hit chAt dieu hoa sinh truirng thr,rc vflt c6 tinh acid ra kh6i pha chua nuoc. C6n
udc lucrng s6 lin thuc hien cl6 thu duoc tdi da luo.'ng ch6t di0u lioir sinh trudng. l)lta
cl.riLa nurrc sau d6 s0 dugc bo di vA pha chua dicthyl ether s€ dugc lirrn khO trong
clrdrr khdng. Mqrc dtr ptrdn doqur bing dung mOi tir cdrch thuong duo, c su' clpng tu
(;ido li i lt ( lkit t)tit llori Srtth li'mng l ln ( liit |3
trtltrc deltt nll'. d0i ltlri rrri ctlrg l.lr0ng lroi\rr lriio clro nrr',ri lnrirng lttrll, Ngt\y triry, rrt'r
-' -' : , :
ditrtg tltttrc lltrr1'thi birng plruo'ng pliiip tliclr ly It)ng-rirn, Vi0c su clurrg nrOt plra riut
nh0i viio tt'ott-c nt0t cQt nlrri va sLi drrng nri.it lau cli li\rn cho viQc chuAn bi nrlu nhanlr
horr. hiCtr suit thLr ltOi cao hurt,u'il thi tich clung Ln0i sri'clr,urg lai it hcrn. Vi6c sri dung
nh0ng cOt ultti cli-i dtLoc dung d6 trich cho liiu li0t cac ch6t diCu hoir siuh trudng vd
|. :
dit ctttts ciilt tti0l. stL tltay d6i rlirc dio dCn sp pli6n cloan bdng dr.ru_e nriri. ViQc thanh
Ioc chit diiu
hor\ sirrh tluotrg cirng dd co nhidu thi\nh c0ng Lrong viec su dpng sitrc
,i : , ,,,
liy 8iiiy. sitc liy ltip tnting, siiic l<y cQt, srlc hli khi vir sio hy ldrng cao rip. M$t phuung
,

plrhp sac l<y cti the cung cip dr.r clro clurir trinh loc d6 dinh lugng, tuy rrlriCn phtn ldn
r ..r i_ r ,.1, ,
phiii kCL lrrrp rrhiCu phrLurrg phirp cho cluii trinh loc.

2.f. Dinlr lu'r.rrrg clrit sinh tru'ii'ug thg'c v?t

'lltcu sutr tltrii lrirrlr trich vlj lr.rc ciir clrit tliirr lroa sirrlt lrrr<irrg lri viQc <linlr
,i I
lrLrirru. Clrat tliCu lroa sirrh trutin-{ tlrtLc vll. cri tlrci rltLtrc rlirrlr lrLttrrg birng sinlr tlic
rrghifrrr. rrhirrrg plrrrrrrrg plriip lroii lj lrqrc (sirc liy lilri lurirc sic li! ltirrg cuu rip), lurflr:
plrrrtrrrg plriip rrriirr diclr lrgc.

2.3, 1. Sinlr tl'irc lrghiQrrr (llionsslv)

t' t J
Sitth trdc ttghient lit ru<il ltP lltottg.sittlt ltoc clvr2'c dilttg cle tltti'nglti€nt ltottl
linlr ct)o nrct c:ltat ,u'o'i rnot dap fitrg sitth li.

I)i cri nr6t sirth tlic rrghi0nr lilru clung. ciin luu y nhtng ti€rr chrrAn sau:
l. N(r clri clic triCt cho lttrp chit clang t.hri nghiCru,
2. Ntr plriii nit rrlray cinr d6 cti th6 plrirt hiCn urt'rt ltxng uhir chit cliCu hori
sirrh trrttirrg.
3. Nir plrdi trhanh vi\ d6 tl0 thu duo-c rnOt lutlrrg lon rrr0 lhrrc vat tlong nhit.
Stt diill ittts t'tiir rrro rloi vrii rrrtlt cliiit tliCu lroii siulr trrrrirrg tlruc vlit tlirc biCt cung
pliiii nlranh vir di thrrc hiCrr.
, --. ;
4. t'lriit clitttg rltttrc siuh trirc nghiCnr hoirc rrhlLng chiit cri li0n tlulu;rlriri Iti"^rr
diCn ri nrtrc rlti tlriip lroirc li.h0ng co trong cliy.

Co rat rrlriiLr hC thOng sirrh tric rrglri0nr cho rndi nhonr chat tliCu lruri sinlr
. '...,i
[rttolt-c thttc ra[ dir bict. DiCLr ciuart tlung lii chr2n phr.Ltrng phtip sinh triic nglriCnr phtr
l11rp trii ttltuttg tiuh lrtrt-rrtg LiCng bict. Co tlri chia sirrh tric rrghirlnr tlrarrlr 5 nlrrirrr
sa u:

- (,''l). ,'' : nglttCttr clriin

' tr-tc
Stttlt
.: doiu: Sinlr tric rrghiCni dirc bie^t dirng phiit
rJ0
hiCrr rntlt clrar diiu holi sirrlr lrutirg dac thir.
,^' ,..
- (2.1. Sinh' ttiic trgltiitir dung d€ xirc dinh nhiug cluau he v0 hoat tinh vi ciu
:

trtrc: Sirrlr triic rrglriCrn tli0n ta sLr ltlrric bi'^t va tinh nhay cam tliip ring v6'i cAu tr.uc
OrLio li iult ( lrir lJu:t, lloi .\nh li uurtg I lntc I rit l,i
cua nhirrg chirt khiic nhau trong cung rnQt nh6m ch6t <lidu hoa sinh truong.
- (3). Sinh trdc nghiQm ki€rrr tra thri vd phdt hiOn: Sinh tric nghidm c6 tinh
nlriry cirur urqurh vir di4r rirrg nhanh cirrrg vdi tlnh chuy0n bi0t ddy du, cho phep c<i
., i
tlt0 su dr,rng trong viQc phirt ltiQn chiit diOu hod sinh trudng ir rr6ng dQ rat thdp trong
tung phdn doqur sic ky khhc nhau.
:,
(4). Siuh tritc ughicnr tong hoqrt tinh sinh hgc cua nhom: Sinh trdc nghidnr
du lroirc gin du rrlray ciirn di rrhting clrit cti cirr trirc cla clirrrg tlruQc rnQt rrlrr'rrrr chiit
tliCu lroii sinlr tlr.r'ttrr,u cho plrdp xiic clirrlr cluo.c lroirt tirrlr t6ug s0 ciltr rnQl. rrhrjnr dic
bigt trong dich triclr.
... .,. :
- (5). Sinh' tric nghiQnr cri yOu cdu don gidn: Nhtng sinh trirc nghiQrn cr5 y0u
ciu vi thi6t bi chu1,0n bigt vri kh6ng gian th6p, gi6 thiinh re vd cl€ thu dugc miu.

2.3. l.l. Sintr trric nghi6nr auxin

! ,.
2.3.l.l.l.Sinhtrdcn,ehiOursUcougdiQptieuyOnrn4ch (Avena): DuaLr€ukhdndng
kich thich sg sinh truong cong cia auxin vd sg vucrn diri.

2.1.1.1.2. Sinh trAc nghiQm sir th6 hiQn gene thu6c ltr (l,ticotiana): Dga trdn sU thd
. I r | ,
hi;n cua gene _,
tlr€ l<lrrinr (chimeric) trong nguydn bdo thit ld cdy thudc la duo.c
clruy6rr geire diip u'rrg voi cd auxin vd cytokinin. Sp d[nh lugng cua nrdi chit diOu
h' rii sinh trutrrrg dqa tr0n phAn ung uriru ltic dQng bdi anh sdng,

2 3.1.1.3. Sinlr llic rrglrifrrr vi s1r sirrlr lt'ttring tlrirrg tloarr rliQp tiCtr yCrr rnaclr
( lvena)'.l)ua Lrtn kha nlng kich thich su vuon ddi cira auxin. N6 khOng chuy€n bigt
'..: sirrlr Lrirc rrglri0ttt s1r
r hrt nghi0ng cia lu<'rttr.

2.3.1.1.4. Sinh trdc nghi€rn l6ng dqu cove (Phaseolus): Dua trdn kha ning klch
tlrfch su sinh truting corrg cua auxin. Nci khdng nlt4y cirn vrri nhi6t d0 vi c6 thd
tlruc hi.in trong diOu kicn sang.

2.3.1.1.5. Sinh trdc nghiQrm sU tqo rE bAt clinh tr€n cliu xanl't (vigna): Dya trdn khA
rrirng kiclr thiclr sLr tqro ri lrirt clirtlt ctia artxirt tr0tt cloitn tlriin.

trfic nghigur gibbcrcltin

2.3.1.2. Sirrh
2.3.1.2.1. Sinh tr'[c nghiQm ui s\, tq,o cluoug khu' n0i siuh troug l[a rnqch
\llordeun): DrLa tr'€rr klrir ning cua gibberellin kich thich ho4t dQng cua etrzylrte
,;
' d0 tqo dttung khu.
c-anr)'lasc

2.J,1.).'2. Sirrlr tllc rrglri0nr ciy chut chich lh lQrrg (/lalrery': Dpa tr0n khii nlng litttt
gianr lio hoti (miiu viing) cila gibbcrellin trong cdy chirt chich lti r6ng.
(inio liinlt ('lxir Dtiu lk Suth l nnglhtcl'd.t |5
 l'l'l'2'l sirrll tllic rlglritltr Lt'r,tc lra diCp lirrr dilp (l.trc'ttrcry': l)rrlr trirr ltlrri rrirp,r
kiclr
(lrich su'Lro'cliii cria gibbcrclliu lcr truc
rr'dicp rau clicp.

2.3.1.2.4. Sirrh tric nglri€rn nhirrg cay lirn rrhLL cliiu Hir L,itn (Piswn),
lia (0r1.za) vri
lthlt (Muize): Sinh trac nghienr nil,sti dr-rng nlriurg bi0u di lun
dorr genc va dua trdn
khii nirng kiclr thiclr su' vrrun ili'ri cLia eibbcrcllin.

2.3. 1.3. Sirrh tr.iic rrghi6rrr cytol<irrirr

2.3.1.3. |. Sinh triic nghiinr urd sco iOi thin tlru6c ti (Nic:otianu): Itlri klrirrrg dtroc
f

xti ly c.vtokirrin tlri rrr0 lOi thurjc lL sC khorre tao drLoc callus lroirc riit it
callus duo-c
[1o ta. -l'u1' 11;,16,, klri cytokirrin clutrc t"henr viio lhi callus sC phrit tri0rr
rlit nhanh
c[rng1 r,rii srr llia tiurg trong lutrrrg luoi.

2.3.1.3.2. Sirrir trac nghiOru srr thi hiCn gcrrc tlrrr0c lii (Niutti<ttru):
[)rrit LrCn s1r thi
lriCrt criir gcrrc tlrC lilriirrr trrrrrn rrgu1,f11 lliio tltit |til' cil)'
^ .r :,
Illu(]c lil tlrrtrc clruycrr gcnc rlii;l
[Lng vrii cli Lruxirr v:i cytol<iniu. Su. clinh lu.ong cLia rn6i
chit UiiLr Iroi\ sinh tru.uns dua
(r0rr plriin [rrri; rndu ttic clOug boi rinh
sirng.

2.3.1.3.3. Siirh triirc nghiOrn str ud rdng cua lu' di0p c0 cai (/laTrlturttts)..
Sinh traic
rrghi6nr uii;, l.1r.,., tLCn kha niurg cda c;,tol<inin kich thich s1r
nir rQng cua tir diQp cu
ciii.

1 "t | | t ,,. t i
t.-t r.J.4. )ilnrl tr.c rrglriQrn trcn srr vutrn cliri r.rr,rc hn cli"^p cliirr Itirnh (C'f,cirre):
Siph
tr;rc nglrifrrr rur,t'drr'.r tr.0n l.lrii narr-r:, cril cl,toliirrirr lifelr
llriclt srr vrttlr cliii trtrc lra tli,r^p
cliiu ni\rrlr.

2,1.1.3.5. Sirrh tric nglriQrn hiclr tlrfclr su [ao llclircl.arrin

cdn cri c.l0rr tr.iltr
(tltitttt'tilttlttrs) tr()lls t0i: Sp'shtt sirrlr sirc to ckl bclacyanin
tlrrrilig
),Cu cliu cti liulr
sirrh tlitc rlghiCrtr ttiiy dtta tr0n lilrri rrirng kich thich sLr sau
siitt-r:,' StL
si11 betacya,irr
ctilr cytokirrirr trong t6i.

2.3.1.4. Sinh tr.ric rrghiirn abscisic tcid

j
:1 ).t
r | | | .!' ... :.
'q. r. )lilrr r.l'ac ng,rllr'il1 v0 stt trc clrei rriir rrrarrr
.: ; ,,:
lr0t r'u tliclr (l,uctuc'a). Sinh tric
nghi"^rrr dLLa lrCn klrir rrirrrg ctia AIIA uc clrC su.nay
,i i . i
ntAnt cua h6t raLr diCp btii stL
t(tr()ng chc trtre ha cliClt vri su
lthirt triCn cia re Lrdt,tiut,

2'3'l'4'2' sinlr trtic nglriCttr srr rung cudng ld bdrrg (Gos.r1,,piurtt):

SirrS trac lgSi0nr
rriiy rlr'r trc'lihri rrirug cua AllA kiclr tlrich stL r.u'g ri
bO.g. Mi'c dd x', lv AIIA
, t-::;-:-:
-tttth) ttDtttl :
IhI lth.], ll,\i.\ tll 1ru,nry llnc l,u
l6
cang nhieu [hi s1r rlrng s0 ceing gia ting.

2.3.1.4.3. Sinh trirc nghienr v0 uc chd sinh trudng rna lua (Oryza): Dga trOn stt uc
s1r
,i
ch0 sinh trudng bc lri lun criir A[]A, Su giinr chidu ddi bg la lua li€n quan cl0n sq'gia
tlng rrCrng dd n tln cliroc xir ly.

2.3.1.4.4. Sirrlr triic rrghifur sqr cldrrg khi khAu cdy thdi ldi(Conunelina): S6 luqrng vd
nrirc dQ d(rug cr'ra klri khiu drrcrc cluan srit cl€ utic luqng ltiim luqng cua ABA hign
0r9n tf ()ng nrau.

2.3. 1.5. Sirrh trirc IrghiQrn cthylenc

: ,.- ,,
2.1.1.-5.1. Sinlr trirc rrghiQnr diip ung b0 ba dU'a tr0n tinlt sirrlt tru<rng ngang, su'
plrorrg l0rr vd srg uo ch6 thdn cklLr FIn Lan (Pistort): Sinh tric nghiQur rriry dlra tr€tt
l<hir nirng cria ethylcrrc irc ch6 sU vuon ctdi, uc cl,€ sq tnd tnoc truc tlttrcrng diQp vd
kich thich-sp sinlr trut'rrrg ngang. MQt trong ba tinh chdt ndy ddu c6 thd dtrgc dung
dC utic ltrtrng lriirrt ltttrng ctlrl'lcrtc ttrtttg tttatt.

2.3.1.5.2. Sinh tlic giy

ra sr,r sinlr truong nghidng cua thdn va 16 cit chua
nghir3rr
(l )'copersicory': S1t sinh truong nghieng o.ddy ld sr,r gAp xuong cua cu6ng la. Dql
,i,
llgllrong cuil cLr0ug lh ti lQ voi lrirm lucr-ng cl.hylenc cltua trong tttlu.

2.3.1.5.3. Sirrlr triic rrghi€nr kich thich sr,r cltin cua trdi (cir chua, chudi, chanh...):
-l-lrd'i
Ngrrl,Cn l1i cua sish tllc nghiOrn nay lir dya tr0n s1t chin cua tLdi. giatt cltitr cira
cric loai trhi nhu cii chua. chudi,, clianh... ti lQ voi hiinr lLrgng ethylerte c<'r trong tttdu.

2.3.1.5.4. Sinh trlc nghiOur s$ rirng tn'diQp b6ng (Gosslpirrl: Sinh trdc nghiQm
nay giutrg lrhr.r sinlr tlic nghiQm cua ABA kich thich slr rung o b6ng. Ethylene c0ng
kich thich srr fuug. Ivlirc dQ rung giy ra do ethylenc ti lQ v<ri h.rong ethylene c6 trong
nrIu.

2.3.1.6. Sinh 1r'iic rrgh iQnr brassi uostcroid'

2.3.1.6.1, Sinh trirc nglriQnr l6ng tlru nhdt <liru cove (Phaseolzl'y': I(hi auxirt dtLoc 6p
clung v0 mOt phia cria l6ng thi l<lng cong di va theo sau ld urQt chu ki' chdm. BII lirrn
giirur chu k! chrlm khi clrrqc ap dgng rnQt gid trutrs hhi xi'ly auxin. I(lti Illt duqc xu'
lli rrlri0rr lrrrrr. IAA giy rlr stt sitth tt'tnittg cottg sC ttltaltlr lttrtt.

2.3.1.6.2. Sirrh trirc nghiQur sg sinh tru(rng nghi€ng cira phi€n ki lia: Sinh tras
nghi0trr nay dpa tren l<hh nirng cira BIt kich thich sp nghi€ng cua phien la lua. DQ
' '.: la tLra ti lC vo'i hdm luong clra BR c6 trong rttiu.
rughicrrg cua phiCn
(ircio liinh ('luir !)in Iloi Suh lilrtug T hrc I cil t7
 2.3.1.6.3, Sinh lrirc nglriOru v€ sr,L frc ch0:0 nlrtrng n6ng ctQ t6i hio, l]l{ giiy ra sLr
vu<,r diii vii corrg.'r-uy'rricu, ri.rri
dQ tiurg cao, llrt irc crri sinh trudng
tct r'0' Mr"rc d0 giir'r v.rr. 'tlng vii ldur
dr'ri ,r*ii giri rri d6i clru.'g ti lc vrii hiirn
lur.rng lllt c6
trrlug nriu.

2.3,2.lloi Iy tric nghi{rrr

l lli plrrrrrrre plriip lroi.r
ly tr.ac nglricrrr ilrrr.rc dirng plro biCrr ngiiy rriry clC xiic
dirrlt clrit tliCu lroa sinlr lrrL<irrg llruc vlil kj sic
ky long cao {p (lll,l.C: Itiglr
pcr'lirrrnarrcc litlritl clrrornatographl,) va
sic k.v khf cQt rnao clunu ((iC: Gas
clrronratoglirphl). Ngiiy nay lllrl.C chi cluqc
cltrrrg vrii rrhiing clriu tlo (cletcctor)
cd
d0 uhay, cao hoirc vtii nh0.rrg mIu loc rit sach.

2,J.2.1' l'hiit rri6u crriit.sirrr tru'ti,rrg trru.c v{t bir'g sric

t<j, ktr6i ptrii
Sic k1 kh6i ph6 (GC-MS: Gas crrrr-rnratographv
- Mass sr)ectronrctry) duuc
xcttt lt) llltucrltg plrtip t6t nlrlt li
Phlirr tich chit diiu lrtxr sin6 tr,r,t,rg vi .o r1r' giii.
vii chirrh xtic' PhLrcrtrg Plrirp niiy tlii clugc cltng cld pharr
rich IAA vri rrhii.rg 6gp c6dt
cti lic. rluirrr' AllA vir circ chit bii'duir.g cr,ia,(r,
gibbcrcili., cl,tokirrirr,
ll.as5i11e51st.id Vii .iastntlrral.c. Klrdrrg gi6ng
nlru <tdu clo clrrng ctro
lll'l'c' datr dir cho (ic'tltutrng Pltri lruv rniu vir kh6ng ctd diing kh0i p6u,o'g p5iill
pSuc lqii cSit
clrui. dL'oc cliurh criiu ph6.g xar. Su prrar tri6' .uu pt,r,i,.,,g
*i,i, prr6 crro prrcrp
d, clrttrc cltit dirrrg vi cltt'c durrg phir bi6n clc *a. rini ,i,u,,
'i ,ii,,i, lrxrng tit cir circ
rrlrtirrr cltit tlictr llrx'r sirrlt trtrrrrrg tril ctlrl,lcrrc
vi clrit rrly "n
cli.li,,g,t,,r.rc,riic tlirrlr
trang ( iL'.

2.3.2.2. l)irrlr lupng ctlrylene

lr'tlrylcrrc lii rrttll chit lihi rriu cyuir trinli lriclr
n6 ciing dtrn giiin, kh6ng plrrii
cltta cic tlurrll r'6i tricll va thauh loc
nrcrt cich phri'c tap nhu.circ nh6nr khirc. ViQc
rrlr6t khi ctlr'lcnc Lrotlg birth ltoic 6ng nghienr
kin cing it bi nhi6'r tirp clrrit
tt'iclt ctic clriI klrric. Voi stic ki li,hi vi.r cliu '5u
cld ion L,oa,,'L,un Ii'. trll) (flarrrc
iotlizatiorr tlctector) cli cli'rrrg clirrh ltrtrng du<1c
cthylerre vr'ri rn6t lup.rg r.iit,lrti tl0n
natro rrtolc.

2'-l'2.3.1'hrit rrif. ctrit oieu rro:i sirrrr tru.ti.rrg trru.c vit b:i.g ilr)LC
Mic du IU'}r.c krreing prrai Ia prruong phap t6r nrrit
dd plrrit hien vii dirrh
luo'ng chit cliiu hoii sinrr truong, vin
c(r nhfng;lhuorrg phip vrii criu dd dic
bict
dttt.rc tlirrrg' Vicc Plrlit lri0n lrtryrrlr rluirrrg
rlii drrtrc rltrrrg.rc,li,,r', lu.lrrrg tr.rrc tiill lAA,
tt'rii lai AllA vi.jasnrorric acicl phtii clutrc
chuyen hc,d tlrrinlr hl,razone huyrrli clLrarrg
vi\ b|assirtosteroid tlriinli hisboru;ual.c lruj'nlr
c;Lrang trutic lthil;hirt hicn rriug pturo,,l
ffi, r,u,clTiii lj;,.\*t, tj;",\cTfrJi 18
do huj'rrh quang. l'huong phap niy ldt nhay va <liic trung. tuy rrhi6u y€u cAu nrlu
phai dLrgc lqrc thit sach tlu<ic lihi phAu tich. Siic ky lorr-e co thd t<6r hgp v(ri khOiphd
thainh sti<; ky long lth6i phd (t-.C-MS; l,icluirJ chrouratography
ri --r ,:
- nrass spcctrornetry)
dc xac drrtll nlring liin llop kh0rtg diirr xuiit cria gibbcrellin vii IAA. Mac tlu c6
^ :,

nhiOr.r tru viQt nhung dQ nhay cua l,c-MS vin khdng bing (ic-MS, chinlr vi viry
GC-MS ngiiy cdng du'trc srjl dlrng nhidu.

2.3.2.4. Siuh lrfic nghiQrn rrri6n d jch h<1c

.'. . :
Sinh tlitc ughi"^m nriOn dich hgc lri6n rdt ph6 bi€n vr'r co nhiCu giii tri tlong
kltoa lroc tltttc vit. l'huirng phiip nay y€u cau sg' tinh loc uriu t6i thiiu, rlo do viec
i;
phlitt tich so tttatt lort tlri rit nltanlt vA khdng dirt ti€u, tuy nhi0u gii tlriurh khong tlrrlt
s\r' qtran Lrong. 'l'rong rrhiCu truo'ng ho-p, I IPLC vlrr cin thit5t dC thu dLrtlc nrfiu s4ch
. ,: . , .,,t ,. ,:
va Rct (ltlit tltu duo.c cti tltO dltng d€ so siinh voi cac phuong lthirp sinlt tric nglri€rn
hoir l1i. Nhrin,{r tro ngai trong sinh trdc nghidnr rniEn dlclr lroc lir cr-r thi cllu d6n sg
uuc ltl(.tt)g vttgt rluri': ve nrftc clQ khdng nguy6n vri nhfi'ng vtn d0 klrirc. l)o d6, vi0c
clton nr6t nrQt k6 ltoqclt lirnr si.rch uriu rdt .An thi6t d0 ldm cho sinh tric nghiQm phi
hgp voi k! thuit plrin tich hlrdc nhu GC-MS.

2.3.3. Xic d!nlr cu6i cirng

D0 xic dinh rach loi vii nrdt chir Aie u hor\ siuh trurj'ng thgc viit dil dlro-s sinh
.! :
tt'itc nghiCttt cati cri ttroI ureil xdc dinh cu0i cirng v0 htlp chAt nay biutg phu'ong phdp
sric lty. l(lri dLrrrg ttlt|ttt-e;rhr-rtrng plrap sirrh tric rrglrii'nt l.lri vii.e xiic rlinlr cu0i cung
I _,.;
liltt)ng carr tlii0t vi rrlrt-Lng ho-p chdt gi6ng nhu chat di0u hoir sinh tluong dd dr-Loc do
ct'.ii cirng uiiy tlrrrirrrg ilrrtrc tlrtrc ltiCrr vrii sic Ly lih0i ph0 lray
lLlirrr-t. SLt'xtic tlinh
,1. :
" sitc k1'ltlti
'kdt 'hqp vtii hoirc sitc k! lottg. Phuong plrrip nr\y cho m6t kCt luin chlc
chr'rn v0 chit dtruc hhrio sirt.

2..1. Kdt lurlu

C6 nlriCLr butic clO ly triclr, lt\nr saclr va sinh tric nghiOnr chit dieu lroa sinh
tlrf(urg tlrrrc virt. (.'iie lrrrric rlir cti tltC tlrrr.lc ttirrr tirt rrlrrr tr.orrg lritrlr 2.2. Nlrrtrrg ciiclr
d6 x{c clirrh clrAt diCLr hori sinh trurrrrg cirng riit cla clqurg, tui theo urpc tlich cr.ra nha
ltglriCrr c[tu ntii llltutrng plriip nao duuc ip cluug. l)u clro cri dilng phuung plrall phAn
ticlt ttito di chirrrg rr[iit tlti viCc clrrriirr [ri rrr0 tlriclt hr11l tnLric li,lri llhiin ticlr ki rit rpran
.i
Lt'ortg di .
trich duo-c ltoiut chinh, de liur sach uriu uri khirng bi tro ngai bdi
11'
phrrong phap dang clung, d6 trich va lanr sach nrAu dugc t6i da vd d6 cung cip clu
, i , ,,1
Itrr.tttg tuiitt dat clrat ltrrrttg ilrt cho viCc xlic dinlr cu0i cirrrg l<lri sirrh tliic rrghiCrrr.
lliCn tai co nhicLr k! thuAt phdn tich lriQn dai duqrc dirng trong nghi€n cuu chAt di6u
hoir sirrh tru(rug tlu"Lc vat lray phirr (ich lroa ht2c n6i chung. Cac thi6t bi cin thii5t d6
thttc lti0n citc trLttic pltiiu ticlr lrtliur chinlr lii sic [y ldng cao iip, sic ky ltlri, sic ky
khi - khOi phd vrl cQng hu'ong rir hach (drn (Nuclear nragnctic resonauce: NMR).
Circ thi6t bj nriy cho plrip rhav diri cldu do ho{ic phuong phhp phdn tich <td nhd
rtghien c[r'u clat clu'qrc nrpc dich cu thi.

lr40 thrrc viit

I)uug rn6i lv tliclr

I

+
Phin doan vii ldnr sqch biug cdch tihch ptra nuoc
r,<ii pha drrng nr6i hirr.r co hoirc sirc ky c

I
I'hiin do4n vA li\nr saclr bang
J 1,,
sirc kj lorrg cao tip (l Il)l,C)

-l-6ch
thirrh nhicu ohan doan

Sinh tlirc nghifrn (bioassay I'r'ic nghiCnr l1i htia cl6 rlin
:iic lllriin tltran thu dutrc d0 x inh vi dinh luo-ng: GC
inlr hoat tinh sinLr hoc ;C-MS. LC-MS. NMIT

llitth 2.2. So'cli t1, trich t,it xtic'dinh chat cliitr hc,td sinh trtrd'ng tlnrc vtit

'

lh Snth liturrg l huc I it 20

 cAU r nuc r.roA Froc, ,,*,, .,?l,'Ji,'u. uo nNr-t l ruoNc stNrt r_V
CUN CAC NI_IOM CI IAT NINU I IOA SINI.I ]'RU'ONC'lHqtC
vn'r.

'l)'ottg chuung nay

chu ydu ctO ciip d6n cdu truc h6a liqrc, sinh tOrrg 61rp vti
lrnlr hrrong sinh ly ctia tiinr nhorn clrdt cli€u hoa sinh trr.rirng
thyc v{t c1(qtr li auxin,
gibberellin, cytokirrin, abscisic acicl, ethylene, brassinostcroicl.
salysilate vir
.iaslttolta(c. Cltic chiit lhLrQc Itltonr polyanrine vi ciic chAt khiic s0 clutlc tlc cip d cdc
chu'ong sau tu! thuOc vio vai trd sintr ry cria'6. Nrri,
chung du,o.'g corg criip ung
lieu luung cho tdt ca nlrDng ch6t clicu hoa sinl.r rru<rng rhu.c viit
cra uiet.oit.rrg t,int
chuCrrrg (hirrh 3.1;. Chtit di6u hoa sinh rruong c6 tdc clung kic6
t6ic6 se t6d hGn tdc
dong liich thich tr'6ng <16 thAp vd rd'g dan d6n i'h rruo'g
t6i cta. t<tri n6'g d0 vuo-t
clua urtrc kich tlrich t6i da se giy arrh liu<ing fc clrC.

.(!
rto
(O.

.3
= do chdt dt6u hoa sinh ii
llirrlt 3' | . l)rritry4 cortg cli[rt rt) rrtti'r' rrq)trdlt i'rtt tritjtr rtirtrt
tlii v6'i rtirtg clo c'hit cliiu hdct sittlt tt.u.u.nH

3. l. Auxirr
Auxi'lii thurlt ngii chung clqi tji€n cho lop cua nhfr.rg h<r1r clrit rlrrrrc dirc

lllll ll'll "'l' do,,s a r

!r
-,;,!;ffio--i*fii0,,*
u,i

iinh lrucrrrg khac ben canh stt vuon dirri, nlrung str vuou dtii cluoc
xe'r li) rhe. ch6t
{rt' Auxin ntii cltung rttattg tfrtlt acicl v<ii rnQl. nlriin kh0ug [riio lrr)1 l,ui,. ulin[.Ii,,
-___=__i=>-'-
xuiiL cua chring.

3.1.1. Sinh t0ng hg'p auxiu

Sau l<hi phirt hiQn la lAA, nhidu lro-p chdt goc inclolc ciing rti du,crc plrir
l,if',. I!J,,l ie,, ho,.,rrf,
rrlr tha.h
n6 th.\"I. I I r\T '-: -r-.
Noi , :
chLr'g ngudi ta chdp nhir' ring ilI1;;G;ig-t,";r,
i!ll_ct"_
, ---.
.lAA.
a^ ,a -----':-
-,,.^i...^I-
!l!,"ph,"r, nr6t arrrino acid, tlong lrat pliin vti nh[.ng,r.,,, ,inl, tr.'D,g h,.,4t .r,r"[ffi.
2l
 lll0 Phatl sinh cltdi, lth0i str l<.ho'i cua ld, lii non riarrg
lttn, hQt dang phar tridn vir trr.ri
rlrrr eorr tluirrrI klrti iutiirr, lrlrrr cirr.lror), vii o.ry lrol.l.

l^ll ll
cu.-'5.n-cooll
\- \ft-' Nll'
ll
lrr,ptophan
' ( ().

'l;)11'illil"

'l ll
rr ptarrr inc
I rrdolc-3-pvruvic acid
l12 o1
Cf-l cu.-c-NOSo:
I v-'\N-' $
A nr isc I II [tr"or"
ox idase
I
(i lucobrassic irr
( ll. (itt,Ot f lntlolc l)\,ru vate
14-'',t
1..-
,1 I
NIl| ,\ dccarboxr'lasc
.,l._N-.ll

II
I rrrlo l-J -ct ha:r o I
/ --cot Myrosinase

'()
tr'\
t.it -ir-Cl l,-c.'\
ItrLlt
'lc cllrutrol -\-^Nt ll lt
i)\i(lilse
tl I rrdolc_3_ircetonitri le
I ndo l-3-accta ldclrl.dc

t/l o./-\ / ,2-tt',O

-\ /l,,,1'
N itratase
lrrrl<rlc acct ir lrlclr t,tlc'\
dclrvrllrrgcrrasl' \.
,' NllJ

l^1- tt
or. coorr

tl
Irrdol-3 -acctic acid

l'lirrlt i.2. So di .sinh tort,q lto.1t crtuitt

('ri
ttl'.rl lrt: llrorrq ciic cttz.r'tttc tlriurr girr r,iio t;rrii tr.itrlr sirrlr
t0rtg ltr.r' atrxitt
ttltrL ciic c-!A]']:.-1!-llltlrlllrr tlc'clt'[rox,r'lasc.
tlyPl.rrPlrirrr trirrr.si!!]r_LL!-.lic: iunillc (]xi(lilsc,
irrtl.lc p)r'._r,irtc dcr:arbox)4asc,
.l' E"l"
acc tltldclrrrclc r l),h.vdroger rase ui tralase. 'll'rrnc,
virir ttitralase. nrr:i rrlJ -,,---r-*^"-,i.]^ll
li.ongqLrirtrirrItt0rlgltcrpauxitttl0uc6sg
tharrigia cLia l<Cur. Cl6 hai;r]dr,.,')t€Jhfirli tl0 hiCrr,loi
t,.ipr,,1,t,on thdnlr IAA tuf
tltco loii tlrttc Yiil (llirrlr 3.1). ('rrrr tluong tlru nhlit
bicn dOi trl,ptopla' tlirnh
irrdulc-'l-PyruVic acitl b0i cnz1,'111s tr-vPtopharr transanrirrasc.
li0 c1j clu,o-c clccarboxyl
lrtiir btii itttlolt: 1.1;'tLrvitlc dccar'[ro.r;'lal;c thiirrh indolc-i-acctaldeSyilc
(lAu\l<.1), 16i
22
clLrqc biOrr d6i tlranh tAA clua con dudng indolc acctaldchydc clchyclrogcnasc. Con
dutttt-e tlru hai IiCrt cluatt tlcin su dccarboxyl htia ctia trylltophan [lrtinh trypLartrin brii
trypto;lltan dccarboxyltsc. k€ do duqc bi6n clOi thanlr lz\Ald btii anrinc oxidasc vir
cuOi cung thiurh IAA boi IAAId clcltydr<lgenase. 'frong nrdt viri loi'ri tlruc viit, IAA c6
tlre clLrqrc t6ng hclp til rthitng tien chAt khdc khdng phiii lii tryptoplran. () dua leo,
irtdolc-3-cthituol clii duqrc tirrt thdy vir klri xu ly ngo.,qi sinh co thi ilin t(ri s1r Lao ra
lAAfct do cnz)'nle indolc ctlranole oxidasc. NhiAu lor\i thuQc hg Cruciferae I'to/ac
Brassicacede ch[ra glr,rcotrrassicin ducrc bi6n cl6i thinlr irrclolc-3-acct()nitrilc bdi
cll:/-)lllc tttyrclsitlltsc va cutri eurrg thanlr Inn boi cnz),lrc nitrulirse. IVlqic cltr co rrlriCu
hC thirrrg l<ltiic rtltittt t0rr tai trurtg ciy. trry rrlri0n phiirr l<irr glri ulriin drrt.rc lii str t0rrg
lrop lAz\ tLr tiCrr clrat lii tryptophan.

'l'rurtg
tint t6ng hgp cia ciic auxin la d circ urd phAn sinh. lA non, nriinr lr oa,
, , , .i
h0L darr-u lllriit lliirr. rrrtit luo. ng riit it auxin chLtlc cli chuyCn clCu circ crr cpran. Su vitt
clruyin triry tltco rrliu nr6, tutrng Ling vii ctl tinh cuch phiin crrc vtii vlin tilc 5 - l5
,,1.,_., : .
nrur/ h. ltat it thay auxiu vdn chuydn theo chi6u tu duui l€u vii thco chieu ngang.

3,1.2. Citc auxin plrd bi6n

'l'u ciic nglriCn cuu cuu Darwirr (1S80) ui lriC,r tu(.tng qunng hrLung dirrrg clin
.ciic lihrirrr phii ctrir Wcnt (1926) vi rrrQl clril cti lrort tirrlr sirrh lroc trrrrg rliCp tiCrr y6n
ruirch.zlvcrra dir dln dCn su phhL hien ra IAA trortg uuiic ti0u cita ngtLt)i tlo c0ug ctia
K0gl va ilzragcn-Srnit (193 l). IAA sau clti cld cluoc plran lap tir nrcu llia, tlorrg rrltiCu
loii thuc vat bic th6p va l.lrrrc vat bqrc cao khirc. Nhu viiy IAA dir rlutrc xcrn nhu k)
luxin tlrLtrc plriit lriirr suni nhit. Ngi\1'na1'. LrCu cirnh IAA ntri sirrlr edn co nlriCu
irrrxirr tlrtr.rc tirrrg lrup rrvi rrlriCrr ttrrtc diclr klrirc rtlrrru. Atrrirr tt)rrg lttrll lir nlrlrrrg lrtrp
chit cri hoal. tinh trl(Ing trr nlitL lAA, nhung kh6ng hodn toiin LtLong trr r'0 cirr Lrirc.
Ar.rxins co LhC clutrc clria thiurh 6 nlion sau (hinh 3.3):

- Nlrirng din xu6t irrdolc: ludolc-3-acctic (lu\A) r,a lnclolc-3-Lrutylic acid

(1Bn ).
- Nhirng [rcnz.oic acid: 2,3,6-trichlorobcnz-oic acicl va 2-nrethoxy-3-(r-
d ich lorobcnz.oic ac icl ( Dicarnba).
- Nhri'ng chloropherroxyacctic acicl: 2.,4,5-trichlorophenoxl,acctic acid
(2,4,5-'1') va 2,4-dichlorophcnoxyacctic acid (2,4-D).
-
' Picolinic acid: 4-anrino-3,5,6-trichloropiconic acid (-l ordon hay
l)ichloraur ).

- Nhilng naphthalcnc acid: u va p-naphthalcncacctic acid (u vti B-NAA).

- Nhirrrg naphthoxyacctic acid: u vir p-naphthoxyacctic acicl (u vr\ p-NOA).

2i
 ('()ol
f^l - -fl ('rt, ( (x)tl
I
t'.,1-,- ct
(l1".,7'^-,.-OC|1.1

\A,.r
II

il
ludol-3 -acet ic ac i.l lrrdole-3-trvtlric acid 2,J,6-tlich lolobenzo ic acid 2- rret lror1,-3,6-d iclr loro-
(t/\.,\) (lllr\) bc rrz-oic ac id
(d icarrr ba)

Din -tuit ittdula Benzaic acid

()cll.('o()il (rcllrcooll
\_l
__.cr()otl
.,i- -.., .zl'....(ll
itl lil
tl LI
Clrt\. Cl )-.' Nll2
CI
2.4,5-tf ichlofophcrroxy 4 -anr i rro-3, 5.(r -tl i c h I o ro-
acctic ac id 2,4-dich loroplrcrroxy p ico lin ic aciil
(2.4.s-'l') accti'.- ;rc icl ('l'ordon or p iclr Iorail)
(2.4-r))
(' lt I rt ntlt lt e n ( LtJ'.rce t ic uci (l I'icolirric ucitl

('t tr('ootl (l)Ct t,c(x)

I

e li CD hoxyacetic ac itl ll-Naphl hoxyaccl ic acid

tr,-Naplrtlralcrrc aeetic acid [-Naplrtlt;rlcrre acctic acitl cr- l'.lapht
(u-NAA) (lj-NAA)
Nultltthnlcne ucirl ltl rr1t I t I I t o.r.1, nc e t i c ac i d

,/-,ro.-,,tCl
tlll
t^) *"
coor.r
Qu i trc lo luc

Itiulr 3.1. Ctitr trtic ctiLt tlt6l so uuxitr n,iti itiiirt

Cirlorol;lretrt)xlacrclic acid' picolinic acicl vi) nallhtfralellc acid ld nhi.ng

tlrrrrlc 1r'ir cri lhLrirtrlt rlitrrg lr'orrg rrirrrgl rrclriilr lriCrr rrir,v. lAA. rrrrplrllrirlurcircctic lrcitl
(NAn). irrdolc---J-[rutl'ric ircirl (ll]A) durrc dunq clro srr 1rf(.]ng rd vei str dqru trai. Su
tltay tll6 rtlrirrrg rtltirnt l<ltiic ttltau l"r'Cn nlqcll \,dng hoirc rrraclt rrhiirrlr ch c(l nlrii'ng inh
lttnirrg Irtattlt lCn lroal tirrlr cri:.r irrr:;.in tlrri v..ii trlrtirrr l.rhcrroxy acid, l)o tirrlt cholr loc
ltllifllg pltcttcrxf irectic acicl dilc biet nhu 2.zl-i) va 2,4,5-'l'dir duoc ding
ctra clr(rrrir.
r'6t pli6 biCn cl0 lt\nt tlrrrt)c trir cri lii lring trong. lrlriCrr nihrrr clua n.tirc dir hi0n rlay dA
lratt clrC vi l.hirrrg stI dung. Nlri'r'rr.q clriit ni\l,cirrrgi di cluoc rlirrrg rit nhicrr Lrorrg chien
tlaltlt ltita hoc. Qr-rinclorlc lir tlrudc uri chon loc do cdng ty llASl; cua Drjlc sin xu6t
vir clrtttc dirng lairl LhLroc trir cti lurrg vuc trcn rudns l[a. Nhirug ho-p ch6t
pltertoxyitcclic acitl tlrrlurrg liit LrCn.'l'hi'ir ra. plrin l6n cti lii rCrng hai 16 rndnr kh6ng
llri phan lrr,ry nhirrrrl lroP clriit ni1,r,i v:i;,tlrtLuirrl dur,rc ciirng vdi nilng tlO thAp. Chiit
cl6c nriitr cla cattt r.lit drttrc rlirne tlC liirr, rLrng lir ciiy Lrorrg chien trirnlr Vi€r Narn ld
lin t li ittlt ( h,,t l)tt ll,ttt Surlt !r.t,t,ry Ilurc l,it l,l
lurQthdu lrqrp cua 2'4-l) tu do vii n-trutyl cstcr cLra 2.4.5-'l'. ll'()ng (lr.rri (rirrlr t0rrg lrqrp
cLra 2,4,5-'l' vti nhiiug plrcnol gOc chlor clir sinh ra trlri0u siirr plrAur phu khitc nhu
nhlng chlolodioxin rAt nguy hi6nr cho nguoi di6n hinlr ld
2,3,8.'9-tctrachloltrditrcnzo-para-diox in (hinh 3,4).

llitth 3.1. ('tiu !t!o cth clioxin (2,3,s,9- !etrucltlot'odibenzo-prt.u-dioxin)

Attxitl Iti"^rt tli0rt trottg circ t0 brio tlrrrc vit clu(ri nhi0u hirrh thLrc lihlic nhau:
auxitt ttt do, tiCn attxirt vi.i auxin litrt ker. Chc phuong phrip ly trich th0rr-q thuirng se
l.ltu dtruc auxilr t0rrg sO 1ca ba krai trdn). Circ hinh tlrrLc trich auxin thco kiiu l<lru6ch
tiitt tlttttl'ttg tlttt dttr.rc auxiu trr do vii auxin liCn liCt l.lruirrrg k6t clrit vtii ;rrotcirr,

3.1.3. Nhff'ng:inh hu'ii'ng sinlr lf

ALrxin li0n cltran cl€n nhi0u rlLrii trinh sinh ly trong cAy. Vaii inh lrudng cluau
ttr.rttg cria auxirt clic\u ht)a cdc clrrti Lrinlr sinh ly ctla tlruc viit c6 tlrC lii rl0rr plrtr sau:

l).
VLrtrlr dai t6 bao: C6c bLroc tinh huong lOn su vtLon dai do auxin tirc
(
dong co l.h€ t6nt trlt nhr"r sau:
- Ciiirrr tinh clrOng clriu cua vrich tC brio d6i vo-i s[rc cirng. Di€Lr nr\y g6y ra
do stL bt giiy rrlriirrg liCrt kCt khOng cOrrg lr<ia tri gi0a xyloglucarrs vri ccllulgsc
Lrong vich ti buo.
- Cti rnirt stL thiry ttdl vi rrltltng chc clo uuric vcri td biio. Ngay ca klri rip suat
.t; ., ( ..1 , .:,
I'llillll Ulill'l lf()tl8i tc buo
'. lilr0rrg tlray d0i. tlrC nirrrg nutic Llong ti [riro tlrrrrc xri ly
vrii auxiu lrti nCtr 1'Cu lron do su giirrn thC nirng iip suit.
- Su giirrn th0 nang nttoc clto pl.rcp nrro'c di cliuy€n vao ben trong te bdo vd
tao ra mOt ap suit vi phia vach t6 bao cti tinh clio lao ra su vuon cliii.
- Khi su vtLcvn <iiii hoan tit, nhlrng li€n kt5t khdng cCrng htia tli giira cellulose
vi\ trlrifng ltolvsacclraridc trii lap Lrri lai. Quii trirrlr ni'ry hltong thuarr rrglticlr.

(2). Qtrlrrru lrtttirrg dtittu: (]tratrg lrrLtirrg dQrrg li'r sr,r pluit tliCrr criir rrr0t rrrir
.:.
LltLtc val lttttrn-u ver llltia siug d<l su diip [rng vui thOng luorrg tlrrc titill Iroirc gr.adicrrt.
1'hco giii thuy6t choloclny-went thi iinh sing chi6u tu' m6t phia s0 gdy ra su di
rl:'
chuy0n cua auxin v€ phia tdi, do d6 nong d6 auxin r,0 phia tOi cao htrn phia dugc
chi6tr sang, Str pliirr plrdi auxin t<h6ng ddu duoc xcrl li\ trguydn nlrin giiy r.a su
nglriCrrg.
( ireio liiuh (hdt Dii, I lui St,,l,in,r,g Uu.rc t an ?5
 (3). Dia hutrng dqing: Lir sp hurittg dQrrg ctin nrQL cu cluarr tltltc vqtt dlip utlg
,,(
vrii tltrrrg lr,rc. r..,wCu urQt cliy cltrr;c tllit ui-rrn ngirng, clriri cua nti sC rrghiCrrg lelt pltla
tr'0u ngrrtrc chiiu voi trilng ltrc (rlia huong tlQug irnr), trui lili ri sC nghi0ug xu6ng
eur trorrg luc (dia huting d0ng duorrg). 'l'hco lhtrl'Ct Clroloclny-Wcnt v0
, , 'i
theo cltiCLr
riia huong clOng thi thiin vri 16 clirp ung vrri lrr)ng h,Lc ticlr li1, IAA vC lthia Lhip hon.
'l'rotrg thiin IAA lticlt t.hich s1r sinh truOng tren irt-rl. dtiy cua tltarl va larn cho thliu
: ,,
lr!,ltiCng vC plrirr trt'rr. l(lri cril clrrip ri tli tlri hlra rrirrg dii;r filrg criit r0 <liri vrii trr,rtrg
lr,rc bi rrrrit tli vii klri tlrrt clrrip ri'rrt'r lrri tlri tirrh di;.r Irtrtng tlirttg clrtr.rc plrgc lr0i.

,' :
(4). LJu tlri chdi rrgr.ln: Ch0i ngon cltLrz-c biCt la ngul'On rrliiin kh0rrg clr0 sg
pliirt triin cLia clr0i bCn. l(lri cit chbi nllon, ch0i bCn se phat tli6n.'l'rry nlrien iliecr
. , :, , ^ plriit triCn trt;i lCn va fc cliO srr plriit triCrr cua rtlrfitrg chOi bCn
thtri giun. elriii bin
luoc sau. SiLu klii auxin cluoc phirt hiCn, hant lurrng cao cua no trong dirrh ch0i cirrrg
cli rlurrc ghi rrhin. ULr th6 clroi ngort dir dttr-rc ghi nhirn u nhieu lot)i thr,tc vilt vit <lugc
,.: ,..i , {. ,.1
rlii'u khiCn lr0i clriit diCrr lroa sirrlr tt'tLtruq.

(5). S1r ttrqrng r'6: .luliLrs Vrn Saclrs (l8tt0) cho ring trong Iit Iron vi\ trltiLttg
, t..
elrOi lroitt clQrrg cir chfa rn6t chiit tliitr hr,ra sinh truirng cd khir uitug ddtt truy0n va
l<ich rhicli srr trK)ng 16.tnn cling tli clrrqc bic{t lir clrit cti khir nirng hich thich s1t
.;
trLoug r0 crra carrlr giarn vii cirng rlh ctro thriy lthii rrhng itng cllrrrg trong tlrttc ti0rr.
l.'lliirng auxin t6ng lrcrp LlirLcrng drLo-c dirng thny,vi IAA ttr nlrieri vi chLrnp. khOrrg bi
plrirr lrril lrrii crrzyrrrc IAA orirlirsc lr;r.1, 1111ii 111, cnzynlc l<lriic vi:;f lirtr l;,ri ltottil tttir
tr()n-q lni)t tlriri giirn dai. Ap dlrrrg atrxirr rrgoiri sirrlr cti t.lri llrelr llrielr sr,r'turTrtg ld vit
:irL plriit tricrr sttnr cira r'6. triii l4ri s\r'vu(rr) ck\i cira ri rtrii chuug bi rrc cl16 trir khi tilr
dtrng voi ttong i.l0 clu rrho. Sr,r iLc chC sinlr tlutrng crra auxin tlruong co liCtr tluatt cl0tt
.l
srr l(iclt tlriclt' sirrlt' t0n,u lttr-p ctlrl lcrre .

((r). Srr siirt sirrli cthl'lcnc: Str l<ich thich sarr siuh e-tlt)'lcnc gay ra do ar.txitt
':. tirln trerr ci\ chua boi Ziurrnclttratt vii Wilcoxttn (1935). Ngt\y nay.
tlrroc ghi nliin dlir
auxin clir clLLtrc [riit la chit ilieLr hixr sinh trttirrrg kich tlrich su'sirrli t0rrg, lrqp cthl'lcnc
tlCn r{riiu loai thLrc viit nltr.r dqu xanh, l[ra, cti long vuc...

(7). Sy phirt trien tltii: Sg gia ting ltich tlrrruc trhi chir yiu clo su lro rQrtg cua
,i,,
tC biro _qiiv ra. ALrrin cir lien quan rlen srr llit r0ng cira tc biro va clting vai trt) c<r ban
tlong vi"^c rluyCt tlinh sy phirt tliOu cua tri'ri. Vai trt) rirauh rn0 c[ia aLrxiu troug suphft
rriirr ctilr lnii llirrn lrrri yCu t0. llrir rrhit li nloi (luiur lrf girrir srt ;llriit tli0rr lr0t vrii
.t : I

liiclr tlrrrtie crriri cirrrg vir lrirrlr rllrrg tliii, llrrr' lrlri lir vigc frp tlrrrrg irtrxirr lCrr tliii rriro
dti ri nlriLrrg giai doarr dac thu cua srL phrit tri0n se giiy ra srr tlirp irrrg. Vi dq 0 diu tiy,
ntii phoi nhi vi'r plroi trong bd qua san xLrAt auxin, 116 cti chLry6n ra ngoiii vri kich
iirrb lrinlt( hit l)ttlrt llo<iSurlr'ltrtotrylinrc Itit 76
thich sp sirrh tru'drrrg. Vi tri cria b6 qui trirr trdi c6 nrQt anh hudng l6n d6n hinh d4ng
rrni. l]6 c1ur.i tliiu tiiy ninr tr0n rrgoi\i d6 hoa thit qui vii <16 dirrg thc clQng. Khi trich t6t
cu tli qul thi tlai l.hOrrg plrrit tridn.'tuy nhiCrr, u0u tticlr tit cri bc{ qua vii rill dlrng
auxiu lCn dd hoa tlri tlai plrirt tlidn binh thucng.

(8). 'l'r'inh cpri sinh: l.i\ su plrriL tricn cua truri kh6ng clua cluir trinh thq tinlr.
i,
Arrh hutirrg cuir uuxin ldn tlinh clua sinli thi khdng ndi bat bdug hnlt huring c[ra
gihberclIirr lCrt tlrrrr tlirrlr rur;'.

(9). SU rpng: Ndu cirt Lrd phi6n 16 norr thi la sE d6 rtrng. 'l'try nlii0n cudng 16
se kh0ng n,,rg dtro-c xu ly auxin nhrr lAA. Sq lpng lir ld clo su thiutlr ldp tirrg rtri
'rcu
vir hi6n tucrrrg ni\y bi chi ph5i btri auxin. Sr,r rpng sE gia tirng khi luong auxin ndi
bidn birrg hoirc lon hcrn auxirr ngoqri bi6n. Xri ly ar.rxirr vd phia lh cua tartg rlrnglt\m
giirnr s1r liro lroa. vi phia thAn cia tdng ruug kicli tlrich su lio hoa vt\ gAy ra str rtlllg.
Su giiinr auxin lCri sinlt trong Ia hoitc cac cil quan khac cr"ra cay sC giy fa sU l'lrng.
Vi0c xrr li Nnn lray 2,4 D cirng li\nr giAnr sr,L lr"rng trAi.

(10). SqL tlri ly auxin co thi l;irn thay dOi giui tinlr
hiQn gio'i tinh: Viec xu
cria hoa tlCn nrQr s6 toi i ciiy vir s1r thay ct6i giOi tinh nr\y chLgc ghi nhin c6 li6n cluart
clCu sp kich thich sinh tdrrg ho-p cthylcnc. Khi xrr lj aurin trgoli sirtlt tll'i lirrl tilrrg s0
'li'utrltg Dai
lLLirug hea cii trCn lio biu bi. Circ rrglri0n c[ru cr-ra Khoa N0ng NghiQp,
lloc ('iirr 'l'lrtr cirnt glti rrlrarr drrrrc lirrrg arrxirr rtgoai sirrh larn {arrg s0 lttt.rltg ltoa tltrc
lren ch0rn chCrnr.

3,1.4. Sr.r'ltltiitr hriy :tuxirt

Su hi0n cli6n hay c6 mirt cua auxin c6 mQt iurh lru<rng sAu sac [0n sg sirrh
.l
truting vir pliiit triin cua thrrc vAt. Cac dqrng li6n I<0t thu4n nghich vii bAt thuqrn
nglriclr cr'ra irrrxirr. cnz)rlrrc oxy h<ia va quaug oxy ltoir la nhurrg 1'.iu t0 plrin lrrry htlac
liirn bit lroat auxin. lr.rrz-ynrc IAA oxidasc cld dugc tirn thAy tr0n nlti0u loeii thuc va-t
co lthr'r nilng ;rltlin lru.t' lAA. Siin pham chinh tao thtinh trong chu tlirrh <Jccarboxyl
Irtia oxy lrria lir 3-rnctlrylcnc-oxinclolc. Ngot\i ra c<)n cci circ chit khiic trhtr
3-lii,dr,r*",'.,.,lir'' oxinclolc. indolc-3-rncthanoI, indole-3-aldehydc vtr irtclolc-3-
carboxylic acicl. 'l'r'ong clru trinh oxy lriia dccarboxl' hoa klrong hoan toirn ct6i vOi
IAA tlri tlru plrint chirrh ll oxitttlolc-3-acctic acid. Nhiitr loiii ciy cti
rlrrcrc san
cnzynrc IAA oxiclasc co th6 phfin hiry dugc lAA. Circ hgp chit pltcttoxy acctic acid
,i,L
tlruirng rirt bCn vl khirrrg bi cnzyurc IAA oxidasc phiin huy. l)lrirr ltin cfly llt rQng
sorrq tit rliCp lilt0rrg tlrc plrrrn lrtr1, 2,4-l) vir 2,2i,5- l'. l'lrin ltin 2.4-l) vir 2,4,5- l'dtLtlc
clung thOng thuolg lA nhirrg acid tu rlo. nruoi vd mu6i anrine. CAn chu y rang nhlLng
acicl tu tlo rit 116 bay lrtri vii phiiu tiur.
(irtio li inh ( hdt l )pu l lo,lt .\utlt l) tntttg I htrc l dt 21
 .].2. Gibberellin (GA)

cll3

Itirrlr 3.5. Cciu tric cti. ent-gihberellane ld sttin co.bdn ci, cac zibbereilin

Gibbcrcllirr lir ruOt nlr6rn chdI cli0u [rr\a sinlr truo.ng thu,c vl,rt c_ti suou
crrt-8ibbcr9f!4!l!-lhinlr 3..5) vii kich thich su plran chia tC tlao hiiirc sr,r vrro'n dAi tC
-llitp. Ngoii ra clt[rrtg cdn eir nhfrug chuc ning Ci€u lroa khac tucrng tu nhu
gibbcrcllic rrcid (OA3).
J Acctr,l CoA
fc-J-t,,t.tut" I
+
Mcyllorr ic ncirl rsopc'tcrryr
.t ./ ^m
I\lcvalorr ic ucid 111, rollhosph n tc Isopcn tcrrl'l Al\'l l)
.:Y*_-)
lsulreut.*l t pyroph.syrlratc ---+ ---+ Farncsl,r pvrri'rrrs'6atc
->
l)irnctlrr'l:t
v l11, l gr1'rophosphatc
\ t'
0
(icran1'l p.r'roplrosphatc V io laxa rr t h irr
+ \
liarncs]'l py ro;rhosplrate 9-cis Violaxarrthin
.t
(i era rrl lgc ra rrv l pt,rophosphn tc
Xarrthoxin
..t €6- Clr{t Liru ch{rn \
()oPr,l gtt' roplursprhulc sin lr truii'rrg gdc uniurrr y'.llA lklchytlc
+ \
crr 1-l(:r rr lcrrc
Allscisic ircid
+
crrt-li:rulcrrol I
Chit t,irrr chirn silrh trurirrg
t,, <--)6--- | gic p1 r'ir,iitliirc r,:l triaz-ole
r,rrl-l(:rur, rnrl |
'l ctcycl:rsis
J "----)i-l lrrlbclrlidc
crr t-l(tr rr rcrroic ucirl ot-l
I

7u-lr1'dnrry li.l u rcnoic acid

I
v
'!
I

+
Gr\12 akhlrvde

Gibbcrc

IIiltlt i.6 Sittlt tirrg hop gibhcrallirr, (.t:tokiniti vit Ab.st.i,sic. uc'itl ti. t)tevqlotric o(.id
:: .- -:::--:--::.-
(Jtutt lr t( It,it ltt,t llthlSt lt lannr.Tluc Iit
28
 GA: la gibbeLellin du'oc thucrng rnai rr6a dAu ti6n vd dd cluo.c dulg c6o nr6t
h0 thirrg sirrlr t'irc rrghii.r clruiirr. erirlr-vi vrtrr 116 dMi dien
r. (iibbcrcllirr drrtlc phd, ltlr diulirjrr llAltl !,r.
Giblrcrelltt Jiriikut'oi. 'l'ir lihi rlutrc Plriit lti0n, no tlir rluo-c tint tlriy tt.Ctr uhi0u toill
thy'c vit rrlnr tlrtrc vit lr0t tirr, ttlfCvet htjt tr.in, d,rougiiG-reu, t.,o luc, n6ur vir vi
-{:mmn.

Sinlr liittp
3.2.1. Sinh hr;'p rrihhprr,llirr
tdng hrrn gibbercllin
i ta chip nhin rin
in drrcvc tdng hcrp tu nrcvalonic
acid tlorrs nhine ch6i no sinh tru6'ng tich cuc va trQt dang phlit triOn. Chu
lrittlt ltrevalonic acid l<hdng chi ct'r lien quan <1€n siuh t6ng hop gibberellin rna cdn
li0rr clr.ran cl0rt sinlr t6ng hqrp c),tokinirr, abscisic acid vii brassinostcrgicl (5iplr 3.6).
Sau klti tnevalottic acid bien ctOi thanh nrevalonic aciil pyrophosphate rdi rhanh
isopenLcnlil p1'trrllhospltate se tirch ra thco hutlng tritrg lrclp cytoliinin, a[scisic acicl
vi\ cott cluirttg klttic tltco ciic buuc ti6p theo tle tao tlrtirrh cnt-liaurcnc sC clil cliu su
tlrirrlr lill ctic plriirr tLr gibbcrellirr. Qrrii trinlr tOng ho-p gibbclcllin cs tlri bi irc clr0
btii c:ic clrit lirrrr clriltr sirrlr trtrdrrg tnrrrg [rtrri'c clruyCrr lrtiir l"(r gcrarrylgcr.auyl
pyloPhosphatc thiirrh copyl pyrophospha(c. Circ chdt k)rn chAur sirrlr Lr-Lrdng g6c
pyrirtridittc, tt'ittzolc, tctcyclacis vir inabcnlldc ciiug rrc ch6 str triin rtOi ttr
ctrt-kaut'cnc tlrtjnlt cttt-l<aureltol, tu cnt-kaurenol tlrinh enL-kaurenal, til enl.-kaurenal
thanlr ent-kaurcnoic acid.

Qui trirrh sinlr gibbercllin cia nim G fijihtoiva thrLc

tOrrg hop vit biic cao
co thC clria thernh 3
-eiai doan chinh:

- (--hur,6n hiia rnevalorric acid thinh ent-kaurene.

- Chuycrr Irtia cnr,-liaurcne tlri'rnh gitlbercllin pr'ototypc, GA12-arcrchyde.
- clrLrl'6rr hoa (iA12-alclcliyclc tlranh C2e-, r6i rhiuh cre-GA vdi con cludng
l<liirn-q l3-lrl,clroxy'l h6a vii con duorrg l3-lr1,d;ey1,1 hoa srrrn c;- ci:rc vie,tr.i khirc nhau
vd sau cLurg thtrnlr circ dqng GA khiic nhau..

Str cltttvCtr lrtit tir isopcnLcrtyl diphosphatc (ltl,roltltosphatc) tlCn ciic clang
t ,.i
grlrtrclcllin clrrtrc tlif n la chi ti0t nhrr tronq hinh 3.7.

(ittio Li inlt ('hLir l)ih llni .\rnh liwtr+ I lttrt I tit 29

 - l,s
ANI
(lll20l,l) GAI
lsol)cnlcn\ | (liltltospl)ltc _.-> cllrollP GA2
-> -_-->
( iclanr lgcrarrr I dilrhosphatc
Copall'I cliphospharc

(Y' =
<- < GAJ ( cA3
zt:
cnr- 7u- l 11,

I
I

I
I
I

Y ll -Co,ll " c()2H

(;i1? (;4.],1 -( ittlh('lrtc

/\,/r\ t t
/\t\
"'\iffL
[qft) ,,,rtffI;
2.3-drrJchydro (iAe ( iAt,l

t i
. I
tl c()2lr
--G45-*- c,ln ---(,n4
; "rf"
iiit r rr..(c,2t.

rffi-qx'
I
(,

lf^'r,_j1t-_fi;'',,nr "l"ill,j"'" "n,

;.,n
l!-lllltt"l'q I 1-h1'tlnrxl I lrtirr srirr

II
I'\
': ;':
-(j'lt ] cor ('( )21 |

cA rr

(;,\ ,r-ca t:llx, I i t0 (i429 (iA3

IIitth 3.7. So'tli .rinh tong lttrlt Gibbe rallin

;:-----------
Itr ,, !ritth I tnir l)itr Lt.i .\uh tr,;,,;rt nt!, ti
l0
 t'A|(l l'l t;rr111,y
r# \'q{
(iA-r (l:.P)
a-/;-A .,\ ,,^F-{/-\ \.'"'
ln
.
r): Qlllh,*
|..,,,
\ r/rrr
(;A4
lt \
I ll L()tll
(l;.1')
,n,

/: ,,,\

'q#44_trfr"ry
(iA5 (t')
()

t .L.^d-{-\
t,l.-zt',A
(iA 7 ( t:. P)
cjo I

CAe
*f YJSa""
(l:.P) GA 1s (r)
-.n,

ti,lI\.F
)-i1-1
r
'--
co,
(iA;1(1,) ... c( ),t i

(iAtr(tr) (;n t,t (tr)

{ rtl
to.,r[-\-olr
a^ 'r-/\ .'\_ !tro
,,,4is> \l-l!,/\,,'-
:I I lr(
(iAtr,(|.l'l ,:, t,i, t t trl I I

((. )rl
( ( )rtI
ll ( (|,ll
(ir\r7 (l') I
(iA re ( I'l (iAlo ( t')

c( trl I

((),1| ('ort I -,1i, tto,"

( l') (iAzj ( l'j; oA){ (|r.t')
otl

Jl{ :'q{Ps z'fr

!()
()

('( )tl I
cortl iil
c())tl coltl
( l') (i4.7 ( t)) (P) CA2e ( l') (;Ato ( l"t)
.{
rll .,f
)t i )l I
-.(

q4* CM
r iA3l (t') (jAJ2 (1, )
uA;3
( (,,11

(l))
( ()rll
^i',lj
!ll()/-\
rt-lr .\
J.-l.lti\/=
: ii I rr
ct
t:. r,
'(
ir "l
I

I
iA-.;1, (lr) i.rr2t t
'.r_re rr , (iA40 (r)

,Gtfrlt =ffid)il"dtr,#
i,,,i, tt""
(;At r (ti)
,,,
:
co.rl
ll

r;,r.,!-tflj
Y-i;i \f\,,;,,,
CA,lt (F.l!) (iA,15 (l')

- (F-): Chi circ gibbcrcllin c<i nguOn gOc

tu ndrn
- (P): Clri ciic gibbe rellin c,5 ngudn g6c
tu cdy xanli
Ilirilt 3.8u. Cdc Gibberellin ri,GA1 d€n GA6

.tl
 |tr/,.

(. ()t | |
o
(= 0,11
(i Ar, (l') (;A.D (l:) (in (P)
il co2rl
r8 Gn 50 (P)
Ito
olI
()-g )

i- co,l I
(iA1r ( l') (;451 (P)

1)
/.+
g()

\ lH, Lr:.X L:\*X )*.^+X \

IPffi(#TfuI,$*
:#
:,',{' .,t'){-,.,,, <;r"' (# -
un'11,,, 'u'Jttn", ,''i'uu'

aR
I col y)!X'f (::/\

.ffi
f\:..'1'
Corll ' rr COrll Irilli ti-o,rt f i'i tcu,,, r 'Cr)rll
(iA(,(, (P) jnt (iA67 (l) (t,)
(iA(,8 (P) CAo; (p) .,{rtr (;n 70 (l')

(;A?r (l)) (ln?2 (l') (i17i (p) colll (iA:l (l') (;A?s (P)
'r .rrrI
'1rl
oll (J /-\.,(rlt

fficr#q#
()ll 9llt,
V,'tl en,,
9''tr o.ll(l
/* '::r.ifr
'"l LUl,, .r Lvzrr t ll c()?ll I ll co2ll

o
(iA;(, (1,) cA77 (P)
CA77 (P) Cin 78 (p) CjA?e (t,) Cn E0 (P) .oll
/---\ )l | .r /-----'-\ llO^ ,-
''cd#,r^4,rp
(

i, l, db r'd# -i

r) ,-'1 (, a-\ t'rr - Y"o

fj*,
,,.qt-'mK-ftfr
(ir\s6 (P) GAET (P) CABR (P) CAse (l) (iAeo (P)

- (l;): Ctri circ gibbercllin c6 ngu6n g6c tu uAnr

- (P): Chicirc gibberellin cd nguOn g6c ti ciiy xanlr
Ilinh 3.tih. Cac Gibberelliu ti'GA4, cten GAs,

(irict lii ('luir l)iu Ilui Suh li utttrg lhui t qii 32

 COrll
cAe.r ( I')
oH
o
.\\,
-cH,
gc)

cL)2l I

Gn e7 (P) GAes (P)

.*tr (;n t{)r ( l') &-,,|

clto

(iAlo!
L1/2r I

(1,,)
cortl
(;n ror (1,)
() U pn-
7,',
ct:-?
( iA
lor,
c( )'l
(r,)
()ll
I
Or
YX,,,,( ir\ 1,,7 (l)) (iA ros (
Corll
t')
V*
''t*-<
(;A t0,, ( I')

o- c o- c ,
,\\> ,
,\\,
C()| (o I ti
=A<
lt'' I = -{-.
co?ll
CAl1r (P) cArr+ (P) oA 5 (P)

9ll o o
/ -'r ?R
so
co:ti tl
corl I c0,t I
cA (, (P) cA, ,u (P)
(;A
tro (1,)

,rt( )ll ()-------: ,,\Oll ,\ .r\Oll

i /'-1.\()rl
4/rK ,\ ^F, 'r\Oll r\Ol I

+:l-*nz
i ''
co

lto2c
iH
CO

lt
\,,,,, coJl I
(;n rrr
cort I

( i.,\
1';
(iA 1r2
(in t2.l

\\

II
I to,c cortl tl
OrC co2tl
(;A cAr:s U /\ (iA llo
r27 t29

- (F): C'hi cic gibberellin c6 nguein gOc tir nAnr

- (P): Chi cac gibberellin cir ngu6n g6c tn.cdy xanh
Itittlt 3.Bc. (lcic Cibberetlin ti.GAsr den GA 1j6

i.l
 3.2.2. Nh*rrg rinh huong sinh lf rria gibbereilin
cibberellin.c6li€n quan d6n nhidu qua rrinh sinh lli trong cdv. Tuy nhi€n o
nhirng chi. loai voi nhfng y6u 16 khdc nhau s0 quy€t dinh gibberellin dac hi€u
hi6u
qui nhrirt. Gibberellin dnh hunng rJ€n nhi€u qua trinh sinh tn-rdng vii ph6t ridn
cua
thuc vdt nhu su phrit tri€n rhdn, su niy mdm cua h6t, midn rrang, tr6 hoa- phan
h6a
gitii tinh, irinh quii sinh, ddu triii va lfro hoa.

- .4nh hriing
trAn sg phrit tri€n ciltt lltqtc vfir s6ng: Citc gibberellin da bi6t
I
dcu co kha nirne kich thich su vucrn dii cua rhan hay su phdn chia 16 bao. Su kich
thich vuon dii
crla GA rhti hi0n rdt 16 rr€n nhtng cdy non hoqc bQ phdn non, d cdy
dii trudng thanh hay co quan dd gin thi 6nh huong si kem di. Nhin chung. GA kich
thich su sinh truong cua nhi6u loai cdy d4c bi€t la nhirng ciy lun. Khdc voi auxin,
rinh huung vtron dai cr]a cA ldn thuc vdt s6ng thi 16 hon tr6n chc doan rniu dugc
c{r[. I hrrc vat dap ung voi ciic loai gibbercllin khilc nhau cing kh6c nhau. Do1 voi
truc ha dicp rau di6p. nnh huung kich rhich su vuon dai cua cA6 khdng rd net, inh
huun-r kich thich su vu(m diri crla GA.t, GAr va GAr nranh ddn va GAe tgi c6 iinh
hudng manh hon cd. Trong m6t vai trudng hqp thi rinh huong kich thich srr v\ron
clii truc ha diip drra leo cta gibbercltin cirn_q kem hicu qua.

ltn tinh trgng lilt,: Co nhieLr bi0n cli thi6u sinh tong hop GA
- rl'nh hu'rtng
cli clLrt.tc phrit hi0n. Dnl ta lirrh trang dun gcnc. kiclr thLrdc cua cirv bi6n di co the chi
,: . .:.
bang rrror llhan niim ciy binh thuonq va srr lun chu ycrr ki do long bi ngin lai. cic
dang dirt bien lun nhu d6t bi6n bip lun (Zea nra-r,s L.) tll tit ct5 vi\ lua lun (ory.:c
sutivu L.) 'lan-ginbozu vii whiro-c. cA n6i sinh ki6rn soat hoat d6ng cia bip vd l,ua
li'r C;\1. Viic xu li CA n-qoai sinh liirn cho cac criy nriy cao rrcy lai binh rhugng.
cirrrs cir rihirng tlang dot bi0n lun khting dip ung vdi vi0c ip clung gibberellin ngoai
sinh vi\ cri-r vlin l[rn sau khi xri lf .

-.'lnh hrdLrg ftn sy'nri.v n,6,, crru h6r t,ti ntitn trarrg; Iri0n nay cA duoc
LriCt lir nlrtng chit co ktra nirng kich tlriclr nri.r nriur r.,i phd vr-r rnicn tran-e trdn nhitu
loai ciiv trong. (ii\ cir the kich thich hoat rlong cia c.ic enziymc tlril,phdn ildrolase
trtlrrq htlt ngi cOc. Cr\ ngoai sinh tirc ctting ldn ltip alcrrrone ctia hdt ngfi cot ri
kich
thich str sirn sinll snz\'rnc q-anrr,lasc dLl {iic dong lcn srr phin htn,tinh hot thanh
drrirrrs dorr,'l'iic tlong rriiy cti tilc dunq kich rlrich nii-t,nrarn vii phii rir rnicn trlng.
Kht'.ri lil-r,trtltcrtl] rttiirnsrirnkhi lili rtii (ir\r.(i.\cirrrccirthikichlhichsrrnur
rrtittt cLtit Irrt lttu tli,-ip rnrr khdne clin xu Ir lirrlr siing clti. i;l\.,i,,g co t6ci ttrav rnj
clieLr kie rr nhi0t do thip hoirc ngiv tliii di plirl vd rni0n trrng,

- ,4tth htr&ng l0n srt tr(t hou: Cibbcrcllin cti klui nirng tlrirc cliiv qua rrinh tr6

il
 Itoa rrong nhi0u loai rltuc r rt. I)oi voi nlrlna civ can r'8u cirr ngtiy dii lral tliii qrra
diiu ki91n lanh truoc tro hoa rhi klri xu ly GA trong nhirne <liiu ki6n kh6rrg ciiur irng
chfrns. ss- Lu'trttg lttta ra trtt Itol. Anli lrrrtirrg rrir cri licn quarr d.1rr su kicli thich tlrrir
trinh phin chia t6 bao va irroLr diti tci bao.

:.,
-.1ttlt l|rt sy pltiin hda gii'i titrh, tritth quo sitth, dQu trdi vd liio
ltu'ii'rrg
hrio: GA co the larn thl1,dOi giiri tinlr cua hoa luone, lu nhu auxin. c1'1e(inin vri
cthllcnc. Tu.v- nhidn. GAcil hieu qua rrguo.c voi auxin vii cthylenc. GA lt\rn tang so
hoa duc trdn dua lco. Cz\ cirng gii1, n€n hi€n tuc.r'ng trinh qud sinh vir tqo nOn trdi
khong hot. GA cirng girrp cho trai to va tri hoin su lio hda. Cac loai trai nho khong
h$t o Nhit. Uc. M1r vii chiiu Au tlrrrdng c6 xu lf GA3. Bing c6ch giiim lao hoa. GA
gif cho vo trdi canr quit trrsi l6tr hcrn, ch6m nrdm khi chin va keo dai tho'i gian btio
quan hon. GA cr)ng liirr cho vo tiio dep hon, cay kidng tr6 hoa so-nr vii tirp trurrg.
GAr ctne co th0 ': giup qui trinh san xudt rnalt trong cdng nghiQp san xulit bia hiQLr
qud hon va ngdn hon 2-3 nga1,.

3.3. Cvtokinin
Cytokinin la nhfn chit adenin duoc thav th€. n6 kich thich s ,,Illan
chia,t€ ban va nhfrng chuc nins. di€u hoa sinh tru&ng khiic gi6ng nhu kinetin
(6-furfurylaminopurinc).

ti€n dugc phin lep t& DNA tinh tring c6 trich dut/c thanh
-C;'tokinin ddu
trune vd ducrc p.oi lir kinctin boi vi no c6 khd nane kich thich su phiin chia ti bdo
hav su ohdn bdo (cvtokincnsis) trons rn6 l6i thudc la. Cvtokinin c6 neuon c.0c tu
nhi6n duoc phdn l{p diu ti€n tu ld zeatin
hQt bap non vd duo. c CAi
-(6-(4-hydroxy-3-nrethyl-trans-2-butenyl-arnino)purine). Ngiy nay, hiu h0t
e)

H2 cHa
NH-C-C=.C-CH?OH
^ ^'H
N/\r-r\
\N-
,. N
- H

Hinh 3.9. Cdu mic cia Zeatin

Hi6n nay, co nhi€u ci'tokinin tOng hoc rluoc bi6t. C6 3 chdi thong rlrrrro l'l
kinetin (6-furfur,v-laminopurine), tsA
(6-benzylaminopurine), va BPA
(6-benzy Iamino)-9-(2-tetrahydropyranyl)-9H-purine) (hinh 3. I 0).

Cvtokirrin dd duoc tirn thAv s hau h€t fruc vAt b6c cao. rOu. ndrn ki sinh vd
---
Gpo Trinh Chh Dteu I loit Snl: Tnmng ThLrc I dt 35
 \hdn,e k-
sinh, vi khudn, va ctng co trong phin lon rRNA cia vi sinh !at va re bdo
d0!g rlg! l-ii€n tai co hon 200 cyrokinin ru nhi0n va tong hop dd duoc phdt hi€n.

N4N
...,, y, _
o, Nr'li,N,r_NH
N
NH n G-' *N--N
6-( hr rfury lam inolpurine Fcnzv larn inop urine

--n N'. N
-.-N '''1. 'N-'i -"1
tu

__ | ;-ltl ' \._-/j

(6-benzylam ino)..9-(2-tetrahydropyranyl)-9H-Purine
_
Hinh 3. I0. Cdu m.ic cia 6-(fitrfurytamino)purine (Kinetin),
benzyluminopurine vdBPA (6-benzylanino)-9-(2-tetrahydropyranyl)-9H-purine)

3.3.1. Sinh t6ng ho'p cytokinin

Cvtokinin co nhi6u nhit mi0n phdn sinh va vung phiit tricn cri hieu
quri lidn tuc bao 96rl r0. lil non. trai dang phrit tri6n vd h6t. Ching dugc .rem la
dugc t6ng hqp o r€ va vAn ch ddn ch6i bui vi i€u btio c6o cho thdv rins
-FoFlnin au,r. lim thdy n nhuacLTE, nhi8n. cy'tokinin dii ducrc tim thiy nh'idu
thiy ring chting c6 thd dugc tdng hqp o do. Su sinh
Lrong rnr.r ,;tia triii va hot cho
t6ng hop cytokinin li€n quan d6n cic buoc khoi diu cua chu trinh mcvalod!_Aei
'I0n is-1_[ppgrtgrrl phosphatc (hinh 3.6). Buuc ri6p theo. isopentenvl se k

-
AMP d6 tao thanh isopentenyl AMP. Chrit ndy sau tl6 duqs bidn d6i thanh
---
isopentenl'l adenosine theo sau bdi nrQt loqrt phin ung khric de tao thirnh cl,tokinin.

-- Nh0'ng rinh huri'ng sinh

3.3.2. lj ctia cvtokinin
l.l
Cri nhi0u cytokinin duoc tirn thiy trong ciy, tuy nhien. chi, lodi va nhlrn_e
yeu 16 kkic si quy6t dinh c1'tokinin ndo l:i hieu qud nhit. Sau day li nhfn_s dnh
huung sinlr I! cria cytokinin:

- Phfin chis l0 hdo vit tgo thdnh co quan: Vai tro chinh cira cytokinin trong
.it-r li kich thiclt str phin chir tc bio. Crllus co thc duoc t1o thl\nh ban diiu chicdrr
rturitr ltt.ric l'tokinirt riCng k-. 'lin'nhi0n tl,-i tlul,tri su phat triin cira clllus. su kOt
hop ctra aririn rii cltokinin tt) ra cin thiit. -l-i l"s auxin/ r:l'tokinirr sc arrh hrrong luln
.. :
str tlo crrllus. rcl hirv clrOi. khi nang tiri sinh ci1'tu callus li nrdt cong cu k! thuit
sinh hoc tlrdng thudr:rg dnng d6 chon lo,c nlrlrne. cal'khdrrg roi diiu kifn khd han.
stlcss do nrirn. binh. thu6c cd hai' nhirng.r'ciu to khirc.

(*'-ln,;;irtl;in hi.s, ), Ii"-,Jt*, n l6

 - Sri'rtity uri,rn, rr., nto'rQrtg cfta ft bdo rti co'quutr:Kinetirr co thd giup htit
'.: :
rau dicp nav nrirn vuo-t qua inh hudng uc ch€ cua 6nh sdng do xa. C1 tokinin ducrc
phin chia rd bro. ruv nhicn virr co rrhttno rrJon-s ho-p thriv
bi0t la chiit kich thich s1r

dugc inh huong cia cytokinin l€n su md rQng cua r€ bao. Cytokinin kicir thich su
rno rong t€ bao truc ha dicp duoc cit tir cdv ci ctii. bi ro. c6y lanh va nlrielu cir soug
'.,..
lu dicp khic. Str rtto r0rrg ttlir tc Lrito Li do su hip tliu nuric qlir,ra do srr gniur tlti
nang tharri thiu cua tC btlo duoc kich thich boi su Uien doi tro lai cua lipid du rrt
tron_s truc ha di0p thanh dudne kht (glucose vei fructose).

- Sg tu'1trtg r0^"yd
plttit tri6n rd; Cytokinin c6 th6 kfch thich hoic uc ch€
st!
su kh,oi diu r,a phit triin cua r€ ru1, thco nOng dO vA thbi gian xi ly. Kinctin co thc
kich rhich su gia t[ng trgng lugng khd va su vuon ddi crla 16 cdy ddu lupin con, trdi
.i:^
lai hai 1'eu to trcn bi uc che o n6ng d6 kinetin cao. Khi kinctin duoc xri ly l€n rd o-
n6ng dd th6p, n6 kich thich quang ho. p va sinh truring. Tuy nhi€n n6u rd ti6p xric voi
0,47 FM kinetin hon hai ngdy thi sinh trtrdng cfra 16 vd loin cdy s€ bi gi6rn rd r€r.

- St1 phit tri1n n4t vd chAi: C1'tokinin co kha nang kich thich choi b6n va
dac biet ld vuot qua anh hucng uu th0 ch6i nggn. Bang cdng nghq di truy0n nguoi
ta dd lim gia tang dutrc su sinh tirng hop cytokinin trong cay thuoc ld va
Arabidopsis. Mot gene cua vi khu6n giii nrd enzyme isopentenyl AMP synthase.
rnol cnzvrnc dap ung vrii quii trinh san sinh cytokinin, cung voi kich thfch gdy soc
nong di dua duoc genc nay vao cal, thu6c Ll vit Arabidopsrs. Gene rnoi niy dugc
, I -0-.
hoat hoa bAne cdch clro ciy churCn qcnc vilo di0u ki6n nhi€t do 40-45"C rrong nrot
thoigian ngan vii k€t qua ki ldm gia ting ham luErg zealin riboside monophosphate.
zeatin riboside va zeatin lan lusr ld 23,46 vA 80 lan. M0t vi dU khrc ld vi6c dua nr6r
gene giai rnd enzyrne bi0n dOi IAA tu do thdnh mot arlino acid li6n hc.rp bat hoat.
Duoi dnh huong cua genc ndy, IAA tu do bi giAm, do dd kich thich ch6i b0n phdr
.; .; , t.
tri€n. Uu th€ chdi ngon duoc di€u khi6n bdi su can bdng gita nruc do cytokinin va
auxin n6i sinh. Co hai gia thiet v0 rnoi quan hd cua cytokinin l€n uu ih6 chdi ngon.
Cia thi6r thri nh6t cho ring c1'tokinin c6 thO uc ch6 enzyme IAA oxidase trong ch6i
bdn vd do d6 cho phep tfch lfry auxin va gAy ra su vuon ddi crla choi ben. Gia thi6t
thu hai li
cytokinin co th6 khoi dAu co ctrd l;en quan d6n suc chria o ch6i b6n bing
cdch kich thich su vdn chuy€n dinh dudng. r,itarnin, khoang vd nhirng chdLt sinh
trusng khac di tac clong len su sinh truong.

-
Tri hodn s4' Ido hds vir kich thich sqr vfrn chuytn chhl di,rh drdng vi
nhirng hqp chfu hA'u ca; Cytokinin c6 th6 gi[p ldrn giim quri trinh lao hoa khi rach
Li ra khoi thdn cAy vd hoat dong nhu ld ch6r thay th6 cho sy cin thi6t cria 16 dC gidIrr
lao hoa- Cl,tokinin cilng c6 khd nang thay the inh huong cua iinh sang va lam giirn
Oiio Trinh{tuit Diiu HoA Sinh Trudng ThEc l'{jt 37
 sLr k-io tri nguydn ven mang tonoplast (rndng brin thim bao quanh
h6a bang ciich duy
khdng biro). Khi cytokinin duoc xir Iy ldn lii cdy ua vdng hoac rri di6p vdi gio
truoc
khi dLra ra dnh sdng,
ti€n lap th€ s€ duoc chuydn h6a thdnh luc lap va k€t qud ld c6
su gia tang sr,r sin sinh di€p luc to. cytokinin cfrng ldm giim su ldo h6a cua
hoa cit
canh va rau tuoi. Kinetin co khi nang kich thich su v6n chuyCn cia nhtng ho-p ch6t
hfu co trong Ia di cat vd giir trong taii. Khe nang cua cytokinin kich rhich su vin
chuy€n dinh dudng va tao ra suc chua di dusc bi€t tr€n nhiiu iorii.

3..1. Abscisic acid

3.4.1. Sinh t6ng hopabscisic aciel

or-t

't
)H oorl
]H:
c:o
6'-hydro .\y me t hr'l-AUA i^o
/-u
I
t/Oll
f,, 'l/
I
.\u ilo\ i/
crHr { \_rtl
1- 'oH
A bscisy l-p-D-glucopyranoside
(_) i Oll
IJ:
I)haseic ac id

oH OOFI
II:
-l'-clihi drophascic nc id

f]inh 3 I I Su chu1,t;n ltoo ttia ahsc'i.sic at,itl

isopicnc llrinlr 3. I l) d slur trons luc lap vl) nliirng biio khiic
thc'o cott tlLririrq chir trirrh rricvaionic acid. Co hai corr clrrong d.i tao thiinh ABA sau
g d0
klti Itrcr:tlort'c tlro tltinh isopr-nrcrrvl p-rrophosfiiaTFllTLrlmi_F-Tlrilil]Lbn 6,r,ru
__,__L
__ a -_ -:_-:__. .
. :.__.____:__,__j__ __-__ , j_ __- -- -
-
@itop.nr.t'"t1,1".r1**'prrr,.qir-t,rti,-[t^rocdl--]rior,]ilnh-
() o |ritt|; ('t:,ii 1,r,i , I/r,i' .llr,/, l r.r;,.E ; ".:.1fr-
Brrcqf=pUqp]:os]1hatc r)roi sinh
Con dusng rl]y hai la tu isopenren),|
pirophosphalgl4o-lL\1nh carotenoid roiqua rnot loat Ui6n Adi drsi
3.4.2. Sg hit hoat ciia abscisic acid
ABA co th€ duoc chut,€n hoa theo hai con ducrng khdc nhau: N6 co tlr6
ducrc bi6n OOi thann abscisl'l-p-D-glucosepyranoside crla niot phan ung thudn
Irghich hoae no co tlre bierr doi khon,e thudrr nghich thanh 6'-irvdloxvrncthll AIIA.
phase ic acid houc -1'-dihvdrophaseic acid (hinh 3.ll) ABA cing co ttrd trir horr btii
,:,
su ket hop cira glucosc voi nh6rn carboxy-l cia ABA d€ tao thanh ntot ABA-glucose
ester.

3.4.3. Nhfrng dnh huong sinh li cria ebscisic acid

ABA phan phoi rong rong gioi thgc vat nhu thqc vdt bdc cao, rdu, tiio IUc,
i ., ,^ - ,I
ndm. nroi day cDng thdy trong ndo chudt. Tuy nhi€n n6 chua duoc tirn thdy trong vi
khuin. Abscisic acid ld ntdt san phirn tu nhien li€n quan d0n nhidu linh vurc sinh
.:
truong vd phat tri6n cua thuc vdt nhu r?c ch€ sinh truong, kfch thich sy phdr tridn
phdi binh thuon-q, tich lfry proteins trong h6t vd sg <ldp ring ndi bat vdi stress. ABA
hoat dong nhu rrrot tinlr hitu ciro bi6t ciy troug tinh traiig stress, tuy nhiOn, no cfrng
li6n quan voi nhfng qud trinh sinh ly th6ng thuong trong cdy. Sau dAy ld nhlrng dnh
huons sinh I'i cua ABA:

Khdng xir ly

Sv hdp rhu kati dvqc di dong nhd

bottt phu thudc ATP di K dvqc
Nuoc
tao ndng ri hdo bao v(

Yr= Yr+ Y,

,.lctd hitr crr trhtt nrrtlic'ocid lthine nin tin tlo r)iing X'& r,io lint
T-hi ndng nuitc rro
rhic.h ltrrtron I1- khi K- di ,'Ao
giam thi ndng nvoc. cho phep nrtlc tli vdo va
Hdu no di quang lop va ho hrip .xar ru
I

I ** ry ABA J
Y
x,i'ry ABA

Nrdc.

Nuoc

..1ll,l ric e ht; h/tm protofi phu tlruoc n BA ngdn tlitng K- vdo ri biro vA
.l l l' \'t r or tltlp l u h D! cun It o. nroc rt ra ngodi gdv nElsy giam
kltdu vin ding stic truong lan khiu dting loi

Hinh 3.12. Anh hud'ng cua abscisic acid lAn stl d6ng m6' khau

Guo frinh Chir Dtur lloa Su t lironp Tlwc l'dt 39

 - Ddng no kh.i kh,u.' vai tro quan trong cua ABA nhu ri
m6r chdt cdm rmg
vui srress da duoc bi6t trong nhiiu nirn qua. xri Ii ABA ngoai
sinh gdy done khi
khau trong didu kien siing vd dul' tri cho d6n khi ABA bi Juyiin
h6a. Trong di6u
ki6n stress do thi€u nuoc ABA co thd gia tang l€n 20 [,n.
Su stress do thi6u n*'c o
16 cirng sa' sinh ra ABA 16; cli ctruydn d6n Li va
roan cay cci lam cho khi khiu trong
Iai. Khi ca-v nhan 6nh sang trong diOu kien kh6ng bi stress,
co mot dong di chu,v6n
ctra porassium vao nhfng tc bao bao v€ nhu vao bsm
ATp-phu thurc voi K. ndm
trong rranu plasma cua t0 bao bao ve trong khi H- va cdc acid
htiu co nhu malic
acid du<vc chuy€n ra ngoiri. Khi di6u niy x6y ra. thii irang thirn thiu
'an tro ndn im
va re bao giarn th6 nirrg nuoc, nuoc divdo td bao vd ram
cti ttiu mo(hinh 3.12).
Khi ciy bi srress sd ldm cho K* di ra khdi tii bao bao v€ trong
khi IJr vd ciic acid
hru co <Ii viio lam cho khdu d6ng lai. ABA ngdn ciin su rno khi khiu
trong diAu ki6n
sang birng ciich cin tro quii trinh tr€n cho d€n khi bi chuy€n
hoa hodn toan.

- Brio vo ch6ng lgi

mtjn vi nhiQt dQ: Lrim ruong ABA gia tang khi
stress ilo
cay bi stress do min, ranh va nong. Nhirng su bitn aoi nay ri
nguyen nhdn crra su
r,f1eu vi€c xu l]' ABA ngoai sinh c6 thti rdm cho rot *oiol ciy chong
1uoc, iai
dieu ki€n lanh vi min.

- rlliEn /rcrg.'Trong diiu kien ngiy n-ean. ham luong ABA gia rang
rrong Iii
vi nram chtli dir dnn d6n su rnidn rrang. 1'uy nhi€n cfrng cd
rruong r.,qp t.ng oiiu
kien n-uiry ngin giy ra su midn trang trong vai roai rai kh6ng
co su gia tang ABA ndi
sinh' Viec xri li ABA ngoai sinh l0n mirn'ch6i va lcn hor dii kich thich
nli€n trane
cira chirng.

- s4t'rung, sy ntiy tn6m ciu har vd srysinh trrrtng: Tirkhi rnoi duoc phdt
hidn. ABA (luoc xcrn la chdr gdy ndn sF rung lii. rrii v;i hoa. -l.hdt
ra iliiu ao tnong
ho:in toin dirng ABA kh6ng tnrc ri0p anh hurrng r€n su rung. AtlA co th0 tric
'a
d0ng giirn ti0p tdn quii trinh ltio h6a truoc tru<ing rhanh vi ldnr gia
tarng su san sinh
ethl'lene i'a cthylene ddnh thrrrc nr6t s0 gcne rien quan ct6n su rung.

Nsri'- nuy ngroi ta bi6t rirng ABA co nhidu dnh hrr<ins r.6
sinh I1,. sinh
h6a Vii phan tri trorrg h6t cirng nhu co rnirt pno u6n tronq l[rc 'rarr .rrv
hdt phiit tricln.
rrlricl. vi-ri tro trrrc ti.-ip cria AtlA cltii cric quii trinh ni_r,rin cr\n chLra ro.
'oi
3.5. Itlrvlenc I{"C-CH"-
ethl lene

IIinh 3 i3. Ctiu tric ctia e rln,!ene

(it'io l) itt!t t'ittir !tr,ir, llo,.lSi,ttt Tnto,,c IJ,.<liE

40
 Elhy'lenc li\ nr6t hvdrocarbon kh6ne, no don eiin. no kich thich s
trii rii.gir ra rri0t dip i'rng b0 ba trong nhlmg cd),dqu ]l;i I an ria-+arrg+s€-Exn su
uc chd vuon dai, gia ting sF nghi€ng va s.inh truong ngang cua thdn ddl irng r,o'i
___==..___=:__
!gry_!g. Ethylene c6 ctlu tnic dcrn gidn nhat so \/oi nhtn-e chit di€u hoa sinh
trudng thuc vdt da biet (hinh 3.l3).

3.5.1. Sinh t6ng ho'p ethylene

utro.-

cltr-s
('ll" Li€n quan d€n
I rl''>()r()rt i enzvnre kh6c cH1-s-cl11-cl tr-co-coo
'\l f l\t\'ltt
t-]
oo \ f "'''
lv1 I R- l.l, \ \/, n-cir-coo-
t /iasazrinase \
\/.
tl Y R-(0-('(X)
I t R k,,u,r,'
Nllr
An,4 t-
cH.1- s- cH'- cH' - cH - coo
f",-j-:-__-l
I ctt' I

i ['6>-oJ
l\-{l
I or{o}r
J
I

t_____!11!
A fll..l ttur.ltttsidrrsr.,
NI].,
Adc t-
\ ct.t1-s-cHt-cHr-cl-t -coo

------* pell,sm incs

orjofl
AdoMcl
()iloll
N4l'.\

ntr9..^,^t,-4() -Cl I., COo

lzt..
-
t-t2c coo
MACC

ACC: l-am inocyclopropane- | -carboxylic acid

SAM: S-adenosylmethionine
KMB : 2-keto-4-methylbutyratc
MACC: Malonyl ACC
Met: Methionine
MTA: M ethy lth ioadeninc
M l lt: Mcthylthioribose

Hinh 3.11. So do s{ sinh tong hop ethylene

Ethylenc ld chat dieu hoa sinh trudng th;rc vqt d dang khi, n6 c6 !i6n quan
,.i
nhi6u toi clc qud trinh sinh trudng ya phAt lri€n cua thuc vdt tir su ndy mam cia hdt
C a Trinh Cfuit Oriu I lo,i Stnlt 7i(qns Thtrc l't)t 4l
 d€n ldo h6a va ch€t. Su sinh t6ng hop ethylene Id m6t chu trinh dugc thd hi€n tot
nhdt so voi bdt ki chdt didu hoa sinh trudng thuc vdt da bi€t. Ethylene'rluoc tdng
hop tu-Methiorr,rrc nhu minh hoa trong hinh 3.14. Methionine duoc bi6n d6i thdnh
-=-.-.-
S-adenosylmethlbnine (SAM) theo con duong cia enzyme SAIII synthetase. Mot
phAn cua SAM ducrc lAp lai chu trinh trong chudi sau: Methyl thioadenosine thanh
Methll thioribosc, thanh Methyl thioribose-l-phosphate, thanh 2-keto-4- methyl
butyrate vii lro lai Methioninc. SAM cring-di theo hai con duong khiic, con dudng
thrr nhdt li
bidn d6i thann S-adenosylrnethyl thio propylamin b<ri enzyme SAM
decarboxylate va di vao sg sinh t6ng hop polyamine. Con dubng kh6c ld thdnh
l-aminricyclopropane-l-carboxy,lic acid (ACC) theo con duong cira enzyme ACC
., . ,.i ,;. ,
synthase vd biin d6i theo hai huong, n<i c6 thd duoc bi€n ddi thanh ethylene do

-AQ9_gf{qf hoirc thinh malonyl-ACC (m6t sin phAm cuOi bdt ho4t) theo con
duong cia enzyme ACC N-rnalonyl-transferase. I-ICN dugc phong thich ra trong sU
bitin d6iACC thdnh ethylene c6 thd k6t hop voi cystein d6 tao thdrnh cyanoalanine
' va F[2S theo con duong c.ua. enzyme p-cyanoalanine synthase. Qua nhDng thi
n-qhi€m drroc d6nh diu cho thiy rang methionine duoc sri dung theo hai cach: C-5
...
n6i voi sulphur duoc ldp lqi chu trin[ C-3, 4 dugc diing cho ethylene, C-2 trong
HCN. vi C-l duoc dirng de thanh lrip C02.

l.;
C6 nhiiu nghi6n crru v0 sinh tr)ng hqp cthylene tqrp trung u buoc giua SAlvl
va ethylene. Co nhi€u ydu to co ttrd t<ich thich su sinh tdng hop cthylenc, mQt trong
:4,
nhftng 1eu td do chinh ld ethylene. Quii trinh ndy duoc ggi li qud trinh tu xirc tdc
siin xuat ethylene . Ethylene truoc tiCn kich thich ACC oxidase theo sau bdi sg gia
,;,
ting dirng k0 hoat tinh cria ACC synthase. Ngiy nay; protein cria ACC synthase vd
ACC oxidase cli cluoc thanh loc vir gene cua n6 ctng dI dugc tich rong trong nhidu
rn6. z\CC N-malonyltransf'erase c0ng dd duoc thanh loc vd dic tinh h6a. Tu khi
I . ^^
nhirng scne vc1 ACC synthase vii ACC oxidase duoc thanh loc, chfrng dd duoc sri
, i
dung dc'l bo
' sun_u cho su chin cua trai. Nhtng kha ning cua no v€ di€u hoa nhirng

qud trinh sinh ly ciy trdng cDng dd duoc phdt hi0n.

3.5.2. Su kich thich t6ng hg'p ethylene cua Aurin

Auxins didu hoa su sdn xudt ethvlene dd dusc nghidn cuu bdi Zinmerman
ri Wilcoxon (1935). FIo di th6y'ring khi rip dung heteroauxin (lAA) ldn chdi ciy'
drl kich thich su sinh trudrrg n,uhi€ng cua nhlrn_e ciiy dugc xti l! vir khong xir lj khac
dLroc clit cilnt trong mot thirn*e kin. Su quan sar ni1' cung vrri khd nring cua aLrxin vd
cth)'L-n(' giil'ra rnot sf, anh ltrrnrg tuong tLr dI cho thiy ring nhtng diip ung niy li
tlruirc tirrh cua auxin co thd clLrtyc ga1, rn do crh';lene. V trrdng niy da khdng duoc
lLrr,r 1i cho nrai ddn 29 niun sau khi Morgan ra Ilall (1964) rrinh bay mQt quan h0

song phtrone giai thich nhiurg dap frng cr,ia aurin vn ethl'lene. Cd hai tdc giA CA dQc
Li,.j,t lriulr (' ::ir I)trtt !hu S,nh Tt:RD,,' rh,,. t i' {'
Lip phrit hi0n klri nirrg cua auiin kich thich sirrh lOne hqrp ctlrllcnc. Ngr)1 nar'. noi
. |:
chung rrguiri ta clrAll nllin lirtg auxili kich tliicli su sar srnh etlrvlcnc drrrrc drCrr lroa
r li

boi nruc do n6i sinh cira auxin (ltinh i. I 5).

..1('(' .\ -.\ /rrllrr ltrutsl,'t a.s.,

l'ri c nhin kfch thich T:ic nhAn kich thich

l rirrlhtxrns r at li r a ltou ltrrc Su ch in cua triii
Strr:ss do kho han vii ngap . Erh-vlene
: .. :
L()n lrung va nlilnr Dgnn' lan cons
S(r ch in cua trii
l-io heia ( l-a vii hoa) Tic nhi n ic chi
Strcss do nhi€t dd Cobalt choridr'
,-.: , ,^ : !,

Blassinostcroid. au r in, cytt'k in itt. ulcu litgn vcm Knl

clh)'lcnc v2r absc isic acid 2 J] L
Ca lciu m
Strcss Ozone
'l'ic nhf,n Ic ch0
Anrinoctlioxr,vi nyl glycinc ( AV(i )
Aminooxyacctic acid (AOA)

IIitfi 3. I 5. T6nt tut so' do su' sinh t6ng hop elhylene

liitt rlutttt din tuc nhan kic'lr thic'h va i'c cltc

3.5.3. SU's:in sinh cthylene do stress

Khuynh hu<ing cda stress ai tictr thich sdn xuiit ethylencdii duoc thay bang
rhuar ngu "stress cthylene". MQt s6 lugng lon cia stress nhu hoa chAt. kh6 han.
ngap lut. ph6ng xa. thiet hai do cdn trung, b€nh, thuong ton co khi, va nhirng loai
kh6c dd cho thiy kich thich sdn xudt ethylene (hinh 3.15). Stress ethylene co bi6n
dudng cin bdn vri duc.rc san rudt boi tC bdo song. Khi tC bao chAt khdng co su sdn
sinh ethylene.

3.5.4. Nhung dnh hu'ri'ng sinh li ctia ethylene

- chin ciio trdi: Nhfrng nguoi Ai Cqp c6 dai dd khOng bi6t rang slr san
Stt'
sinh cthylcne gia tang ld k6r qua cua vi€c gdy vtit thuong bing crich cit nhfng trii
va dc kich lhich su chin. Ngay r)ay str chin cua trii di ducrc biSt li do cth-r lenc dicu
hoa. Ethylene cing d6ng'rrrot vai tro thi6t y6u trong su chin crla trdi dA giri. Thuit
ngir "clirnactcric"(gid man duc) d6 am chi nhlrng triii sE chin trong sU ddp ung vtri
ethylene va "nonclirnacteric" (chua gid, chua min duc) arn chi nhfrng trdi s€ khdng
d6p ung voi ethl,lene. Ethylene tu n6ng do kh6ng th6 phdt hien cho ddn 0.1 dcn I
pLiL kich thich su chin.cfra trdi b6t 16 m$t sU tang tinh mdn duc trong h6 hip (r i clu
ffi 43
tao va l0). rrui lai nhDng tr;ii khong rnan duc t6ng hcrp rdt it ethylene vd khong duirc
kich thich clen chfn (vi du nho vd cherry). Thudt nglr climacteric (m5n duc) dugc
,'. ,. .- .l
'.
dLrng lain diu ti0n d€ chi
' su gia tang trong hd hap cLia lrdi, tuy nhi€n ngiry nay no
bao hdm su gia tang trong srin sinh ethylene. Sp t6ng hqp ethylene dI giim dang kd
trong nhirng ciy ca chua chuydn gene boi su th6 hi6n cta nhlng ciu trirc gene
khdng nhai cAnr voi ACC oxidase, ACC sl,nthase, boi su th0 hi€n cria cau trfrc gene
nhay cam rsi ACC deaminase va trong tdt ca cdc truong hop tri hodn mQt cach co !
nghia su chin cua trai. Ngdy nay, voi nhirng ciy chuy6n gene dI cho th6y mric dQ
, t, ,.^
giarn ethylt'ne crir Ii6n quan dOn su tri hodn sp chin. ACC N-MTase ld mdt enzyme
c<i kha ning clap ung voi su chuyin drli cua ACC thanh MACC bdrt hoat. Sg phar
hien niiy cho phep nhing nhd nghi€n cuu phdn lap gene ACC N-MTase va dilng
) ,,1
gcne niy d€- nhirng cay chuy€n gene keo dai sU chin cria trai ,ei6ng nhu v6i gette
'l-uy
dearninase . nhi€n d diy rndt gene cua cdy duoc dung thay vi m6t gene cta vi
khuiin dC bit hoat ACC.

- SLr phrit lri6n ctZq cfry con: Narn 1901,

-----"---
di minh hoa dap ung bd
Neljuborv
ba cho tha_v rdng ethylene uc ch6 su vuon dr)i, kich thic'h sr no r6ng theo chi6u
. ': theo chi€u ngang. Ngiy nay ngudi ta biet rang ethylene co
' ' tri€n
ngang vi su phat
thO irc ch6 hoflc kich thich su vuon dai cua than. rE hoic nhlrng co quan khiic. Su uc
che su vuon drii di co bieu hi6n nhanh va c6 the dao nguuc. Ethylcnc cflng co bi6u
tri6n kich tlrich su vuon diii o thdn va ri.-luy nhi€n, di€u niy xiy ra ti t6c dd chdm
hun la su uc ch0. Thoi gian giira hai su ki€n keo dii dd tluos ghi nh0n tron-e nhi€u
nghicn c[ru dan dCn gia thuy0t ring su kich thich sinh truong cira ethylene co the la
mOt dnh huong gidn tidp.

Ilinlt 3. i5. Etir)'ierrc kich thich s'tr ddp r'ntg bo ha ciq cdy dau Hir Len
 D[p ing b0 ba kich thich biri eihvlcnc co thi hoat ddng nhu nrQt co clrt t6n
tai trong ci1'con. Dhp Lrng bd ba dua tren tinh sinh truirng ngang. sir phong lcln ra
su Lrc ch0 chi0u cao than (hinh 3.16). Vidu: Khi mdt cdy ddu Hd Lan gip phai mot
thanh car: irhrr bd nrirt cung cua dit, <Jii, hodc r,6t nao d6 gidi h4n su nhd l6n crla 116,
no s0 dip irng voi su sinh truong bd ba, cho phdp n6 ddrn thung hoac di chung
quartlr chtrorrg rrgi.ri. do do cho phcp no \(ron toi lrc nrat ciir i,a phir rricir. Khi su sirr
sinh cth,r'lcnc tronq vung rn<ic cau cua ciy con lcn.f,t cao giup no du,v tri hinh moc
cliu chat che (dau lla l-an hoac ddu) cho phip n6 vuol qua ap luc r'6t lli khiva chanr
vcri dit nri khirng bi ton thuong. Khi cay con nh6n duoc 6nh sang do. su sdn sinh
ethylene sd giam. Kdt qui lei m6c urd. Didu nAy hgat dQng nhu rndt co ch€ an toan,
ngtin cansu rrd tnroc cua moc vi thucrng trin den cay con truoc khi nhd ra.

,- Srl'ltlttg: Su rung co the duoc dinh nghia nhu sg t6ch ra cua rndt co quan
hoAc m0t phan ciy tu cAy cha rng. Quii trinh rqng rdt quan tro{g trong ndng nghi€p
boivi su rung holc khong rung cta hoa. trdi va Id inh hudngddn nang suet vd hi€u
qui cria qdri trinh thu hoach. Ethylene duoc xem ld c6 vai tro tu nhi€n trong vi6c
didu hoa r6c tid rung. Ba yeu t6 ldm bing chung vd vai tro cta ethyiene trong sr,r
rung dd duoc d€ nghi bcri Reid (1985). Truoc nh6t sg sAn sinh ethylene gia tang
truoc l:hi ryng trong nhieu cs quan crira ciy dang rpng. Thri hai. xu ly ethylene tr0n
nhidu lodi ho{c nhtng hop chdt phong thich ethylene kich thich sU rung. Thu ba,
nhing chdt uc chd su sinh tdng hop ethylene hoic ric che hoat d6ng crla ethylene sG
, ,i
uc cne su runs.

-jWAJu!: Nhidu ndrn qua nguoi ta dd quan s:it ring kh6i tu gd tht'c ddy
tr6 hoa trong cdy khorn vd xodi, Ngdy nay irguoita biiit ring ethylene ld thdnh phAn
co bin trong kh6i gia tang qu6 trinh tr6 hoa. Ethylene trong phdn ldn truong hop ric
chtl rrO hoal tuy nhi6n nd c6 inh hudng kfch thich trong cdy kh6m (dua). xoai vd vii.
, -, i .
Cdy cir th€ dugc xu I! voi cthylene trgc ti6p holc thdng qua vi6c xir dung nhtng
chdt ph6ng thich ethylene nhu ethrel (2-chloroethyl phosphoric acid) ld mQt d4ng
,;
tdng hgp cua ethylene khi d the ldng voi pH thich hop thi khong sdn xuAt ethylene.
Tuy nhi€n.'khi pH duoc ndng l€n n6 phAn hny d6 tao thdnh ethylene hoac gian ti6p
voi auxin n6 kfch thich sdn sinh ethylene m6t c6ch tg nhi6n.

:.St! ldo hdt:.Ldo h6a c6 thd dugc dinh nghia nhu rnQt tlrdt bqi chung cua
' ':
nhi€u ' .:
phdn ung tong hgp truoc ,id kii pha
tC bao ch€t la rndt cria su sinh trudng cAy
tr6ng keo ddi tu giai doan chin hoan todn dtin cht5t vd duoc ddc tfnh h6a bdi su ph6n
': to, protein, hoac RNA cf,ng nhu nhfng ydu td khric.
rd di6p lrc Ci ld vd hoa duoc
xri If
voi ethylene ngo4i sinh kich thich qua trinh ldo h6a vd gia tang su sAn sinh
:
^:' chrl y rang
ethylene. CAn c6 nhfng truong ho. p ngoai 10, ethylene kh6ng don l6 li€n

Aiao Trinh Clrit Driu lloa Sinh liuong Thuc l'il 45

quan d€n qu6 lrinh lio
hoa. Bleecker ra ctv (1988) cho thdy rang tr€n ld cua cdy
Arapidopsis bi6n di khdng nhay carn voi ethylene (etrl giit di vdi toc clQ chdm hon
dang hoang dai; tuy nhi6n chirng van gia cii, dieu nAy cho th6y ring cthylene kh6ng
; : , :' ! 'T'L: --- -+.t -: -^ '
phdi y€u t6 duy nhat
l2r ' c6 li€n quan d0n 16o h6a. Them -.: vdo- d6 co nhi€u cdy to ra
Ithong gia tang san sinh ethylene trudc khi lio hoa. Nhlng truong hoo niy cO rh6 lAr
clo su -sia ting tinh nhiry ciim cira cay'rui cthylene hon ld su gia tang hdm luong
cthy lcnc.

- Nhii'ng inh huti'ng sinh khic: Irthylene cing co li€n qiran d€n quang
l1i
. , ^,1 :
hop, hd hdp. virn chuy€n. mi€n trang. su niy mam cua hot. nhu ch6i,
') uu th€ choi
nggn, sinlr truring ttl bao, nudi cdy rnri, thanh tnp phoi, sU nghi6ng, su khoi sinh rE,
nhtrng co quan dU trir, su thrinh l4p g5. sillrlrc ch€ rnim hoa, su phdt tririn gidi tinh,
dia hLrong dong, srr ri nhua vd su thirnh llip nhua mu cia ciry.

3.6. Brassinosteroid (BR)

3.6.1. Phiin logivir ciu trric h6a hoc

' 5cr-cholcstane bralSinolide

rl
6-deoxo Crq-BRs

Ho.'
- -
("',Hcy'.'-'u4-' i),, L,J ?"{
,r^\./\/
HOr.
/,,\rO/,,./dO\.{O
tttttl
6-oxo
/^r ttl
'oH
t"./HC tl
no'' VHo#not"
Y o
oH(
rll
,/\-a\./'
-oxallctone
tll
'oH
l) OH
\o
Vrtng lJ

Hinlt i t 7. i\,ihirng sr kluic bilt t'i catt tric lrottg t'ottg A, r'dng B
.,'ti tnttclt nlrdtth cuo cit' br{tssirtttsleroid tr.r nlti0rr
 Cac hcrp cttit nR. ru nhiin co ciu tlric co bin tir rnqch khun-q,5cr-cholestanc'
thinh 3.17). Srr rhav ddi criu rrlc co bin cia mach khung si hinh tlianh nltr'ing hop
chat r oi rtlttns cltirc niutg khac rthau.

- l''itng,4.' boiso luorrg vd vi tri

Su biirr doi crla vons A bi chi phoi chri 1'6u
cua nlrorn hydroxril lhirrh 3.17). Nhin chung nlrirng BR co nhonr diol hrtirorvi ki
cin (d vi tri C-lu ra Ci-3cr) tlrrrirng c6 hoirt linh sinh hqc uranh nhu BL ri\ CS. M6t
rii IIR clri co gr6t nfiripr h1'clroxll tr vons, A (o vi 1ri C-icr hoirc C-3p) nhu
tvphasterol ('l'\') r,a leastcronc (-l'l:). Ngorii ra con co hai BR co nh6nl 3-keto nhu
3-dehvdroteasteronc (3D'1-) r'a 3-deh1,dro-6-deoxoteasterone (6-deoxo3DT). m6t BR
i
co nrt)t nhorn 2p. 3$-epoxy' nhu secas(erone (SE). r,i hai BR c6 nh6m hydroxl'l o
vong A (hinh i.l8).

- l/brrg B. Voi ba dang duoc bi6t la 7-oxalactone (7-oxa-6-onc), 6-oxo

(6-ketonc) r,i 6-deoxo (hinh 3.17). Nhin chung. loai 7-oxalaclone c6 hoqt tirrh sinh
hoc manh hon loai 6-oxo. vi] loqi 6-deoxo co hoat tinh sinh hqc yeu nhdt.

- Mach nhdnh: C6 tdrn nhom mqch nhdnh d6i vdi nhirng BR co ngu6n gdc
tu nhi€n nhu md ti trong hinh 3. I 7 voi t6ng so carbon lit 27 ,28 vi29.

3.6.2. Sinh tdng hop brassinosteroid

Brassinostcroid trong tu nhien thuong la nhtng steroid co 27., 28 t'a 29
carbon. Khung carbon cua Ilqch nhdnh ndy cing phd bi6n trong sterol thuc v4t. BL
{amdt steroid c6 28 carbon voi ciiu tnic suon carbon tuong tg nhu campesterol,, m6t
hop ch6t cb nguon -eoc duc.rc t6ng hop tu mevalonic acid. Cdc BR co 28 carbon nhu
castasterone, ryphasterol vd teasterone duoc xern nhu tidn ch6t sinh tdng hop BL tu
campesrerol(hinh 3.19). Llnr diu tucampesterold6n teasterone r6i cliuy€n O6itnann
typhasterol. lyphasterol s0 duuc hydroxyl hoa o vi tri carbon thir hai de tao thdnh
castaslerone. SF oxy hoa Baeyer-Villiger d vi tri C6 cua casl.asterotle s0 tqo thanh
nim l99l d6n narn 1997, Sakurai vd cQng tdc vi€n da dung hQ
brassinolide. Tri
th5ng nu6i c6y, mo riri phAn hoic h6r chLra truong thdnh cua cA1' dua can
(Carharanthus roseus) !d xac dinh chu trinh sinh tdng hon brassinolide tu
campesrerol (Sal<urai va ctr,. l99l;Sakurai vA Fujioka,1997 Fu.fioka vi Sakurai,
1997). Ilai con duong song song di6n ti sp bi0n cloi tu carnpesterol cl6n brassinolide
dd tlugc dC nghi do la c,on cltrd'ng oxy hoa C6 s6'm vd, cott dttd'ng oxy htia C6 nttott.
Trong con duong oxy h6a cl6 som co nhtng ch6t bien dudng bi oxy h6a tqo thdnh
tu campesierol lir 6o-hydroxycampeslanol vd 6,-oxocampestanol- Trong con ducrng
ox1,' hoa C6 rnu6n, rnQt h-o. p ch6t khdng co chuc ndng ketone o C6 la
6-deoxocastaslerone di duo-c tao thanh vd bi oxy hoa d'5 Uien d6i thanh castasteronc.
cai t;inn Cnh Diiu Hoa Snh liuang Thtc t tir 47'
 C2s BRs (24o-methyl)

?nl -
t-5" n
Ha'r^
no'"' Ho"t.-
ll rlrsrnolrrle ([]l- )
( attsrcrrrne (C 5J 2-l:piCS

?nt. n, ?*
l-\{,^v
|

.t. 6" ,^.1,,^.. 6H

I

Ho\. L - -.' ' ."'#

til .,'L
p,o'' "i -' HoA-f/ d*'A
2 3-DIcprCS 3-DchyrlroTE

H O,"

HO' nO.\-fi--- 6- DeoxoT E

j- l- pr-6-deoxoCS 6-DeoxoTY

'l'r"''(
6nI

no/ i'i
o
ftronc
Se€asttronc Clthasffonc 2-DeoxyBL

Crn BRs (24B-methyl)

.J_1t-
n
iii
n/\-'\' 'l

i 'o
'l E-l-nryristate (R =
C13t117) lJ-Eprl)l' lJ-l:piCS j.?l- DicpiCS
I-l--i-lauratc(R = C11ll1;)
Cls BRs (24-methvlene)

i
.\.,..r,W Fl
l| ,'^tl{&'
Ho.a^Jr\#
til
o Ho \-il*'
JJ [.pi-6'lcoroCS Dolicholide tDL) GII'oxolX
i)rrlrchostcronc 1 l)5 )

Gio lrit ('h.it Dt,\t ,'lo,i S th 'll.rorry I htr i'dr

 C.e BRs (2{d-cthrl)

OH

HO- HO'
)

HO .U HO
HO
o
lli'llonro l \ 2li'llorrto Il:
lli-llontolll. 2li-lltrrlo('S

C1e BRs (2{-cthylidcnc) C19 lilts (24-mcthl lcnc vir crlrl 25-ntcthl l

HO HO-

HO HO
H
2 E-l l0nroD l. 2 8.1 lorno DS {>Lhm>2ll-lxxrnDS 25-N4ctlD ll)s(25-Mcl)S)

to
3-tipi-2{eoxy-?5-Mr:l)S
2.3- l) rupi-25-\1cl)S 2-llo\\ -25-McDS

C27 Blls (no 24-su bstitucn l)

H,

,. O
-o
28-NorBL

,/9'L
l:
,.^,v,,
HO- | '-'),
ctt,((tu
- 4\r
! -,r-
HO
qlt.coo --' - o
H6
TS303

Hinh 3. t 8. Cdng tlnrc cau tao cia cac brassinrtsleroid

dao frirrt, o,7rtlti, tlod S:,l,h 7 ruong Tttttc l'ir 49

 Nlevalon ic acid

.tl

6-Dso\uaathi!tlarone
gHl a)
I
t)
t '/t'(
o"
*

I
r-k\
- -.-J,,...--
I
ii cl)
Ho' ' x -'
+o

o
| - x)-,..
Y -r-l
r' -f -l
6H

l,-.l-../-'
o..-.-lr.-',
.,1-l)ch;"dro-6-dco\olcilslerunc
oHl
t-
V rl.6rr
--i-) I

r
_t \.]-'
3g \"--;'---'
iT
o- Duosirtr Phlsterol
t-t t J FI lr phnslRrol

+
H

-_>
lr - [ )('r] \ (rclltl.lSltrotlc FX C.r>r,r:tcrtrrc
(ru Ilr dt trrYc r stlr te't trnc

tlinh i tt) So'li,: sr sittlt lorryi ltop bt'assinolilt

(,u:ui,li,tl, (lt.it l\titt l!,.1t \tu\ h tnttt:: fh\tc l''t! --i0

3.6.3. Nhu'ng inh huo'ng sinh lf
cfia brassinostcroids
BR co arth lttrtiltg trrin Ithi€u dirc tinh sinh 11 cua thuc rit voi nong dir rir thip
nhu auh htro'ng l0n su sinli trurirg nghidng. kich thich su vuon dii. BR rit cin thidt
. .:
cho su phit tricln binh thuirng crla thuc liit. n0u cAv thicu BR se din dcn nlitlrng bi€n di
bal thtrong. Trorrg none nrhi0p. BII ducrc tht nghi6nt gia tang sr,r c'hin ra gia laltg
rtittg suit ttrui ctra ntu rti:irr. trrrrrt tinlt diu triii cung rtltu gia tirng rriurc. su3t liot crra nlir
i. . - ,:
coc la ci1 lii cri (Atrc. 1989). Ill{ kh6ng chi gia tang ndng sudt nra con _qitp ciiv
clrong chiu lai strcss elo ntr-ri trrrirng nhu khiing bernlt. uc ch€ su lQt xiic cua con trun!..
gidrn dQc t6 ilo thuoc cri. chong chiu min. chiu lanh. BR con co anh huong l6n su gia
-: tong
tiing hc.rp protein. DNA va RNA.

3.6.3.1. Anh hu'6'ng ciia BR lOn su'sinh trud'ng nghiGng

Khui diu

D6i ch*ng

IlL. l0-r0 M

BL IO'9 M

BL l0-'r M

Hinh 3 20. ,1nh htil'rtg c'tia nong do BL l1n sy nghiEng ctia phi,ln la ltia
r Ktroidau a D6i chung

IB{J
o BLto "'M e BL to-"v
16rJ s Bl.ro*v
l{o r29.E

120

9,-
.? ov
..l
v6a
40
n
o
.2'rE4qE!q
iTaZ?.:&.-:=
t5=aE=6=2 - Gl(,ng hie

Hinh 3.2 t. Sv nhgy cam cia cac gi6ng lua vdi BL t€n st1'nghiEng ctia phi6n td ."

Sq sinir trudng nghidng cua phi0n.16 lrla di tlugc dirng nhu phuong phap
pnO fidn nhdt cho sinh irac nghi0m BR vdi tdn goi thuong dirng ld rice-lantina joint
/esr (LJT) hay rira-lamina inclination test (h\nh 3.20).T6t ca cAc BR diu co anh
huong l€n su sinh trudng nghi€ng cta phi6n 16 lira phi6n td hia cing t6 ra rdt vi
nh4y cim r'6i BR. BL vd homobrassinolide (homoBL) c6 titc dQng l€n sr.r sinh trac
nghi€m na1'rnanh hon IAA l0000lAn (Abe vd Marurno, l99l). Cric gi6ng hla duoc
(;fio liih Ch.it DEn Iloi Smh Titone ThLrc I it 5l
 trong o Vier Nam nhu t{6p Thom, Ndng Thom Chq Edo, Tai Nguy€n, Thom Lun,
IR50404 vd MTL 147 ctng to ra r6t nhay cdm vsi BL (hinh 3.21).

3.6.3.2. Anh huring cria BR lOn su vutrn cliri

C l,ria
160
t Cri ldng vu c
140

t20

.100
EI
'" ,ic 80
E ,:60
,G:
!.:40
'l lzc
s0
F.l ,J
'-.1
kr tr qr
414
.1 aO

Hinh 3.22. ,inh hrong cia BL l1n nrwron ctai cia be la ltia vd ci ling ,trc
(**) Sr khdc bi|t cd 1t nghTa o m{rc I% trong phidp thtr T

Ill{ kich thich sLr vrrcvn dai trOn nhieu loiri thuc vat voi ndng dQ rit thdp. Sg
ruon ckii cu.r lting thu hai trCn ckiu co vc duoc kich thich r6t manh boiepilll-. K€t qud
thinghiQm.riiy cirng drroc phirt triin thanh phcp sinh tric nghidm vdi long thu haitrin
dqru cd ve (Kohout va ctv. l99l). Vi0c xri $ epiBL da kich thich sv vr-rtm dii cia truc

thuon-e di0p dua lco dong thoi kich thich su tfch liy duong trong truc thuong di€p
(Naka.limavd'fo-v-ama. 1995),Xril1i Bt.voi nongdott0.l-l0pMkichthichmanh
I nre srr vrron dii cia diep ti0u, tru trung diQp.trong t6i. BL I - l0 pM kich thich sg
vucvn dai cua bclii lua (hinh 3.22). gia ting sd lA. nhung lqi uc ch€ chieu cao cdr lia.
Anh hrrong nay cing dutrc quan siit tr€n co l6ng vuc voi mdt su nh.1y carn mqnh hon
d lua. Nhin chung tsR khong cti iinh huong r0 l€n su vuon dai cira 16, n6ng clo BR hoi
cao dd giy,irc cbi srr vuon di\i cua 161cbron. N.iv{ vri ctv. 1998, 2000-2.001).

3.6.3.3. BR cin thi6t cho su'phit tri6n binh thudng cua thu'c vf,t
Srr thi6u hut IIR din den bicin di cav lirn dii duoc plrdt hi€n o ciy
It'rthitloJt.si.: hicrn di rirr/7 (Choc va ctr'. 1999). det2 (l.i vi\ ctr'. 19961 l-i va Chorv.
1997t. rhlll ri
ctr. l9t)S). lri,.in di /t-b (hinh.].lj) tr0n tliiu I'Ll l.an (Nonrura
(Choc
rri ctr. 199,'). iii biin tli rfu1,trtn ca chua di dusc phit hi0n (Koka va ctv,2000).
Viic srr li f{l da giup nlrr-rrrg ciii bien di lrrn ni1'phir trien binh thuung tro lai. Dieu
rrrir chtrrrg'o BR co rrrtlt rai tro sinh lr rit qrran trong, tronq vicc phat tri,-in binh
tllrrong cr'irt ''ar'.
-i;, D,;; il,r' s.'h r,,",L ' j2
'i;7;;'t,
 Iko Ika + llL Ikb IKb + BL WT
(Nomura el al. 1997)

I'linh 3.23. Biin di tltieu brassittosreroid lkb drgc phqc hAi mng BL ngogi sinh

3.6.3.4. sr ch6ng clriu vai diiu ki6n khic nghidt cria m6i trutrng, tinh khring sf,u
benh vi
tinh ch6ng chiu voi thu6c c6
- Dtip futg vhi itiiu ki1n lqnh t'ti tinh chiu nhipt
Nh[ng h0t lira va rrra lua non khi co hai Li vdo dung dich BL c6 nong d6 ru
| 0-i den lO'apprn giup nra chiu lanh rot hon. phun hoic nhirng cdy dua leo con va ca1,,
. ,:'
ca phoi voi llL
o giai doan Lrd hoa cfrn-e girip ching gia tang linh chiu lanh. BL co th€
ldm gia tang hdrn tuqng difp luc t6 rrong li bip khi duoc xri ry d nhiQt clg t g0c. su
r r l. . i
phuc hoi cua cay bip
khi xu I'i lanh cfing ducrc quan siit (He vd clv. l99l ). cdy cdi
sau
diu va cd chua duoc x& ly voi cpiBL cfrng ch6ng chiu cluoc v6i nhi€t do cao rdt tot.
Licn quan din co ch6 clrong chiu nhicr niy c6 su hinh thanh /up protein (heat slrock
protcin) khi cdy duoc xu l!
voi cpiBl (Dhaubhadel vd ctv, 1999). BL (105 - l0-6
ppm) c[ng girip tdng su phuc hdi cua cay lua rrong trong dung dich mu6i stio-zooo
ppm (Abe. 1989). Cdy l€ hay c6y bcnjamin duoc xri lV vdi hqp chdt c6 chua BR vd
propyl dihydro.lasrnonate crng ting duoc tinh chong chiu han-(Karnuro vd
-eia
Takatsuto. 1999).

-
Thth Khring sdu b1nh vd thth chilng chlu v6'i thuAc co
BL cfrng girip cay ch6ng chiu lai vdi b€nh dOni van r6n l[a, hio muOn tren cir
. .,i.
chua. th6i nhun bip cai. vd mOc xdrn dua lco khi phun voi n6ng dQ tu l0-r d6n
lO-"ppm. BL cfrng giup tang cuong hidu qu6 cria validamycin a6i voi b6nh donr vin
tren hia (Abe, 1 989). BR co cdu rrfic rat gi6ng ecdysteroid. nhom hornrone l6t xac cua
l.;
nhr€u loar cOn trung va dong vdt ngdnh chdn khdp. Chinh cdu trric tuong trr ndy dd
giirp cho BR ngan cdn su l6t xiic cria mot so con trirng, vd BR duoc xem nhu la rn6r
antiecdystcnrid. FlornoBl co c6Lu truc gAn gi6ng vdi ecdysone (hinh 3.2a), no dt Iirn
Grio linh ('har lloi Sruh TntongThvc I rit )J
keo. ddri su lot xdc cua con gidn giai doan nhong tran k! cu6i th€m I I ngiy (Khripach
vi ctv, 1{)99).BR con ldm gidm t6n thuong do thudc c6 d lira do simctrin, butachlor
va pretilachlor hay o lira mi cio simazine.'fuy nhi€n. BR kh6ng lam gidm ton thuong
ch<r voi thuoc co g6c auxin nhu 2.4 D vir MCP md con gia ting hi€u lUc
cdy trong <1oi

cua crrc thu6c cc g6c auxin nay (Abe. 1989).

HO,;

t-l\J
Homobress inolide

Hinh 3.24. Cdu trtic cua brassinolide vd ecdysone

3.6.i.5. liich thich sq'sinh tdng hop ethylene

U Dtii chung 18
0.i0
o D6i cnung
tZ l0p\ f tll. to
-; {} ri a 1OpM BL

Ln -)n rr- 12

(.; 2,"
6U. l) a
i8
;
E
(Jn ln FA

E
c n n(

t).00 0

l.iru Co l0rrg r trc Lr.ra co l6ng wc

tlirilt J..15. Atilr htrctru, ciu llL lt)n sv hinlr Ilinh 3.26. Anh hvottg ciut BL lEn sv hinlr
tltiutlt etlr.t'!etrc tt ltio vi ci lting ,'tx' ticlt !uf HC.t{ o litt't'i ci lirrg urrc
(+| t Sr Alt,it' hitt t'i.t ngliu rt nic |'It tntng plti'p thrr'l ('+t Srr khtic' biit ci 11 nghfu i nrrc l')i' trong phip tlni T

liirn gia t:rrre str sinh tong hopcthrlcnc o trrrc hu dirlp tlitr xattlr hine crich
lll.
kich thich str ltoat dt)ng ctil crlzltnc ACC s1 nthasc (Artcca vi cll, 1988) t)1- kich
rIich str sirrlr tthl'lcnc tir chtii l[rir trorr! dicu kiin nha lrrui cirrrg dii dtrclc blo ciio
(l:rriii. lJt)l) thicli sirrh trirrg lrup cthylcrtc- criir BR drt dtrtrc bicit li do gcnc
Srr kich
clii.tr ktri.in. EpiBI- larn sia tang str san sinh c-th1'lene u truc ha ditp drlu ranh ddng thoi
r.tii srr th0 hi0n cua ACC si nthase cDN,,\ la VlL-ACS7. Trong khi ir\A anh htrong lOn
srr tltc hieln cLir i i\CC svnilusc cDN.'\ lo p\rli-r\CSl. pVR-ACS6 vi pVR-.\CS7 (Yi

t i:,i,t li'i:h t rr.j; i)r.ir //irl -\rrlr 'liut:lrt ih:it l ,ir

 Vil ct\'. I 999 ). Khi xti lv ts I{ r'o i rrc\rrs do cao. su sinh r6n-e Irop e thr,lcne eia rin_e rit f d
cr lua lii co lOng vuc (chon vii crr,. 200r)(hirih 3.2j va 3.26). Sau khixu lj,BL vo.i
ndng do tu 0. 1 d0n l0pt. harrr luonq ethr'lene sirrh ra tir lria tine ltp rAt ro.
O co l6ng
r.uc. khong chi hanr lLrong ctltvlcne tins l6n r0 r€t ntA cdn co srr tich l[ir,
cr,"n'de
(HCN). Su tich lirl'c1'anide o co cao hon o lua li clo hoat clons. cia cnz\'nre
B-cl anoalanill svnthilse. cnz\ nlc phin hLiv cvaniilc. 1c{u htrn cy lLra.irrh lrutipq irc c6C
ctla Il[- none d6 cao lOn str sirtlr truting l[a drrrrc giri dinh la do srisrjrr sinh ethl.lcnc.
'l'ronc
khi tli' alrlr hutlltt nril lrr.itt co l0rrt r,uc drrtrc \cnl la anh htrtl.rrg cua su sin xuiit
eth\ lr'nc va su lich lllr, c)an'dc (hinh 3.27).

Sg'ri'c clr0 Sr1'^ric chC

Ethr'len Efhvlcnc

ACC 0ridasc ACC 0ridasc

ACC

nthasc CC nthasc

i\leth ion i
;- ^." '' !
Hinh 3 27. Gia thu1,s1 vi cct ch1 tdc dong cua BL d ttiq vd c.o ling vuc

3.6.3.6. Khi
n:ing ri'ng dgng cria brassinosteroid
N-aiiy nay'. cri r6t nhidu nghrcn cuu tim hieu vd su ly trich' sinh t6ng hop, co
, : , ,.1
che lac dung. su br€n duong, tong hop nhan tao, vai tro sinh ly va nhftng dnh hudng
cua BR len su rh0 hi€n gene va bicn d6i protein. Cac n_ehi€n cfu ung dung
brassinosteroids vio thuc ti6n sdn xu6t cing ducrc rdt nhi6u nhd khoa hoc va nhi sin
xu6t quan Ldrn.

(ibo Trinh Chdt Diu lloi .\nh Trrong Thqtc I rit 55

 Nhi€u cdng ty n6ng duoc di co nhfng thi nghi€m thuc ti6n ae fnao s6t hi€u
qud cria brassinolicle, 24-epibrassinolidc vd 28-homobrassinolide l0n cdy tr6ng. K0t
'| : 3,l-epibrassinolide lam gia tang ti l€ niy mAm khi xu ly l€n hot. O Nga
qui cho thly
va liclarus ntl duoc dang ky chinh thuc tld xu l! hot cho nhiirr loai cAy rrong. co vir
cdy kyi sinh (Kamuro va Takatsuto. i999).

BR con tlugc su dqng dd lam gi;t t;ing s0 la. diSn rich la. trong luona tuci Ia
trong lutrng khCr cua lir vir 16. tuOi ki, s6 luong cua choi hay canh hiru hi€u. Nguoi ta
con dirns no d6 lam gia tang s6 gie tr€n bong cua ho hoa thdo, sd triii trCn hoa miru,
cay an qurl va cu dd ldm gia lang ndng suilt. Krishnan vi cgng tdc vi€n ( 1999) ctng dd
tarn gia ting cluoc s6 luong h4t chic, kich thuoc hat vi trong lugng hat lua IR50 bing
. ,., i'
r iic xu iyi kdt hqrp giira BR va bcnzyl;rminopurine. Cic sdn phdm nong ducrc cua BR
thuong li
nhirng sin phim co ch[ra BL. castasterone, epibrassinolide hay
-ein day co
ch[ra thgn T'S 303.

3.7. Salicylatc (SA)

cooH

LIinh 3.23. Cou trtic cia salicylic

Salicvlatc la mOt nlrirrrr h('tp ch{it cir hoat tinh tuung tu nhu nhu salicylic acid
(ortho-h1'droxvbcnzoic acid) (hinh i.28) lLr rndt phenolic thuc viit. Nhung phenolic
iluuc dinh' nghia nhu nhirng clrir co vong hvclrocarbon thonr mang mot nh6m
hvdrorvl hoirc diln xulit crit nti. Saliolic rcitl tl;.r duoc tim thiy phin bd rQng othuc
, .i
riit bric uao, cho rh.1n nuv nir cL'i duoc nhin dieln tron_q hon 31 loii thtrc vit. Salicylic

aciil clir Jutrc tiur thal' o lii vii nhirng bQ pltin sinh san t'.r.ra thuc viit. vtii mirc dd cao
tr()ns nlrirng ctrrrr lroa cua cil sirrh nhir-1t va nlrring cin bi nhidm clo rninr burnh gi1'
hoai tr,r.

1.7.l. Sinh tdng hr.r'p srlicl'lic acitl

['htr tlirrlr slrikinrrc ucitl siir]r rl cinrr:rrrric lcicl rt\i sirLr (l\) biin dtii thc.l lrli
clich rl,-i tlo thi\nh salicvlic acid. Clich thrr nhar thcrr corr dtrorrg cLin srr tuo thanh
bcnzoic lcid. ciich tlrfr hui thco con tlLrun_e cua srr tao thi\nh O-coLrntaric acid (hinh
3.29).

rir,ir: li,inlt (::.i: lJn:u ll,t t:-th liurL,rg llurc l,ir i6

 cooH

Phenl,lalanine

,f

[-\, -,V-'oo'
\:/
s-Cinnam ic acid

O-Coumaric acid

saricvric a?ilJ

Hinh 3.29. Clru trinh sinh tong lwp salicylic acid d thu.c vat

3.7.2. Anh hu'6'ng sinh lf

Salicylic acid da cho th6y co anh huring trdn nhidu qu6 trinh crira thuc vdt. Su
tr6 hoa. sp phrir sinh nhiet trong nhlrng cily -sinh nhi6t, vd kfch thfch rinh kh6ng benh
la nhirng quil trinh rnd salicylic acid c6 nhtrng dnh hucmg chinh. Aspirin
. (acetylsalicylic acid) hoat dong gidng nhu salicl,lic acid vi n6 dugc chuy6n thdnh
salicl'lic acid trong dung dich cia nudc trong thuc vdt cfrng nhu trong h€ th6ng d6ng
vAt.

Ortio lritfi ('hit D u llti Sruh liutngfhuc t'At 5j

 - Ai'nh huting l€n td
hot: Salicylic acid c6 thd kich rhich su tr6 hoa. c6
sqt

thong tin cho ring SA duoc xu lj ker hop voi kinerin va IAA rhfrc diy sq t4o mAm hoa
ciiy thudc lii trong nu6i c6Ly m6. 1-uy nhien cing co nh&ng k6t qui tr6i nguoc vd udn
,:. , :
eli nay va cho rdng co nhi€u hcrp chdr dnh hu0ng l€n su t4o mim hoa ndy khong ri€ng
gi SA.

- fulai quun hp gifirt su'lic-vlic acid vti str phdr sinlt nhi6t trong ciiy: Su phat
sinh nhi€t trorrg ciiv hoa loa ken duoc xiic dinh ldr do SA- r6c dong. SA gdy ra su ph6t
nhi€t d0 lanr hay hui nhung hop chir amin va indolc c6 n'riri {li hip dan con trune d6n

-eirip
cho su thu phin. Nhi6t d6 xung quanh hoa c6 thd tang l€n l.l0C.

- fulii quan hE giiro salicytic trcirl vir tinh khdng blnh trong cd;, : p1U, u2l,
loai cay khhng b"^nh gioihqn su phatui6n cua vet b€nh rhanh mot vung nho quanh noi
xarrr nhi6rn cua rnirn b0nh lfic dau voi m0t vdt hoai tu xLrit hicn. td ueo tu ch€t ad uao
vc drrrrc goi ln phan ung quii nhay cam. I Ii6n tuong niy thuong cli kcm voi su san xuit
ra mdt loai protein co trong luong phdn tu nho. Tinh kh6ng benh vd sg san sinh ra
protein nay co thd duoc tao ra do SAhodc acetylsalicylic acid ngay ci khi khone co
nrin bcnlr xiirn nhi6m. Trong quii rrinh phiit tri€n cua nhtng ciiy si€u nhay, cdm dtip
trng vcri rniim b0nh. mot s6 luong ltin SA di ducrc sinh ra ttr cinnamic acid trong vins
rrhrr cin crtr ', il bclnh ht-riri tu.

3.8. Jasnronatc (.lA)

ttinlt 3.30. Cau trtic ctiq iasmonete

Jasmonate la rnot nhom dic bidt cia nhftne htrp ch6t oclopentanone voi hoat
tinh ttrune ttr nhtr (-) jasrnonic acid (hinh 3.30) r'ir mcthvl estcrcira nir..lasmonatr cld
duoc phtit hi"3n tronq 206 thtrc v iit dai die n cho l j0 hq gorn duong xi, reu vii niim
loii
, ., l : ,
cho thiil mnq ch[rns phan phoi rong lrong gioi thuc vat.

1.8.1. Sinh fdrrg ho'p,, chut'On hoi r,:r vin chuvGn jasnronatc
\l.ie rlLr cir it hiritr birit vC i i":c sirrh ,|,,ln [,rp jlsntonirtc. r.hirrrg cfr clir clr,r tha-r
trtrrtg clirtit thirt. lri non. triii clrrra chin lii chtlp rj chLra mor hdm lirons cao Jr\.
.litsttlonatc'dtrrrc sinh torrg hgp tu linolenic acid boi rnrit loar phirn frne trong hinh
i.3 l. Str vin clru\'€rr. su clinh vi trong nOi biio. r,a su dic\u hoa qLra trinh sinh tjne hop

(i;it; liit:lt('l;ir /)i,r I s,i.\!rh ljn:,:t4iiuc i,it

5S
.iasrrtonic acid thi chua duoc bi€t nlfiiu. Hi€n tai chua co nh&ns nrinh cliune. trrrc tiip
:.-rl
r'€ su r an chul cln cua JA tu noi tong ho-p dcln noi tdc dong.

\--l---.-,- -\----\'--\---\----\--cooH
I-inolenic ac id
I
I
I
.. -r - -\--\..- \ --\.\. -.-COOI I

oolt
I 3-H1'dropcroxl,-l inolenic ac id

COOH

(+)-7-Jasmonic acid

Hinh 3.3I. So di sinh tong hop jasmonale ti Iinolenic acid

3.8.2. Nhii'ng rinh hu'ri'ng sinh lj' ciia jasmonate

Jasrnonalc cho thdy c6 rnQt dnh huong rong rdi trong c6y. No lhe hi6n cd inh
. Vai tiic dQng cira no giting
nhu ABA vd ctlrylcnc. Vicc xu lj JA ngoai sinh co dnh huong uc ch€ su sinh truong
theo chi6u dc.rc cia cdy con. sy sinh trudng chidu ddi 16, sq sinh trudng n6rn rE, su sinlt
trudng md dang nu6i cAy, sq phat sinh ph6i.-sy ndy mAm cria hdt, sq ndy mdrn cua hot
': su hinlr thdnh nu hoa, sinh t6ng hop carotenoid, t4o thdnh di0p luc 16, t6ng hgp
phan.
rubisco va nhtng hoat dong quang hop. Bdn canh dnh huong uc che, JA cfrng c6 inh
huong kich thich hay gay cam uns tr€n su vuon ddi cta hom mia, su phan hoa trong
rn6 nudi cay, sU t?o thanh rd bdt dinh.' phd vo mien trang cta hot,, sg nay rn6m crja hat
phin. su ndy rnam choi mua d6ng. su chin cua trdi, sr.r 16o ho6 cira v6 quA vd ld. su
rung lil. su tqo cLi, su cuQn tua xcin, sg d6ng khdu. pha vo thoi vo sac, phAn rd diep luc
:,^,i : ,
t6, ho hap. sinh torrg hop ethylene vd protein.

r:dqTnnhCl t Drcu ll, tSnhlruungThuc I tit 59

 Jasmonic acid co anh huong tr€n sg th€ hidn gene trong nhi€u loai. Protein
sinh ra bdi ;asmonic acid li protein du trt trong clAu nanh, chdt irc chi5 proteinase ga.'-
ra do v6t thuong cua cd chua va khoai tay (Farmer vd Ryan 1992), protein dg trt
trong hot vir protein mdng th€ dAu (oleosins).

lvluc do jasrnonic acid nQi sinh cirng gia tin_s dap ung vrri nh&ng kich thich iu
b0n ngoai nhu bi thuong. luc co gicri. tuc nhdn gay ra <irl mam b€nh tdn cong va stress
cio tham thiu.
t'
3.9. Cdc chit di€u hda sinh tru6'ng khrlc
Ngoai tdm nhom chinh da dugc d0 c+p o trOn, ciic chit thuQc nhom
, ':' &c
polyamine, cac chdt
, ch€, sinh truong hay. lam chdm sinh trudng cua thuc vdt cing
li nhirng chit chi phdi quan trong ddn sr,r sinh truong va phdt tri0n. Nhtng chit niy
se ducrc d€ cdp o nhung chuong sau voi nhfrng tinh chit thuc ti6n cua no l€n sp sinh
truong va phirt tri€n cira thuc vit cDng nhu nhiing ung dgng cua ch{rng trong ndng
nghiep.

,',t.:., Ii I't't,,ir !' lt' :tu.i: .'t, f,*'iilii7il 60

 . Cluro'ng I
\AI
TRO CUA CHAT DIEU I-IOA SINH TRUONG
TRONG SINH TRUONG VA PHATTRIEN CUATHU'C VAT

4.l. Diiu klri6u su'rr:ir rrrinr cria hrlt rir sr.r'plrit triirr ciin ci1'con
Dc hieu 16 nhirng v0u to ditu hoa srr nav ntdrn vA quir trinh sinh tnrcrng sau
do cua civ con. truoc nhit clring La ciin lriiu rd nhinig qud trinh lien quan trong suot
giai doan nir1,. Nhlin giong bing hot lA phuong phdp clrinh cira qu6 tlinli sinh san
trong tu nhi0n vd la phrrong phiip duoc dp dung tren diQn rQng trong n6ng nghifp do
tinh hi€u qui cao cta n6. HQt la mQt noin dd chin, khi rung khoi cAy tne no chria
.:
ph6i vd' chdt dg trf dugc bao bec bdi vd h6t. S1r niy rnirn cria hQt c6 th€ dugc dinh
nghia nhu rnQl loat su ki6n xdy ra khi nhtng hot kh6 kh6ng hoat rlQng hirt nuoc, kOt
qud ld gia tang hoat dQng rrao OOi cirit vt\ khoi lao mdt cdy con tu phoi. D6 khoi diu
.:
sg ndy mam,, nhtng ti€u chudn sau c6n phii co:

vi c6 khd nang ndy mim).

- Truoc nhdt hot phii s6ng duoc (ph6i cdn s6ng
.':
- Nhfng di€u kiQn moi truong thfch hcrp nhu nudc ddy dn, nhiQt d0 thich
h-o. p. oxygen, r,a trong mQt vdi truong hqp Anh sdng phdi dugc cung c6p.
- Mi€n tr4ng chfnh phai duoc khic phuc.

Trong nhi6u rucrng hop, d6u hi0u cdn s6ng dAu ti€n cira h6t ndy rndm lir sq
nhri ra c[ra rd rnarn rir vo hQt. C6 nhung trudng hop ddc biet, ch6i li tinh hiQu cira sU
T ,]
song ddu ti€n, vi du nhu hdt Salsola. Theo sau su nhi ra cua r€ mdm cdy con mgc
nhu mQt sinh vdt duoi rnit d6t vin chua dqa viro quang hqp cho sU sinh trudng. Khi
cdy nhu len tu dat. quang hqp vd hoat dQng sinh dudng b6t diu.

:
Bon giai doan li€n quan d€n su niiy rnAm cfra hQt vd sg phrit trien cria c6y
con ld:
- Su hut nudc.
- SU tao thdnh ho{c ho4t hoa cdc enzyme.
- SW bi6n dudng cria nhfng ctrdt Ou trft, su vdn chuy6n ti6p theo va sr,r t6ng
't
hqp cua nhtrng ch6t rnoi.
- Su nhf ra cua rE mim vd su sinh trucrng cria cdy con.

C6 nhfrng chdt ric che vd kich thich sinh trudng nQi sinh inh huong truc
.i .:
ri€p den su ndy mAm cia h$t. MOi quan h€ cria nhirng chdt sinh trudng thuc vAi dsn
ie ua f6t hop ld dqa tren nhtng quan h0 cua n6ng dQ nQi sinh cira chring voi nhftng
giai doan phdt tritln d4c bigt, nhirng inh hudng cua vi€c rip dung ngoqi sinh va m6i

Cio Trinh Chrit Diu ltod Snh Tnmng Thuc I dt 6 |

quan cria nhfrng chat sinh truong thuc vdt voi hoat ddng bi6n du0ng. Nhtng chdt
hi
sinh trudng thuc vdt clng c6 li€n quan dtin qud trinh trudc khi ndy mim nhu sg huy
cl6ng chdt du trt, phdt tri6n rc, phat tridn uqc ha diep, kich thuoc tu di€p vir trqng
.' : lt
lugng cua no vd sinh t6ng hgp di6p luc t6 trong diQp'

4.1.1. Anh hrro'ng ctir gibberellin vi abscisic acid

- r iibhcre llin kich thich su nay rnatn'
- ,\lJA uc sh6 ndy nrArn vi tac d6ng nhu m6t ch6t O6i kh6ng tg nhi6n vsi
gibbcrellin.

Drroi nhung didu ki6n thich hop, nhtng bi6n di thi6u ABA dE niy mAm som
truoc khi chin. Nhlrng bi6n di thi6u ABA cho nhirng hot kh6ng mi€n trang vd niy
rniirn nhlnh chong: trai lai nhirng hdt thi6u CA sE khong ndy mirm n6u khong 6p
dlrng CA. ngoai sinh vir ti l€ nay mdm tang theo n0ng dQ GA xn l!'

Su nay rnim cira hot lu6n nhay cdm vsi ci chAt sinh truCmg thuc vat n6i
sinh va moi truimg. Anh hucmg uc chi cua ABA ngoiri sinh l6n hdt thi tucrng tV vA
cong tinh voi su giam th6 ndng nuoc. FIam luong ABA cao trong hQt kich thich
rinh nha-v cdm cua hot tl6n su giam th6 nang nuoc. do d6 lim giim kha nang ndy
tIatn.

4.1.2. Anh hrti'ng cria cYtokinin

Cytokinin co vai rru rjicu hoa qua trinh nal mim. N6u cung cip cytokinin
vi\o tir di€p hqit Cicer arietinum lttc 12 gio sau khi ngim ht2t thi th6y c6 str huy
dOng chrit d1r trG cin thitit cho qud trinh ndy mim. Ap dung auxin ngo4i sinh clng
cho thril cri lion quan ddn su nay rndm cua hot th6ng scotland mqc dir nhtng h6t
nil yclu ciu anh sang do dd nriy mam t6i da.

4.1.3. An h hu6ng ctia ethYlene

- Ethylene kich thich nay rnim cua hor. xu l! tthylenc hoqc nhirng chit
phong ttrich ctlry[-nc- nhu cthephon cri the kich thich nly mim trong nhiiu loiri' C6
srr eia tirn-q siin sinh cthylene trudc qud trinh nav miim. IIdm luong ethv-lene thodt
ra it hcrrr o nhirng hOt co uricn trang htrn lir nhirng hot kh6ng nti6n trang. Nhting
bicn cli (clr) ega Alultidttitsis tltalietru thi khrlne nha-".' ciirn voi cthllene r'i h6t cua
nhrlrns criv nt\r cti ti lc ntil rrri\rn thirp hun cllnq hoang tlai ,

- COr li chit irc chtl hoat d6ng Cua elhvlene, n6 lai kich tlrich s'.r ndy mdm.
:,
Khi rLr l. krlt htrp CO2 voi ethl'lene chirng cDne kich thich su tong nup ctnvlene.
Nhring yir.r t6 khac cirng kich rhich srr nriv mirrt nlitr str hcil .qia. ph1'tochrome.
,t_
a,t,n lii,fi (l .it f\i; lltxi SnhTrLring l'l.luc l'iir 6l
 nhifi dri. c1 tokinin. gibbcrellirr. tusicoccin (m6t d6c td tu niilr) r d nitratc. -l u1
'
..1.:.i.. ..t-..1-..^ I.t.'..-^
rl reil c(ilYicirc r:
iinoitg tllL-t] quati ocn
^ '' l vor c) tof. ntlu.
su ltav nlalr cua not rau orcp
).

{usicoccin har gibbcrellin.

4.1.{. Anh hu'o'ng cira nh['ng chit khic

.i
Nhtrr-s ehrit khilc nhrr potassiunr rritratc. t iourea. fusicoccin. coty lcnin.
lr

brassinolidc .'t"i srrigol cing kich thich su nar, mArn tu1,' n1.,16n vai lro cua cltirns. chrra
rd.
NH, .o t t:c.
\ .//
/ ".'-K ?,, x-_/
HzN----(
" cll:
\
-
lhiourea ll o't"'

:(''
H

o.-t"t
tt. t
llrrtlt , t
1.l. (a! ttu !ruc' ctru tltiourca vafusicoccin

Q Cotylenin A
R=

O Cotylenin C
n
'-
R-H Cotvlenol

HO-. gg Cotylenin E,

^: lrfic cua nhom colylsl1,

IIinh 4.2. Cau
Gfio Tr* Chdt Diiu Hoa Sinh Truong Thqtc l'dt OJ
 Sor-e,olactone
\o)oo
L{
Strigol
cHr

P-/o
,*,
ttA
\-rry
I

orl .

Alectrol
Yoro GR-24
-\ CH:

Dihvdrosoreoleone

tlinh 1.3. Cdu trtic aia cdc chat strigo!, sorgolactone, alectrol,
G R-2 4 vit d i hydros o rgo I e ane

- Potassiurn nitrate: Kich thich su nii_".' nrdm cuah$t, tuy nhi0n cdch tac ddn-e
cua no van cnuil ro.
- Thiourca: C6 khn nang t5i da hoa sr,r ni1' nrim va vrrot qua nhirng nritn
trrnq clo v6 hrit vii srr irc ch6 s nhi0t tl6 cao. Nguoi ta nghi ring nhtrn-q iinh hrrorrg
kich thich cua n(i cri thi do hoat dirng crtokinin cua no.
- l:Lrsicoccirr r,ii cotvlcrrirr: Cii hai chiit rrir d,.-iu co inh hrrons kich tlrich niv
rnarn. str kich thich duoc th[c dav khi ket hop voi G.A,r'a cy'tokinin.
- BIi kich thich str nav minr cria lrdt *itchu'erd vir loai trt Anh huong uc
chJ crja IAA r,a inh san.-e len srr na1'mam cia hdt nal .

t;;,i,) lt'itth ( ir;t l)t';t llL)i St,h 1r:arng l-hv< l ir 6l

 4.2. Sg thirnh lip Ainn ri.cinh giinr
16 Uiit I
Kha nang cua nhiiu loai ci' r.a nhic\u phdn cua cria cay cir
khai ndng tao 16
bing cdch gi;inr duoi nhirng di€u tiien rnicn hop la ydu to quan
trong rro'g
nhrin -eiong cua nhieu loai. Tha'. ri hoac ld co thd ding 'ic1c
larn ngu6n cho vicac
giong khi ducrc k€t hcrp voi hria chir thich hgp, 16c
nhin co hoc 'hdn
diiu ki0n
ntoi truong. Mot ircng nhrrrrg rtri diirn chi'h cua phuong 'a 'hii'g
phap rrhff'gi6ng ro rirrn
nay la cir nioi duoc lao thanh giolr_e nhu cit, cha me.

von Sachs (1881) crro rine co rn6r chdr hay nhfrng ch6t ndo
d6 rrong r6, nu
t'a tu di0p cia cdy da di crrulin dcin 16 vd kich rhich sg
rao 16. lrorr'one nal,rJi
duw Bouillenc va \Ven( (193j) goi la;/rizocaline vi cho ddn ngdy
nay,r,in Id m6t
chit mang tinh chit gid thu1,6t. Nhnng nh6rn kich rhfch sinh truong nrru
auxir.r.
cylokinin, gibbcrellin, ethylenc, brassinolidc clng nhu nhirne
chdt uc ch6 nhu
abscisic acid, nh&ng chdt hodn sinh truong, va phenolic ddu
c6 6nh huong lcn su
khoi sinh 16. Ngal' nay, auxin duoc bi6t ld nhom co dnh hudng
rcrn nhat rren sy too
rd vd da ducrc lhuong mai hoa. Co nhirng y6u to t;en quan
o6n s,, rao r€ r,a chat sinh
truon-e thuc vdl duoc li6t k6 sau:

l). Nhlng di€u ki€n moi rruong va dinh du6ng cria cdy cha me vd
(
cinh cfrt.
(2). 1-hrrc hi6n su clr canh sau khi di chuy6n tu cay cha me.
(3). Khoang rrong dir cruuc chon. phuong ph6p ap dung
vd chdt dac biCr
duoc dung nhu GA t hoic GAa. zeatin hodc isopenryladenine.
(a). Giai doan cua quil rrinh rao 16.
(5). Nhrng tusng rac gifra nhring chdt sinh rrudng thuc v6t da
biet va chua
bict.

- Auxin: Anh huong rdn rr€n su kfch thich ra 16 vi dfr duoc thuong
,i .
nrai h6a.
':
uac char rong hop nhu IBA va NAA dd cho hi€u qu6 kich rhfch rao 16 hcy'
ca tAA.
co nhi6u biio ciio cho rdng auxin licn quan d6n su khoitao.6 bar dinh va
sp phan
chia d6 cua nhtng khoi dau thi plru thuoc vao auxin ndi sinh
hay ngoai siqh. Auxin
clng uc eh6 tao 16 voi n6ng d6 cao. 'I'rong nhidu fuong trcrp, tiJ nhay
cdrn voi
cnat srnh truong thuc v'4t thi quan trong hon n6ng tl6. Nghi€n
cuu vdi m6 rhuc vat
cnu)'en genc cho tha), rang su truy€n plasmic Ri (root-inducing) vao
trong m6 thuc
vat dd gia tang tinh nhay cirn cua chtng voi auxin.

- Cytokinin: Ti r€ cao cia cytokinin/ auxin kfch thich su phat tridn ch6i
ric ch€ su ph6t tri€n r€. 'd
- Gibbereilin: Llc ch6 su thdnh lap rE (duoi nhfrng didu ki6n ddc
bi6r kfch
thich su tao re;.

@ 65
 - ABA vir nhiing chit ri'c ch6 sinh tru'&ng:
- ABA uc ch6 sg phdt tri€n 16.

Chdt &c chti sinh t6ng hop GA co dnh huong nhc l6n sg kfch rhich su rao
:,,{
rC bit dinh, giam GA n6i sinh vir giim sg phiit tri€n ctia choi.
- Ethylenc kich rhich su rhanh lip rE bdr dinh.
- BR uc che su thanh lap re.

4.3. Mi6n trgng

- ABA gay mi€n trang: Ham luong ABA trong hOr, 16 vd mdm chdi cao ddu
r: rj .
din d[n su rni6n trang. Tuy nhien cing co truong hop trong rlidu ki€n ngdy ngan
_!lav ra su' mi€n trang trong vai loai lai kh6ng co su gia tang ABA n6i sinh. Vi€c xri
li AIIA ngo4i sinh l0n rniirn chOi va l€n hdt da kich thich rni€n trang cia chirng.

- Gibbercllin: ldm mdt di mot so loqi mi€n trang bao gdm midn trang do
sinh ly, mi€n trang clo anh siing va mien trqng do nhidt.

- Cytokinin: r\nh huong cira cytokinin len rni€n tran-l khdng m" Cytokinin
ngoai sinh doi khi cO tn6 fnic phqc mi6n trang trong chdi va hQr, tuy nhi€n ddi khi
inh hurin-c ndy cing kh6ng co.

- Iithylenc co the khic phuc miCn trang o mot vai loai.

- Nhiirrg chit khdc:

r IAA/ ABA: Ti le IAn cao co khuynh hucrng ph6 vd nidn trang.
+ l'hiourca khic phuc vtii loai midn trqrng va iinh hudng cua no c6 li€n quan
': hoat dOng cira c1'tokinin.
dcrn
+ Coumarin giiy ra tinh nhay cim OOi vOi dnh sdng cho vdi rau ditip
-ei6ng
rnii hQt r:ira chfrng niy mirrn khong clin 6nh sing.

4.4. Anh huong cira clrit ttidu lrda sinh truri'ng l6n qui trinh lio ho{,
- Cytokinin: Cytokinin duuc bi0t li chat lam giam ldo hoa' no kich thich su
:,
plran chia t€ br\o vn vin chu)'6n cric chat dinh du&ng lim tirng khi nirng ho.lt d6ng
. 1,.
cuit rl'lo Va ic Dilo.

-;\urins: r\urin cti tlr[ linr tiLng. giiirn hoirc khting co tiic tlun,] lin srr_ltio
holi pliu thiroc vao cich xtr l_ rii nong do xt l(. Auxin gay ra srr lio htla do kich
thich sLL san sinh ethv'lene. Auxin n0i sinh giam trrrcrc hoic trong sudt qui trinh ldo
hoi.

t:',hli;t.h (l'. r Llr.;r. i,',,'r.S rrl' lr:nn,'4 Ttv-' | ,t 66

 - Gibberellin: GA larn giam srr pirin ri RNA vi protein. chfng giup gian su '-
lio hoa o cucin la va kdo dai su chin. Neuoi ra cilng th61 rine ham luong GA giim
-' i
lfuoc hoac trong su0t qua trinh lao hoa.
- Ethl'lene. ABA va methvl jasmonate: Cac ch5t nAy drroc bi€t nhu la nlitlc.
chit kich thich su luo hoa. Trong do ethl-rene chi ph6i quii trinh tio hoi ro nhit.
Arrh lrrrorrg. lri'u srr llro hoi do AtsA ra jasrrronatc s:ir pg,.l d[u co licn quan din su
sinh t0na. lrop cth-r lcnc.

* ung dung cia chdt diAu hoa sinh truorrg dd gidm lio hoa: Nhidu roai rau.
qua. trdi cdy va nim duoc giri tuoi d€ dua ra thi tnrcrng. Do do vi€c duy rri chir
Itrong trong su6t qu6 trinh lon trir sau thu hoach, r,an chu1,$1 d6 ginm hu hai la rAt
quan Lron-s,. Cl,tokinin, chit can sinh truone. auxin, gibberellin vi nhFng chdt fc c[6
hoat dong hoirc sinh tdng hcrp ethl lcne co kha niing giam llo hoa.

()rio Tiinh ('htir l)iiu lloi Snh Truvng Thuc I'tir 67

 Chrong 5
.
VAI TRO CUA CHAT DIEU HOA SINH TRUONG
lex cAc quA rnixH srNH sAN cUA THUC vAT

5.1. T16 hoa

) .:,
Su clruydn ti€p tu sg phdt tri€n sinh dudng sang sinh sdn ld mdt pha cuc
trong trong chu trinh sdng cua thr,rc vit bdc cao. Co ch6 cua cach chuy6n ti6p tir slr
.;
phdt rri6n sinh dudng sang sinh san van con chua duoc hi€u rO hoirn toin vd co the
t6m tit theo ba bucrc chfnh sau:

- Giai doqrn diu ti€n lii su tugng hoa, n6 ld mQt su thay ctOi sintr ly nQi tai
vi
hinh thdi. SU thay doi vd hinh thai hoc
rrorrg nur phan sirrh kir,r thco su thay tl,ri
,lC nhin thay truoc nhit ddnh diu sg chuy6n ti6p tir giai doan sinh dudng sang sinh
sin ld su gia tirng phdn chia t6 bdo trong vung trung tdm ngay phia duoi clinh sinh
truong cua md phin sinh. Nhfrng sg phin chia xdy ra o day ln k.it quai phAn hoa cua
i,.
nlrirng tc bio nhu rnd bro quanh m0 phin sinh thirnh khdi hoa so khoi.

- Giai <Joqn thir hai ld su thdrnh liip hoa. sg khdi ddu cia nhfng thanh phin
htra ct\ tlri nlrin thav dtrttc.

- (iiai doirn cu6i cirng ld sg phat tridn hoa. D6 lti s1r phin hoa ciu trric hoa tu
.j
su thanh llip 'hoir tlcln tro hoa va tun-t phin.

"l'rong rrhicu loiii thuc viit, su khtii cliu qud trinh sinh san duoc <Ji0u hoa boi

nhtng 1cu tt) mdi truong rrhu do dai rrgdy va nhi€t dd theo mQt chu k! thay ddi
hiing nirnr. Srr khoi cldu qua trinh sinh siin nt\y cdn bichipnOiUOichit cliiu hoasinh
trrr(rng hay do sr,i trrong tac giira yeu t0 rndi truong voi chit tlicu ht)a sinh truong
, i ;,,
cung vtii cric yclu td khirc.lfuy nhi0n su dhp irne nriy tuy thco loii.

5.1.1. Anh hrro'ng ci'ra nhilngyiiu tri m6i tm'dng lOn sr;'phrit tri6n sinh srin
5, l. l. L Qurrrg k! (photopcriotlisnr)
- Criy'trung gian: Str ttrtrng hoa cia nhlrng ciil'nfr1'phtr thudc vao ki0u di
:
trrrlin vri cril klrtrng tloi htii nlrtt cilu iinh sltng dltc bicrt.

- Ctil quang crirn: S1r tlti hoa ctia nhirns ciil'n;)y phu thudc vdo cl6 dai ngdy.
Nhirng cil,qrrang cirn co the tro hoa theo nhirrts di€u kidn satr:
+ Ciy ngi)'ngin (short-dr1'plant): Nhting ci1'ni1'chi trd hoa khi chu k.i'
16i clii hun cli clai toi hrn ni\o tt6. Cil' g-ioi hnn chu ki tOi Od tr5 hoa.

-iijt i.lt f ,t-". tt.;,:'t,t! /r':,' a.q /l:, r,i 6 j

 + Cdr ngiy diii (long-Ca1 plant): Nhftng cdy'nav chi tro hoa khi chu ki t6i
i,
ngan hcrn do dai toi h.rn nio do.
' + B€u canh cdy ngiy n-san r,a ciiy ngdy dai con c6 hai nh6rn phu khac.
Nhonr thu nhAt ld cdy ngd1, ngan-dai (short-long-da;'plalt) chi trd hoa khi trdi qua
.,: J
di8u ki€n ngdy ngin trucyc va theo sau boi dicu kien ngal' dii. Nhom thu hai ld cd1,
ngai' ddi-rrgiin (lone-short-da1' planr) chi trd hoa khi trii qua di€u ki6n ngAy drii
truoc va thco sau boi didu kiCn ngAr, ngin.

Do dai ngiy duoc cam nhdn tir li cdy do su hdp thu photon dnh sdng cua
ph1'lochromc. 'tinh hi6u cua sg cdm nhin anh sdng sinh ra tu ki sE dugc truydn d€n
nsi hoat dQng. Chit (ao tinh hidu tro hoa dusc goi ld florigcn, chit kich thich trd
hoa hoic hormone tro hoa. Chit tao tinh hi6u trd hoa di tluoc chung rninh bing
nhirng thi nghi6rn ghep ciy trong di0u ki€n cdrn irng vd khdng cirn ung. Khi tri cua
: trong dieu
^ trong ,,:
ki6n kich thich tro hoa nhu chu kj, iinh sang thich hqp hoac xir
cdy
lj lanh duoc ghcp vdo c6y trdng trong didu kign khong cdrn ung co thd kich thich su
tuong hoa. Nhu v6y c6 thd t6n t4i m6t chAt kich thich tr6 hoa tuong tr,r nhau trong
=, co nhfng truong hop kh6ng thu duoc su tr6 hoa khi ghep
todn cdy. Tuy nhien, van
tr€n mdt s6 loii ciy. Cho ddn na1, florigen v5n chua dugc phdn ldp, r,i vd1, c6 nhfrng
:. , : '
gid thul'6t cho ring su tr6 hoa khoi dau bdi su tuong tac gifia nhiing chdt diOu hoa
sinh truong thuc v{t kich thich hodc uc ch6 rro hoa di Ui6t va chua bi6t.

5.1.1.2. Su' thu hin (Vernalization)

:
CAy tr6ng trong virng 6n d<vi s€ tr6 hoa trong di€u kiOn xu lii nhi€t dQ th6p.
hi€n tugng. ndy ggi la sU thu hdn.

5.t.2. Srl tu'o'ng mim hoa

- C6y sE kh6ng tr6 hoa trong su5t thoi kj, c6y con.
- Khi cdy hodn tdt giai doan cdy con cua su phrit ridn. chfng co kha nang
sdn xudt rrrQt ch6t kfch thich trd hoa.
- Dua tr6n nhtng thi nghi€m ghep cdy c6 thd k6t lu4n ring chdt kich thich
:.
trd hoa thi tuong ty hoac rdt gi6ng nhau trong nhi€u trucrng hqp.
- Chir kich thfch trd hoa duqc dC xudl lin dAu ti€n do Chailakhyan (1936),
,.: :, :
6ng co nhieu c6 gang dd xdc dinh vd d[c tinh h6a v€ mdt h6a hgc cia n6. Florigen
ld mQt chat gi6 thi6t quydt dint.t qua trinh nriy. nhung d6n nay vAn con li di6u bi An.

t,.
Cho d€n ngdy nay' nguoi ta vdn cdn nghi rang chat kich thich trd hoa l2r don
chit hoac mot vdi chit ri€ng bi€t, hoac c6 sr,r tuong t6c gita nhtng ch6t diiu hoa
sinh trudng thuc vft c6 thd truc titip gdy ra trd hoa. hay nh&ng ch6t di6u hoa sinh
: ,.^ r,l
trudng ndy lien quan ddn vi€c gdy ra su sdn xuit nhtng chAt kich thich tr6 hoa.

GfioTri r Chrir Driu |loA Snh Tntvng Thrc I1)t 69

5.2. Anh hu&ng cfra ch6t tliiu hda sinh trufing l€n sry tuqng mim hoa, kich
thich vir ric ch6 tr6 hoa
C6c chAt di€u hoir sinh trudng cld bi€t c6 th€ g6y n€n su tuong mdm hoa,
kich thich va uc ch€ tr6 hoa trcn nhi€u loai thuc vat.

- Auxin kich thich sU thinh lap hoa o ho Bromeliaceae thong qua su kich
thfch san sinh ethylene. Auxin ngoqi sinh nhu IAA vd NAA uc ch6 su thdnh ldp
hoa khi drroc dp dung duoi nhfrng di€u ki€n cim ung. Chdt uc ch6 van chuy€n
auxin (1'lBA, t€n thuong mdi ld floraltone) kich thich sg tugng hoa d tdo.

- Khi ethylene hoac nh&n-e hop chdt phong thich ethylene nhu ethrel uc ch€
hoirc kirn hdm su tr6 hoa. E,thvlene ctns c6 kha nans kich thich trd hoa d vai loii
thgc r,4t nhu khom.

- Ci,tokinin duoc dp dung ngoqi sinh sd kich thich s$ thanh klp hoa. Khi xu
1."'' cytokinin len d<2t cffy drr<ii nhftng quang kj' khdng cim frng sd gdy ra sg phdn

chia t6 bab ssm han trong giai doan tuong hoa.

- GA ducrc ap dung ngoai sinh l€n cZiy duoi nhfrng di0u ki6n khdng cam fng
SE kich thich su trd hoa o mQt vai loiri ciy. CA cting lim cdy ki6ng trd hoa som va
t+p trung.

- AflA ducrc cho li chit uc

trd hoa sau khi duqc phat hiQn. Tuy nhi€n
ch,-i
nhirng nghidn ciru ': gdn dd1' cho thdy ring AIIA khong phdi It\ chit dong vai tro co f
nghil cluan trong trong srr diCu hoa qua trirrh rhanh liip hoa. ih,it rt thi vaitro cia
nri trCn sLr trd hoa vin chLra tluoc bi0t rc-1.

- Salicl,lic acid co klrh nirrrg kich thich tro hoa tren viii loiri.-[uy nhiOn co
': cna
chC nii tdc clone lCn su tr6 hoa viln clrrra rd.

O thoi diem hien tai tlri vai tro cria BR ra JA lin su trd hoa vin cr)n chua
t. r .l : , :
drrtrc lricrt rr]ri€rr. nh*n_e, nghidn cLru r'0 anh hrrtin_e cua hai chiit nuy l0n su tro lroa can
ticp trrc rlu<''c liirn r0.

5.3. {nh lrrlo'ng ciir chit diiu hol sinh truo'ng l0lr sr.r'phlit tri6n cua clrinr hoa
hoic thin trong nhirng ciy cti hol lr\ s11th0 hi0n gi6'i tinh
5.3.1. Gibberellin vi sr.r phit tri6n churu hoa hoi-rc thiin
:. dung GA l8n ciy cri
- Ap 1'cu ciiu lanh l:ohc cAl'ngi'r1'drii sE kich thich sLr
thr\rrlr lip hoe tlucii nhirnq cliiu kicn khonq cam frng. (iA cirng kich thlch trd hoa
' ' li! it.;-' ,\, tl,ll,,"';'.,"1,- 74,, r ,' 70
Lrong nhirng cdv rtgdvrgin duoc trdng tlortg dieu kiQn carn tirrg.
- Su thdnh ldp hoa trong nh*ng cdy co 10u ciiu lanh vi cd1'ngiy ddi di duoc
..: ,,.: ,:
di€u khi€n burrg cdch dieu hoa rnuc d6 Gr\ ndi sinh th6ng qua viric su dung ch.t uc
ch6 sinh tong hoo GA.

5.3.2. Chit tli6u hda sinh lru'ri'ng vi sg thG hif rr giri'i tinh
Nghidn c[ru ve girli tinh cua thuc vil dd duoc con n-erroi quarr tAtrr ti truoc
thoi cua Aristol.lc (384-322 truoc c6ng ngulin). Tuy nhien r,'icc Irghidn cuu anh
. . ,:. ..:
huong cua chit dic'u hoa sinh truimg lcn su tlr€ hiqn gioi tinlr cua thuc vdt chi nioi
bat ddu tu nam 1987 do Chailakh;an vd Khrianin khirixudng. C6 nhi€u cOng trinh
, .,: -: sinh r1 r:-.
-:-r- di
cho thdy' r'itr dp dung chdt di€u hod sinh truong ngogi larn thal' ddi gioi
tinh cua hoa,

Nhfng n-ehien cfu d6u tien tren hq bau bi dd cho thAl' ring vi6c xu ly auxin
ngoqi sinh ldrn ting s6 luong hoa cdi trdn nhirng cAy thuQc hg ndy, trong khi GA
ngoai sinh lai lim tang s6 luong hoa dUc. SU thay d6i gioi tinh gdy ra do auxin dugc
,:
cho rang co Ii0n quan d€n sg kich thfch sinh tdng hgp ethylene. Trong nhi€u truong
hgp .tt thay d6i rnuc d0 n6i sinh cua au.xin hodc GA cfrng co li6n quan d€n su ph6,n
hoa gioi tinh cua hoa. Hi6n na1,. nguoi ta chAp nhdn ring sU thd hi6n gioi tinh tr€n
ho bAu bi duqc cdn bang boi hoat dong cria auxin nQi sinh thdng qua ethylene va
gibberellin. l'uy nhi€n cing c6 f ki€n khdng ddng tinh hoiur todn vdi quan diern ndy.
.:.
Sy thay doi gioi tinh tuong tu cing dE dugc nghidn cuu tren cdy cAn xa Cannabis
saliva, thu hii duong, hoa bia. nho. dua huong, bi dao, bi rg, cd chua vd bdng vai.
Cytokinin ctng cho thdy kich thich sy tao hoa ciii trdn cdy cdn xa Cannabis saliva.
'l'ho c[ng cho thAy auxin ldrn tdng hoa duc
Nghi€n cuu cua Trudng Dai IJoc Cin
tr6n ch6m chom.

5.4. Su'rung
Sg rung duoc dinfi nghia ld sq 16ch roi crla mQt phin thuc vdt nhu lii, hoa,
trai. hirt. cdnh ho4c b0 phdn khilc cua cdy m9. Mac dn nguoi ta thuong nghi ring
sU rung thuong xdy ra d6i voi thuc r'6t bdc cao, tuy nhi6n o thuc vdt bdc th6p sr,r
rung vin Iay ra. M6t vi du thong thuong vd su rung lir sU t6ch roi cua nhlng td bdo
thuoc mo chuyon hoii ddc bi6t goi ld tAng rung. Trong nhtng truong hgp nay rnfit
ting phAn biet phai .on sdng rd c6 nhfng enzyme thuy phdn
khA ndng san sinh ra
kich rhich su rung. 0 ddy chn ."-6u de cap ddn su rung o thgc vdt b4c cao vivi str
rung la. c?rnh, hoa, trai va hot. Eoi voi lii co 3 d4ng rung lri nhr-r sau;

l. O nhlng cdy dE rung lii vA cdy buQi duoc trong trong vung 6n doi: Su
tA
rung lri li€n quan ddn nhftng y6u t6 mdi truong nhu quang k! ngan vd l4nh.

Cdo Trinh ('har DEvlkiSinhlnroryThvcIat 7|

 2 Nhtng cny dE rung ld vd cay buoi trong vuhg nhi€t tloi vd 6 nhi€t ddi: SU
rung la phu thu6c vao nhtng thay d6i trong sinh trudng vd cuong lgc cua cay.
3 Nhing cdy thuong xanh trong trong vung on cloivd 6 nhi€t doi: SF rqng
- : .: '
l{i thLroc r € su phat tri0n cira rnua trudc xiy ra 'rrong khi c'j sU mdi khoi ddu cua su
sinh tru'ong trong mia xudn.
- Sp run-e cdnh c0ng co thd xem nhu lir-mot thoi quen tg tia canh.
- Su rung hoa, tr6i vi hdt thuong xiry ra o thgc vat bac cao tac su thudn loi
cho quir trinh sinh san.

5.4.1. Giai phiu hgc cria sg'rgng

Su rung cua la, cdnh, hoa triii va hQt trong phdn ldn loiri cdy d€n truoc bdng
su thrinh lip v'ung hoqc tAng roi tu nhtng t6 bio dAt bi€t. Vi tri tAng roi s lti duoc
nro tti trorrg hinh 5.1. Nhu viiy mu6n han ch6 su ryng cdn phiri han ctre su thdnh lqp
:
tang rol.
..

Soim4ch
CtrOi bCn

86 mach
f:
l ang rol

Hinh 5.l. Ti bao fing rd'i o td

5.4.2. Sinh lj'ctia sy rgng

5.4.2.1. Anh hu'6ng cria nhiQt dQ, oxygen vi nh*ng ytiu tii dinh rlu0ng
Nhi6t rJd chi phdi quan trgng dtin sq rung. Khi nhiQt dd ting cao se ldm gia
tdng su rung. Co mdt nhi0t dd tdi hao cho sr,r 4rng ttli da. SU dap irng vdi nhi€t dQ
cho s1r rung cin,e khdc nhau. O dqu cove nhi6t d6 nay la 250C, trong khi do o bdng
vrii la 300C.

l1i\rr luong ox)- cao cirng lirn gia tirng su runs. Viin d,.i nlir tltrsc gia thieit
:
rring str gia tang oxy s€ kich hoat enz)'mc lAr\ oridase din ddn sU oxY hoa IAA I'ti
lirn giarn IAA ndi sinh trons nro. Gia tnitlt khac cho ring oxy kich thich su chur'€n
hoi ACC thanh ethylene vt\ kich thich srr rung.

(..rit irinlt t-lt,'i !'t,\t llo.l S t Trtroni Thcc l.i! 12'

 Ilartt luorrg carboltrdrate cao se gianr su lunq vd ll-e,rroc lai. Ducri 6iCrr kidn
fnh sine cao s€ kich thich quane hop d0 sinh tong ho-p nhi€u carbohydrate yd liint
cltirn su rung. Ciic qud trinh din d6n su tich [uv= carbohr.'drate thap sE din cl€p su
ru l1.'q.

Cal,dtro-c cutrq ciip rJarn diiv du cung du,r'tri lri lilu hcrn r,a cho rrhieu tnii
hon ca1' thicu danr. Nhnng cal, na), ckrc bi6t co hdnr luong amino acid va nhtng
hcrp chdt chua daur cao ho-n la 1'€u 16 -qir.rp cho su sinh rong hop l)NA, IiNA.
protein vd nh*ng thdnh plrin khac co khd nrng ngdn sg rung. cdy c6 chua clam cao
cing chua nhiiu auxin vd cytokinin.

ciic nguydn td khoring cing rdt cin cho su phrit tri€n crla cay. T'hi0u k€rn
: ,
hav tliicu calciunt
.:
dCu lam la d€ rune.

5.4.2.2. Anh hu6'ng cria chit di6u hda sinh tru'd'ng l0n su rung
- Ethylene kich thich su rung: Ethylene kich thich su,rpng di cluoc bitl rri
l6u' n6 duoc xenr nhu tinh hidu khoi d6u cua quri trinh rung trong cdy sorrg tti di€p.
- Auxin: Auxin c6 thd han ch€ su rung ho4c kich thich sU ryng. Him luong
auxin n6i sinh trong la hoic nhlng co quan khric gidm sd kich thich sr,r rung. Xu ly
auxin ngoai sinh xa tAng rung (v6 phia hi cua ring rung) ldrn giam su lfro h6a. 6.ddu
::.,:
e;in tiins rqng (v€ phia thAn cua ting 4rng) kich thfch su ldo h6a.
ABA ttuoc bi6t ld chdt ldm _eia rdng su nrng.
-
- cA c6 tlrci la*r gia rdn-e su rung niu hdm luong du lon dd kich thich sdn
xuit ethylene. Hi6n tuong gdy ra su rung bdi GA clugc xem lir tilc tl6ng cta ethylene,
GA khdng tu gdy ra sF rpng. Mar khric CA cing giim sp 4rng do gia tlng hoat dong
cua cdc co quan tich liy.
- Cytokinin c6 tirc dune kich thich sinh truong vd gidm su rung. XLr l;i
': tang rung sd kich thich su rung. Xu ly
cltokrnin s vi trlxa cytokinin truc ti6p hoac
o tren ting rung sd uc ch6 sp rung.
- Jasrnonate duoc bi0t la ch6r kich thich sr-r rung do kich thich qud trinh lio
hoa.
- Nhirng chdt kich thich rung 16: C6c ch6t kich thich rung la hic:.) nay ctucrc
t.:. . .i , I
bi€t ph6 bi€n nhu calcium cyanarnid, thidiazuron, arnmonium nitrate, endolhall.
paraquat, sodium cacodylate, sodium chlorate, tributyl phosphorotrithiLrate.
erhephon vd tiridiazuron.

Cdclt t6t nh6t d0 giim ryng Id:

j
l. Cung cap dinh
'. ' dudng
tdi hio cho cay ddc biOt ld dam.
2.'fritnh stress vd nuoc hodc nhtng 1,du t6 dAn drjn san xu6t ethylene holc
Gru Trinh Clxit Dftu lloo Snh Tnrvng Thrc lrit j1
 AI]A.
3. Tranh tich ltiy ethylene va gif t6c clo h6 h6p thrip khi vin chuy.€n hodc du
tni.
4. Khi sg sdn xuAt ethylene l<h6ng thr3 tr6.nh duoc thi ding rndy loc ethylene
, :. . , t
hoic nhlng chat uc ch€ hoat ddng hay sinh t6ng hqp ethylene cing cti hieu qud.

5,5. Sinh lj cria su {liu trfi, sinh truri'ng, phit tri6n, chin vi rgng trdi
5.5.1. Sinh lf ciia su il$u trrii
Su'tiau trdi c6 th€ duoc dinh nghla nhu ld sg phdt tridn nhanh cua nodn mir
thuong theo sau bdng su thu phdn vd thu tinh. sg phat tri€n rriii xdy ra khong qua
thu phdn hoac thu tinh vdi trrii khdng h6t l:i trinh qud sinh. Co thd phin bi6r
-eoi
trinh qua sinh thanh trinh qud sinh sinh du0ng va trinh qu:i sinh dLroc kich thich.

- Trinh qui sinh sinh duong (vegetative parthenocarpy): Su phiit tri€n trdi
. xa)' ra khong qua thu phdn (kh6m, cam navel rvashington).
Trinh qui sinh duo.c kich thich: Yeu cdu c6 sU kich thich cria hat phdn md
1-

khon-s co su thu tinh d6n sau d6 tao ra su dau trdi.

Circ chat di€u hoir sinh truirng nhu auxin, gibberellin, cytokinin. ABA va
ethvlcnc i0u chi ph6i qui trinh nay. Ciic loai rriii nho kh6ng hdt o Nhrit. Uc. Nl.v va
chiu Au thuong cri xu ly GA3.

- Anh huting cua auxin va gibberellin l€n su ddu trii:

+ I'J.AA tang ddu trdi tr€n qui da nodn (diu tay. bi, va. ca chua. hoa hOn_e.
, i . ,;.
illUOC la. Cii DnOr. . . ).
+ G.,\ chng co khd ning ting ddu trdi tr0n nhrlng ciy dirp ung voi auxin va
nhing cily kiuic mt) auxin khOng hiOu qua (vier qudt, cam quit. nho, qua co htit).

- Anh hLrting cua cytokinin. ABA va ethylene lcn su diu triii:

:
+ C,r tokinin tdng hgp: f irng ddu trrii tr€n nho. vdi...
+ AtlA vii ethvlene: Giy ra su rung hoa v;i Inii non.

5.5.2. Anh huting ctia chit tli0u hr)a sinlt trrrd'ng l6n sinh tru'o'ng vr\ phirt tri6n
cria hdt vii trii
- .'lu-uin lri .ra'.rialt lrttd'ng lrrii; Aurin co vai tro lirtn cluan -uiria str phat
tlitin Irdt. kich thtro'c vi lrirrh tllnq ciroi ctrrtg ctra trdi. Ap dung auxin vlio qiai doan
,:
bit kj' nio cua su phit tri€n triii dcru co str dap irng. Ndi nhi r,a phoi trong hdt san
,

sihh ra auxir. n6 sE di chu1,€n ra plria nsoiri vo vi kich thich su phit triin.

ill;o li'inh ('/r.ir llr.'ir llaltSr:i Inrnglhrc I;ir 7l

 - G.'l rd s4rphrit triAn trrii: Ilot lei ngudn giiu CA. Vi vav C,\ duoc xcnr lu
chtil g6p phan phdt triin trtii cirng I'oi aurin.,.Xu l, GAneoai sinh gay'r'a Lrinh cpa
sinh lii y6Lr tO khinq dinlr vai tro crla GA l0n str phiir triin ttiii. GA con iinlr hrrorr-s.
lin dd lorr ur'liinlt dang ctia triii. Bing cdch giarn llo hoa. GA giu cho vo triii carit
qtrit ttroi liu lrcrn cltinr rnr\rn l:hi chin va kco dai tlrtri gian bno quan hon. CA cilrre
l;iur cho r o tiio dcp lrorr.

- Cylokirritt risy phrit tri0n trdi: frdi non phdt trien rit nhanh vii cii srr
i..
pltrin cltiu ti lrao rnqnlt ntE. IIot cua ch[rng chira nhidu cl,tokinin va drrryc xent lli I'clu
to quan trong tron-a. srr phiit tli6n triii. Vidc xri I! cytokinin ngoai sinh cing ldnt ting
kich thuric (rdi, tuy nliiCir anh hrro'ng niir t,hay doi tuy thco loaica1,.

Nhin chung c),lokinin anh hirong nhi€u trCn su diu triii, trong khi GA anh
huong nhieu trdn str phit triCn trai l'a auxin dnh hucrng ca len su dau truii rai phat
tri6n trai. Mot san pham cti tdn la Promalin = BA + GA4 + GA7 li san phinr thuong
rndi cua Abbott Laboratorics dugc durrg tren t6o ae A;eu khi€n dang triii. cu triii,
trong luong va gia ting ning suat triii tao.

5.5.3. Tia thua hoa vi trii blng h6a chit

Viec tia thua giup loai nhung v6n dii co ilr6 xAy ra doi voi cdy irn tnii da
rricln rrhu su tr6 hoa cach narn. nhung tirn thuong v€ vdt lv doi voi cdy vir cirrg cr-r
, ,l .i
anh huong tot dCn kich thuoc trai, dang lrai. rnau sac trai va chat lucrng vucn troi.
Cd thd su dung ch6r didu hod sinh truong Ad Oiiu hoi qud trinh niy. tu1, nhicn n6ng
. ', rit: quan trong: Doi voi NAA hoac NAAm n€n ap dung sau khi
do vii thoi lian xu ly
hoa tr6 d€u trdn tdo vi 5-7 ngdy sau khi tring hoa rung tr€n 16. Vidc xu l-'- clic chit
khac nhu accel (cytol<inin + eibbcrellin), ethrel, sevin, vydate c0ng co hicu qui
nhung con phu thuQc vdo thoi ti6t, tinh tr4ng cd,1', gi6ng vd nh0ng y6u tO fnac.

5.5.4. Su'chin cria tr:li

Ethylene li€n quan ddn su chin dnhi€u lo4i cdy 6n trdi. Th4t ra thuit ng[
"climacteric" (gid min dUc) d0 drn chi nh&ng triii se chfn trong sg d6p Lrng voi
cthylene nhu ci chua va chuoi. Con thuiit ngf "nonclimacteric" (chua.gia. chu;l nrdn
duc) drn chi nhtng trai s€ khdng ddp ung voi ethylene d€ chin nhu nho i'a diiu tliy.
Co rit nhidu thong tin a6 tnang ciinh vi vai tro cria ethylene trong qu6 trinrr clrin.
. .;
l:thephon chdt ph6ng thich cthylene latn ting dQ chin vd chin ddng ddu. Co thd lijrn
giArn su chin d.-i keo dAi tuoi tho cua trdi bing caclr ngan cdn su sinh ti,ng hop
ethl,lene. gidrn hoat ddng cria ethylene hoac lAy ethylene ra kh6i virng gin quA. C6
,:. ,:
th€ uc che qud trinh chin blng cach uc che ACC synthase voi antisense I{NA. C6
thd mt ra v6 vai trd cLia ethylene trong qud trinh chin nhu sau:

@-75
 -
Filhylene didu hoa su chin. -u-cu ciu su chrry6n thdng tin li€n tuc cia
nhlng gene cin thi6t.
-t [:thylene duoc dieu hoa mot cdch tu xuc tiic.
+ Ethylene ho4t ddng nhu ld m6t bi6n hd hon ld mdt c6i ng6t di€n aC Aieu
khrCn quir trinh chin.
Ethylcne ld chia khoa tliiiu hoa o muc dd phdn tu cho qud trinh chin va
+.
ldo hoa. kh6rrg phiri la san phdm phr,r cira su chin.

- sH rung tral
).f,.5. Ngan
.';.
1'rai thuong bi rUng trusc khi chin, ddy ld t6n thtit lon trong sdn xuit ndng
"1
nghi€p. Nguoi san xudt d6ikhi phii thu ho4ch sdm d€ h4n chd su t6n thuong tnii do
rung, tuy nhi€n ddy kh6ng phAi Ia bi6n ph6p tdi hio. Vi€c rip dung ch6t diiu hod
, . ..: ..
sinh truong co the ldm giam sU rUng trdi. Cric sin phdm thuong mdi goc auxin di
som dugc 6p dpng d€ ldm giim su rqng triii. NAA dugc phun I€n lii hai tudn truoc
khi thu hoach giup giam dugc sU rqng tr6i tao. Daminozide (Alar) cfrng duoc phun
truoc khi thu ho4ch tdo giAm dugc rpng trai (hi€n nay kh6ng dung nira do dnh
huong d6n srrc khoe).

5.5.6. Gny n sq rgng trii

Viec ngin str rung trdi ri tr€n d6i khi lam cho trdi bdm chAt tren ciy vd gdy
kho khin khi thu hoach. Vi€c rung cdy co th€ l?im thdt tho6t ning su6t vi co nhtng
triii chua chfn b! rung. Vi€c 6p dgng nhtng chdt ph6ng thich ethylene dd kich thich
su rung trai cherry'. tdo, mdm x6i, dua l€, 6c ch6 vi quyt. OC trantr inh huong khdc
cua cthylene l€n su rung la. rung la vd tao mrl lirm rniit ning sudt d mua sau c6n d4c
|,,r:L.
biQt chri, d€n ndng tld vi thdigian xu l;i. ViQc xu lli 500 vd 1000 ppm ethephon c6
hieu qui rdt ro Ae larn cho trii cherry d6 rung vrii dnh huong d6c t6i thi6u va chdp
nhin thuong rnii. Ndng ctqi cthcphon cao clin 2000 va 4000 pprrr dd gd1'
clucrc trong
hiCrr tuurrg rung lu vu tao mu. O nhring ndng d0 niiy kh6ng rhri chdp nhdn dus.c vi
. ,I ,-
khirne dLroc chlip nhirn trong thuong mii vir lai gdy inh huong xAu cho cay.

{ltio Tritfi ('hrir t)<:'r lki S 'rit 1'.y':rg T!:i:: I it lo

 , Cltro,ng 6
\lAI TRO CUT\ CHAT DIEU }IOA SINH TRUONG
r-Ex qUA TRir.,NIi QUANG I-IQP CUA I'HU'C VAT

6.1. Chit c.in sinh trtlorrg

Chit diiu hoa sinh truong rlrrrc vat l.tao gorn cac chlt kich tltich ciic qua
trirrh silrlr trrrung rii phlit triin cua thrrtj rdt r,a ca clic clrat co hout dorrg trc clrc. Cac
chir uc che co rh€ chi ph6i qua trinh sinh t6ng hqp cira cdc ch6t diiu hoir sinli
rruongnQisinh khdc nliucdc chdt uc clr6 sr,rsinht6nghop gibberetlin. Cic chit rriv
lanr clro qui trinh sinh tong hrrn gibbercllin b! rru rrgqi va ket qud la thuirrrg lirlr cho
ca1 lun lai. ll0n canh clo ctrng co nhring chil cotic dr,rng cart tro trgc li6p qua trinh
.:
',' tliCn
sinh truong va phrit ma kh6ng irc che su sinh tong hcrp gibbcrcllin'

6.1.1. Nh['ng ch6t ri'c ch6 sinh t6ng hq'p gibbcrellin

6.1.1.1. Nhu'ng hq'p chdt oniunt
Cac chdt uc ch€ r6ng hop gibbcrcllin cO th€ k6 d0n lir cblorrnequate chloride
(c1,coccl. thuong mcpiquate chloride (thuong ducrc su dung),,
goi ln ccc).
AMO-1618. phosphon D. piperidium brornide (hinh 6.1). cac hop ch6rt nhom oniutn
.:
s€ uc ch€ qud trinh tao vdng cua gcranylgeranyl p.vrophosphate thdnh copallyl
pl,rophosphate va ngin uin qud trinh tao thinh ciic hop chdt gibbercllirr. CAy duoc
xri ly v6i nhtng hqp chrir onium se cri l6ng ngin. ld xanh hcvn vd diy hon. CAc hop
cSit nhour oniunr con rhfc div quang hrrp thuAn va tAng tinh chiu han ctta thuc vat.

Cllr
CI-CH,. \'--cllr
d,

cH3
Q
cH( '.",
Chlormequatc chloridc MePiquat

Hinh 6. t . Nhintg hcrp c'hat dqi di€n nlnnt ortiutn

6.1.1.2. Pyrintidine
FIai chdt ciin sinh trudng p1'rirnidine thuong dirng la Ancymidol vA

Irlurprimidol (hinh 6.2). Co chii tac dung co birn cua nh6rn nAy lA:
- U'c ch6 c),tochrorne P-450. 1,i5u tO aiiu khidn sW oxi hoa crla kaurene thanh
kaurenoic acid.
- U'c ch6 sinh tdng hqp GA.
- Gdy trcr ngai d6n qu6 trinh sinh t6ng hop sterol vd ABA.

6aoTrirtril,it o'iu lloit s|.,h TiuungThvc l'Qt 77

 Cri.c hqp chdt nh6m pyrirnidine it dnh huong I€n quang hgp, nhung chung
larn giam khd nang sir dung nuoc cria cdy.

Ni

// -\?' //
N OH F

_/rl\
I
(/ FC- o-cHj (,/\ O_C-F
J c4 t-\
C

N N F
I-cH., n
Ancyrnidol F lun imidol

Hinh 6.2. Nhintg hop chfu dai cli€n nhdm pyrimidine

6.1.1.3. Triazole

(>
\//
N
/\
\tl
N

N-N r----\ N-N

//\\l
.,1,/ \ |
cl-<\/l 'FCH=c
\:' iFCH:-cH
H
'-/ 'l H-C-Oll
-C-OHI L|rl' -C-.t,
-, I

cHi!.-cu,
'l
CH.r

cilr
I)aclobutrazol Uniconazolc

* /"\
('> ll
\N-N
ll
\N-N r\
// \\ |
| ( ')-o-Cl{.-CH.-Cll
-oi=i \:/ rr-t_oH
H-C-OI{ |
i cgJ-f -cHl
crr:-i)or,
, L 11;
crr
Tripcnthenol UAS lll

("'>
\tl
N-N
n,c-t'
-\
/--\
F.-l
/H
\-/ lt_c_ol1
I

,,\-ctt,
a\,
tl
o-....-ao
LAII t50973

[{inh 6.3. Nlui'ng,lup chit dai dien n]nnr Iria:ole

6nia l.rinh (-/r.ir ,,_rr.ir /lni .\ar, litt, g.l hrc lit 73
 l'riazole cfrng lii chit uc clrd !nh t6ng hop gibberellin. chung li nhom cirr
sinlr tru<rng co hoat linh cao nhu paclobutlazol. unicorrazole. tripcnthenol. BAS Ii 1.
LAB 150978 (hinh 6.3).Iloat d6ns uc chi sinh tdng lion cthllene va tic dunq sillr
lli ctia no l€n tlruc r,{t co th6 ducr.c ronr rit nhu sau:
- Hop ch6t triaz-ole lim -eiirn su sinh truonq cua c61, bang cach uc chd su
oxv hoa nricrosonre cria kaurcne. kaurr-'nol va kaurcnal. Nhtlrg chit niiy,tl[rr'yc xirc
tric boi kaurcnc oxidasc lt\ cnz-1 rnc cvtocllrontc P-4:0 oxidascr.
^: ngin catt su sinh tonq hop gibbcrellin, rrhing lrcm chrit rriazolc ciyr irc
- Dc
che sinh rong hqp steroll lirn giirn AI3A. ethylene va IAA nhung lai rang harn
luong cytokinin.
- Triazole co ldrn tang h.im luong chlorophyll nhung lai it rinh huong truc
',: d€n quang hnp. Tuy nhidn, triazole lai co nhfrng dnh huong giirn ti0p den quang
tiep
.:
hqp.
- Triazole co fh6 giLrp cdy ch6ng lqi stress 961, ra do nuoc va sullur dioxiclc.
,l,, ,:,
Anh huong na1'co the do triazole ldm gia tang ham luong hoic hoat d6ng cia ch6t
ch6ng oxi h6a trong ciy.
- friazoic co thd dnh huong lcn su giam mar cio quin thc con trung. Tuy
nhi0n. bin chAt crla hi€n tuong niv chua duoc nghi€n ctu rd ning.

6.1.1.4. Nhirng ch6t t<trnc

N N
xl
tl
N
I

<'\'
lrl
\.-
I
/-l
Tctcvc lac is

o
o
/^ U
ll
--1
/\tl
o-c 1r /}-c-ct2
\_____/

Prohexadione calcium

IIinh 6.4. Cau truc ct)a tetcyclacis, prohexudione vd inabenfide

Gio Trinh Chir !rcu lloi Snh Tiuong [huc I it ]g
 Bcn canh cdc chdt cin sinh truong duoc phan nhom nhu tr€n con co ciic
.: ..,
chat khdc nhu tctcyclasis, prohexadione-Ca vi inabenfide. Cdu tn-rc cira chung dr.rgc
mo ta trone hinh 6.4.

-
fctcyclasis: Tetcyclasis la ddn xu6t cia norbornenocliazentinc.'letcyclasis
lirr :,
giarn sinh tong hop GA, uc ch€ sinh rdng hqp sterol. Noi clrung tric dong cua no
tu()ng tu rrhu nhrirn triirzole.
- Prohexaclione-Ca: l.Jc cht su hyroxl,hoa 3p cfa G2s thanh GA1 vi su
hl,roxy hiiii 2B cua GA1 thanh GAs.
- lnabentidc; t-a din xudt anilide cua isonicotinic acid. lnabenfide ue che su
sintriOng hop gibbercllin bang cdch ngan can sr,r chuydn dOi oxy hoa kaurene thinh
krLrrrenoid acid.

6.1.2. Nhlrng chit cin sinh tru'6'ng khdng i"c ch6 sinh t6ng ho'p gibberellin
Nhung hop chit can sinh truong khong uc ch€ sinh tdng hop Cn co the kd
cl0n lti morphactin. clikegulac, cthephon. maleic hydrazilc, din xudt cria acetamide
va clin xu,it cur fhtt.v acid.

6.1.2.1. Morphactin
[.rr
rn<'rl. nlrorn chat can sinh truong bao g,)m fiourcncc.
llourence-9-carboxylic acid r,,a chlorflurenol methyl (hinh 6.5). Nh6m niiy anh
hLrung lin hinh thc cay. Morphactin uc ch6 sg sinh truong cua cay trong khi

-r-libbercllin kich thich su sinh truong cua cay'. Morphactin khring uc ch6 sinh tOng
htrp gibbercllin nhun-e hoat d6ng nhu rndt ctrir AOi khang. Mirc dir morphactin co
anh hLrone sinh ly r6ng trdn thrrc vdt. tuv nhi€n ciich tac ddng cira n6 khdng dic
hieu.

coor'r

i-=
\.4
F luorene Fluorene-9-cl rbory lic ac id

-o-cH1

Clrlorl'lurenol nrethr I

,'] cltdt ctit ttiton trtorp!toctin

Hinh 6.5. rVliiirie lwp
O .io li- rh (-lit Di;n I lo't lnh l'rnrng fbrc liit
6.1.2.2. Dikegulac +
Ciu
^,i
truc cria dikcgulac dugc nro ta trong hinh 6.6. ChAt ni1' lrittt giant ttu
the choi ngon va kich thich choi b€n. Nhtng t€ bao dang phdn chia thi rthav canr voi
dikegulac. triii lai nhLing tC bdo di on tlirrh thi it bi anh hudrrg. Co ch6 t:ic dung cua
dikcgulac r,an chua duoc hi6u 16- no lanr giaur nhlng chdt gi6ng nhu GA. kich thich
nhiine chit gions nhu AllA vi cthvlcnc.

Dikegulac

Hinh 6.6. Cau tritc cia dikegttlac

6.1.2.3. Hg'p chit ph6ng thich ethylene

o
ll--oH
ct-cH2-cHr-\o"

Ethephon

Hinh 6.7. Cdu tuc cua etlrcphon

Ho-p chir ph6ng thich eth)'lene dien hinh ii ethephon hay ethrel (hinh 6.7)
dugc xem-nhu ld ch6t cin sinh tru&ng. r,i ethylene lirm cho than ngin hon vi ddy
hon. Erhephon hay crhrel hi€n duoc dung khd ph6 bi6n. U'ng dung tliu ti€n crja
ethephon la duo.c dilng di3 di€u khi6n d5 ngatr0n ngu c6c vd cdy liry hQt. Ngdy nay
no duoc ung dgng v2ro ndng nghi€p voi nhi€u mUc dich nhu ldrn chin trdi voi rniru
vdng dep hay lim tAng sU tfch lu! duong trdn mia.

6.1.2.4. Maleic hyd razide

Maleic hy{razide (hinh 6.8) ld ch6r cdn sinh truong bing crich ngdn cdn su
, '{
phdn chia t€ bdo. Maleic hydrazide chen vdo su sdn xu6t uracil.
rif
t-l
N-N

' Maleic hydrazile

Llinh 6.8. Cau trilc cua maleic hydrazile

Cr)'o Trinh (-htir Dtiu llai Sinh Tirong Thqc L it 8l

 6.1.2.5. Din xuAt cfia acetamide
,: xudt : cia acetamide '
Cac dan di€n hinh mefluidide, amidochlor, lir
darninozide, cirrelacard (hinh 6.9). Cac hop ch5t nay dusc bi6t ld chdt uc ch(i sU
sirrh trut-rnu cua nhfns. bai co.

o oo
tl
NII-C
-CFl cr-cH/-'. N
--cH2-NI l-ctcHl
cilj
o o.i_
Fl'-cH, cH2-cHl
tl
NH-S-CFr
Mefluidide II Amidochlor
cHl o

oltll/ !rr,I OH

. or r -c-cr-h-c.Ft?-c-Ntt-*\..., c2Hjo-c a\
Daminozidc
clll
l)
Hinlt 6.9. Cau lrtic ctln nnleic hvdrazile

6.1.2.6. Din xuit ciia acid b6o

Cac din xuit cua acid bdo bao gdm cac ciic htlp chdt alcol cia acid beo voi
rnach carbon t* 8-10 va circ methyl cster voi mach carbon tir 8-12 co tdc dung ldm
:-
giirn chieu cao cai1, nhung co ch0 tac d(.)ng chua duoc biet. Sin phim thuong rnai
rncth,vl ester cila acid bco di tluoc cong b0 la co tirc dtrng gia tdng su phan cirnh vd
.: '
iirn giam su phit tri€n chrii.

6.2. U ng dung efra chit cin sinh tru'ri'ng

Cac chat ciin sinh trrrcrng rit nhieu
trrrcrns co rat nhieu [mg dung trong sin xuit ndng san, siin
j. i r'i
suiit hoa ki0ng. giiim dd ngi, liinr thudc diQt co... Cac irng dung thuc tidn cria no c6
thC tonr tit nhu sau:

l .i
- r-Ltrcu ,khrcn
' kich thrrric. hinh dane va chilt luong cul hoa kiing co th8 dirng
CIC-C. clarrinozidc. irncvrnidol. paclobutrazol. uniconazol. tetcyclasis. Trong do
. '' ' :
piiclobLrtraz.ol cil tiic dung nranh va cki btn cuo do do chi can phun xit ruot lirn.
- l.lot trong nlrirng [rng dLrne chinh ciir chit crin sinh trucing rrong nong
rrghiep li nqan d6 ngti trin ngri coc \'a lroa rnau 'iltiv hdt nlrtr l[ra nri. lua. lua ntach
I . !

' ri ki0
dcn '':' r nrach. Ngin dd ngi cfing co the dung paclobutraz-ol, prohrxadione-Ca,
CCC ra ctlrephon.
i\lalc-ic hi'dlazide rlrroc dirns dc irc citi srr nav nrinr cLia cLr hinh vi khoai

'iirJ.t liinh (-ii,it r)].;it I lL:^i \t,th fintDF llrtrc l,ji sl

 {dr'.
- Daminozide duo-c dung lant giarn sinh lruon-s dinh duOng va kich rhich trd
hoa (1989 bic6n o Ml')
- Paclobutrazol duoc ddn,e k! su dung o nhiiu nuoc tru Mi.

6.3. If6i ti6n quan gifta chit sinh tru'<ing ciil,trdng trong quA lrinlr quang hop
ra su phin chia ctia ch6t ttong hoa
* Gibbercllitt td qutng hgp
- Ap dung rrcln lij: Anh huong cua GA lr0n quang htlp khong rd r,d phu
thu6c vdo lodi cay. c6 nhidu ket qud cho th6y c6 su gia rdng t6c do quang hop horc
giirn quang hop.
- Ap dpng l0n r0: Anh huong khdng rd.

* Cytokiniu vd quang hqp

- Ap dqng ldn ki: Kich rhich t6c dQ quang ho-p.
- Ap dung len r€: Ting t6c d6 quang hop.

* Diiu khiin o mrt'c phdn tt tuqng cytokinin nbi sinh vti quang hgp
Cytokinin c6 khi nang lirm rine quang hqrp- Cav thu6c 16 chuydn gene voi
T-DnNA mang gene 4 tld rong hop c.vrokinin c6 chua 10-20% nhidu hon zeatin,
zeatin riboside, va isopentenyladenosinc so viri cay doi chung. cdy chuydn gene co
hoat dong cira hf th6n-e quang 2 trone ning rhylakoid t6r hon rrong c6y coi ctrung.

* ABA vti quang hqp

- ABA co vai trd quan trong trong tinh khdng cua cay doi voi stress crla meii
truong.
- Phun ABA l€n la lhm gidm quang hcro vi hoat d6ng cria enzyme ribulose
biphosphatc carboxylase (rubisco).

* IAA vd quang hop

- IAA kich thich quang hoo.

,'t
* Nhtittg chfit ili4u hba sinh trrring lhqrc v(t khdc vd quang hop
Ethylene vir nh&ng hop cir6t phong thich ethylene kh6ng inh huorrg hoic
iinh huong kich rhich quang hqp.
- BR: Tang tich luy chlorophyll. khi n[ng quang hqp su vdn chuy6n c6c sAn
pham dong hoa do quang hqp.
- Jasmonate c6 thti uc che quang hgp.
- Nhtng Anh hudng cua salicylic acid len quang hop chua duoc nghi€n criu

Guio Trirt Char Diiu I loit Sinh Truory Thvc I'tir 8

j
 nnleU.
6.4. Cic v6n cti vi phdng trit cd d4i
Co d4i c4nh tranh voi cdy ,r6ng v€ nuoc, dinh du6ng va anh sang. Cd dai
con lti noi cho ngudn bqnh va c6n trung dn ndu tl6 t6n c6ng cdy tr6ng. C6 thd t6m
.: . . ..;
'^ di€m
tat cac bit loi cua c6 dai trong l0 sau:

l. Giiim chat lugng ndng san.

2. CiiLrn nang suAr cay trdng va vit nu6i.
3. Tdng gia thanh sin Phim' +

4. Cin trd viEc qudn lY nuoc.

5. I-a vdn rtA O6i vOi sric khoe con nguoi'
6. Gitvi han hi€u qu6 con ngudi.
7. Gilin gia tri su dqng ddt.
8. Girrm su lua chon cdy trdng duoc trong tren mQt mdnh ddrt cho phep.
9..Tham hga.cua h6a hoan.
10. Kh6ng ttep mit.

6.4.1. Phu'ong PhdP Phdng trit c6

6.4.1.1. Ngin ngtra, phdng trir vi nhd cd
Vi€c ngan ngua, phong tru va nho c6 c6 3 tiic dung chinh:
- Ngirn ch+n s1r xam nhiEm cua nhtng loiri co msi.
- NgiLn ch{n sr; t4o hQt cia co.
- N-ein chdn su ldy lan cria co bing con duong sinh san sinh du0ng.

6.4.1.2. Qurin li co dai

' C6 thi t6rn tit cdc phuong phdp co ban sau d6 quan l! co dqi:
- Cldc bidn phdp co hgc nhu ciry bua' nhO bing tay. cit co, nrang pht. d6t vi
cho n-eip.
- Cdc bi€n phdp canh tiic nhu chon loqi ciy trdng' ludn canh, chon gi6n-e, bd
tri thrri vu, mqit do. khoang cdch tr6ng' phdn bon.--
- Phong trir sinh hoc.
Phone trir h6a hoc.

' 6.4.2. Gi6i thigu vi: phdng trir cii bang htia chit
IliCn tai dr-r cri hon ii() loai thu0c co chon loc khiic nhau tr€n thO giui. Co
rh€ pluin loiri thLr0c co thanh 5 nhonr ch[nh sau:

I . Drra vt\o lo4i ca;- slr dung.

2. Duu vr\o nltt-rn-9. anh htrong nhin thA;' tltroc.

O"n frit,Ulrii }ii,, Ilcxf Sinh TrLnmgThrc l'dr 3l

 J. l'lrco phrrong dicn hip rhu.
{. Drra vio linli luu clin hav tinh ti0p xirc.

-i. Dua r,io tinh chon loc.

Cac loai thu6c co dan-e su duns co nhiiu co ctr6 tac dQng khac nhau. Co th0
rtlm tit lirrh lrrrdrrg ctia nil llrcrr bn cr.r chi tiic dung clrinh nhu san: -

L ['.r'c clrC sinh lnrorrg cal trdng.

2. U'c ctr6 hO hip hoirc quang hgp.
i. lJc ch6 nhirng qu6 trinh sinh t6ng hgp.

6.4.7.1. Thrr6c co c6 tic dr.rng gi6ng nhu'IAA

l'hudc co thuoc nhom auxin co tdc dung gi6ng nhu IAA c6 thd kC cl6n la
nhtng phenoxy acid, benzoic acid va pl ridinc

6.4.2.1.1. Phenoxy acid: Kich thich tdng hgp etlrylene vd uc ch6 sinh trudng (2,4D,
Quinclorac. 2,4DR TCDD, MCPA, MCPP).
6.4.2.1.2. Benzoic acid: Dicamba. chloramben co phuong thftc tdc dQng gi6ng nhu
2.4D le gdy ra sg nghi€ng.
.6.4.2.1.3. Pyridine: Piclorarn, triclopl'r vd clopyralid. C6 hieu qud m4nh, luu tdn
l6u. phdn hriy chdrn bdi vi sinh vdt dat. Cfrng c6 tdc d6ng gi6ng nhu 2.4D ld -qAy ra
sr,r nghi€ng.

6.4.2.2. Ntrfrng ctr6t rfc cn6 t6ng hgp gibbcrellin: Nhtng ch6t uc ch6 sinh t6ng
hop gibberellin cfrng c6 thd sir dung- dd phong trir cd nhu flurprimidol,
paclobutrazol, va uniconazole.

6.4.3. Sr; ric ch6 qud trinh sinh t6ng hg'p, qunng hg'p vir hd h6p
6.4.3.1. Nhii'ng chAt ri'c ctr6 na hip (MAA, dinoseb, bromoxynil)
Thu6c c6 ngdn cdn qua trinh h6 hdp cria thuc vdt bing hai c6ch:
- Pha hny su oxy h6a vong phosphoryl nhu methylarsonic acid (MAA) vd
phenols (dinoscb).
- Ngan cdn su vqn chuy€n electron:Nhu nhom hydroxybenzoniiiile (vi du:
bormoxynil).

6.4.3.2. Cn6t *c ch6 quang hqp

Nhidu nh6m thuOc co co tac dQng uc ch6 quang hqp bing cdch uc ch6 sr,r
chuydn hoa ndng luong 6nh sdng thdnh nang luong h6a hoc vi du nhu urea, uracil,
triazine. triazinone. acylanilid va pyridazinone. C6c ch6t nay chen vao qurl trinh khri

ciio hinh ch/i, IMu llod siah Trnhg Thqc I at 85

 nhin PSII (hethong quang II). Nhung chdt nay.voinhtng
frlastoqu inone crla chdt
cau truc khac nhau gin chdt vdo mdng thylakoid lam ngan cAn
s,r yiin ,huyen
electron quang hop (Diuron, bromacil, simazine, propanil, pyrazon).

6,4.3.3. Nhtrng ch6t ri,c ch6 qud trinh sinh t6ng hqo
- Ngin can qud trinh phdn chia td bdro nhu carbamate (propham).
dinitroanilinc (<tryzalin) va dif'enzoquat (difenzoquar rnerhyrsulphate).
- Ngnn can tong hon nucreic acid vd protein nhu alphachroroacetamides.
- Ngin ciin rdng hup amino acid nhu surtbnyrureas (13ensufuron)-
irnidazol inones (imazapyr), glyphosate.
- t,t'c ctr6 tdng hon carotenoid (carotinoid) nhu amitrole, pyridazinones
(norllurazon). fl uridone.
- tjc ch6 sinh t6ng hop ripid nhu carbamothioate (Eprc), cyclohexanedione
(Scthoxyd irn), aryl oxyphenoxypropionare (Diclofop).
- Phd hiy mang t6 bao nhu p-nirro. rhay th6 diphenyrcthcr (ox1,fluorr.en),
bipyridyll iurn (paraquat).
- {Jc chti phdn chia t6 biio nhu propham (l-rnethylelhv} phen,vl carbamate),
oryzalin, dil'cnzoquat methyl sulphate.
- U'c ch6 tOng hqp protein va nucleic acid nhu alachlor.
- Nhirng ch6t uc ch0 arnino acid nhu sulfbnylurea va irnidazolinone. Chung
tuc chc cnz]-rxc acerolacrare synthase
{AI-s). Glyphosate la thudc c6 ph6 rong cln!
cri tlic dung nirv.
- Nhirng chit fc chd tdng hqp carotenoid nhu amitrole. norflurazon.
fluridone. chirng ric ch6 su tao thanh sic t6. vi su trii sinh ch6i.
IJc chd t6ng hqp lipid nhu Eprc, sethoxydim vd dicrofbp. chung duoc
hdp thu bcri ri va vdn chuy6n l6n rrr vi thdn lim irc ch6 vung phdn sintr uang
cacn
tinh hucrng l0n sinh r6ng hop lipiri.
-
Phir r.'tv rni)ng te bao nhu oxyflourf'en va paraquar. chirng duoc hdp rhu qua
chdi lon hcrn qua rd.

6.4.4. Cdng ngh6 di truy6n vi tinh khing thuiic co & thnc vit bdc cao
-l-rons
thuc tci iip ciung cac biqn ph6p trrr co. c6c nhi nghidn ctu lu6n tim
circh triing cao hitu cluii cia qua trinh di8t co bins nhiiu bitn phirp klac phau.
Sau
dir li tiirn tit t iic citrng citc loai hoa chit therrr
rrio rhudc dir-\ co hal cac bign phip
k1' thuiit c(r liin quan d.i li'rrn tiinq hie,u qui diCt trir ccj dai:

- Saf'ener: Dung d.i ndi rorrg tinhchon loc cua cil,rrdng aoivoi nhirng loai
thuoc co dlrt bi€t. Cic loai safener crii duoc dung nhu flurazole vi cl,ometronil.
+'[:lttritzolc': Giirp alachlor co th€ rltrtrc dung nhu mdt thuoc
co chon loc trtl
dr;'r,'*i7j,it D;, t 'oit Snh Tnone hvc I _ir
7
S6
 sor-qhunl.
+ C)ortrctrorril duoc dnng d6 bio r€ sorgliunr
AOI VOi rnetolachlor. Nhrlrn,,.
salcrrcrcri th0 dirng cho l[a. bip. lua rni va ltia nrach.
- linlr klrirrg rhucrc co cr trre trar drro-c bi'g cacrr gi.rn hap
trrrr tlru.c ctr.
tilrtlr r.i
.. fitcn drr()nq. br) surr5t. hcriic phirr I,r cua thu6c co
- civ nr6 r'rr corrg ngrre di truvin: \/icc srr dung corrg n-errc di rr.urcl,-'
.;
cl.ll-l-\cn qcnt'. llir! iicc plriit hiCn ra cic biin di tao
rrlrrr
ra tinlr klrin-g rrl rruoi
ci1 nro co thi tto ra rrlirirng car, khdnr r,o.i rhuoc co. 'c.irr 'hiCrr

ttoo lrtnh ( lrtjr l)rctu lloi Suth |ruongThvclir gj

 TAI LIEU THAM KHAO

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t\i