Mc lc

Gii thiu
I. Cc cht kch thch sinh trng thc vt......................................................................3
1.1. Auxin...................................................................................................................... 3
1.1.1 Ngun gc .......................................................................................................3
1.1.2. Cu trc ho hc v s sinh tng hp..............................................................4
1.1.3. Tnh cht sinh l ca auxin...............................................................................4
1.1.4. Cc cht auxin tng hp...................................................................................8
1.2. Gibberellin ........................................................................................................... 10
1.1.1. Ngun gc....................................................................................................10
1.1.2. Sinh tng hp ...............................................................................................11
1.2.3. Tnh cht sinh l ca gibberelin .....................................................................12
1.2.4. Cc cht khng-gibberelin..............................................................................14
1.3. Cytokinin............................................................................................................... 14
1.3.1. Ngun gc......................................................................................................14
1.3.2. Cu trc v sinh tng hp..............................................................................14
1.3.3. Cc loi cytokinin............................................................................................16
1.3.4. Tnh cht sinh l ca cytokinin........................................................................18
II. Cc cht c ch sinh trng thc vt........................................................................19
2.1. Axit abscisic (ABA)...............................................................................................19
2.1.1. S pht hin..................................................................................................19
2.1.2. Con ng sinh tng hp v s phn phi trong t bo................................20
2.1.3. Tnh cht sinh l ca acid abscisic..................................................................21
2.2. Etylen................................................................................................................... 22
2.1.1. S pht hin...................................................................................................22
2.2.2. Con ng sinh tng hp .............................................................................23
2.2.3. Vai tr ca etylen............................................................................................24
2.3. Nhm cc cht c bn cht phenol......................................................................25
III. Mt s nguyn tc khi s dng cht iu ho tng trng thc vt..........................26
IV. ng dng cc cht iu ho sinh trng trong trng trt .........................................28

TI LIU THAM KHO.........................................................................................35

Gii thiu
Trong i sng thc vt, ngoi cc cht hu c nh gluxit, protin, lipit, axit
nucleic... cu trc nn t bo, m v cung cp nng lng cho cc hot ng
sng ca chng, th cn c cc cht c hot tnh sinh l nh vitamin, enzyme
v cc hormone, trong cc hormone c mt vai tr rt quan trng trong vic
iu ha qa trnh sinh trng pht trin v cc hot ng sinh l ca thc vt.
Cc cht iu ha sinh trng v pht trin ca thc vt l nhng cht c bn
cht ha hc khc nhau, nhng u c tc dng iu tit cc qu trnh sinh
trng, pht trin ca cy t lc t bo trng th tinh pht trin thnh phi cho
n khi cy ra hoa kt qu, hnh thnh c quan sinh sn, c quan d tr v kt
thc chu k sng ca mnh. Cc hormone thc vt (phytohormone) l nhng
cht hu c c bn cht ha hc rt khc nhau c tng hp vi mt lng rt
nh cc c quan, b phn nht nh ca cy v t vn chuyn n tt c
cc c quan, cc b phn khc ca cy iu tit cc hot ng sinh l, cc
qu trnh sinh trng, pht trin ca cy v m bo mi quan h hi ha
gia cc c quan, b phn trong c th.
Bn cnh cc cht iu ho sinh trng t nhin (c tng hp trong c th
thc vt) cn c cc cht do con ngi tng hp nn (gi l cc cht iu ho
sinh trng nhn to).
Ngy nay bng con ng ho hc con ngi tng hp nn hng lot cc
cht khc nhau nhng c hot tnh sinh l tng t vi cc cht iu ha sinh
trng t nhin (phytohormone) iu chnh qu trnh sinh trng, pht trin
ca cy trng, nhm tng nng sut v phm cht ca cy trng. Cc cht iu
ho sinh trng nhn to ngy cng phong ph v c ng dng rng ri
trong sn xut nng nghip.
Cc cht iu ha sinh trng, pht trin ca thc vt c chia thnh hai
nhm c tc dng i khng v sinh l: cc cht kch thch sinh trng
(stimulator) v cc cht c ch sinh trng (inhibitor).
2

I. Cc cht kch thch sinh trng thc vt

Cc cht kch thch sinh trng ca thc vt l nhng cht nng sinh l c
tc dng kch thch cc qu trnh sinh trng ca cy. Cc cht kch thch sinh trng
thc vt gm c cc nhm cht: auxin, gibberellin, cytokinine.
1.1.

Auxin

1.1.1. Ngun gc

Nm 1880 Sacl acuyn (Darwin) pht hin ra rng bao l mm ca cy

h ha tho rt nhy cm vi nh sng. Nu chiu sng mt chiu th gy quang
hng ng, nhng nu che ti hoc b nh ngn th hin tng trn khng xy ra.
ng cho rng ngn bao l mm l ni tip nhn kch thch ca nh sng.
Vo nm 1885, mt nh khoa hc tn l Salkowski pht hin ra indole-3acetic acid (IAA) trong mi trng ln men. Th nhng qu trnh chit tch sn phm
tng t trong cc m thc vt khng thnh cng trong sut gn 50 nm sau.
Nm 1926, mt sinh vin tt nghip i hc ti H Lan tn Fritz Went cng
b bo co m t phng php phn lp cht kch thch tng trng bng cch t
nhng khi thch trng bn di nh ca l bao mm trong mt thi gian nht nh
sau ly ra v t chng vo thn cy khc b bm ngn. Sau khi t cc khi
thch , cc thn cy bt u tng trng tr li. Nm 1928, Went trin khai mt
phng php nh lng cht kch thch tng trng thc vt ny. Went thng c
bit n nh ngi tin phong s dng thut ng auxin, nhng tht s phi k n
Kogl and Haagen-Smit. H l nhng ngi tinh ch c hn hp axt
auxentriolic (auxin A) t nc tiu ca ngi vo nm 1931. Sau , Kogl phn
lp c nhng hn hp khc t nc tiu c cu trc v chc nng tng t auxin
A, trong c IAA.
Auxin ph bin nht, axt indoleacetic acid (IAA), thng c hnh thnh gn
nh tng trng v sau i xung. Qu trnh khin cho cc l non s mc di
hn. IAA kch thch cy ci pht trin hng theo nh sng v pht trin b r.
Vo nm 1954, mt hi ng cc nh sinh l hc thc vt c thnh lp
nh danh cho cc nhm auxin. Thut ng ny xut pht t ting Hy Lp, c ngha l

tng trng. Cc hp cht c gi chung l auxin nu chng tng hp trong cc

loi thc vt v l nhng cht chia s nhng hot ng tng t vi IAA.
1.1.2. Cu trc ho hc v s sinh tng hp
Auxin l mt hp cht tng i n gin, c
nhn indole, c cng thc nguyn l: C10H9O2N, tn
ca n l axit -indol-acetic.
Auxin c tng hp tt c cc thc vt bc
cao, to, nm v c vi khun. thc vt bc cao AIA
c tng hp ch yu nh chi ngn v t c
vn chuyn xung di vi vn tc 0,5 - 1,5cm/h.
S vn chuyn ca auxin trong cy c tnh cht phn cc rt nghim ngt, tc l
ch vn chuyn theo hng gc. Chnh v vy m cng xa nh ngn, hm lng
auxin cng gim dn to nn mt gradien nng gim dn ca auxin t nh ngn
xung gc ca cy. Ngoi nh ngn ra auxin cn c tng hp cc c quan cn
non khc nh l non, qu non, phi ht ang sinh trng, m phn sinh tng pht
sinh. Qu trnh tng hp auxin xy ra thng xuyn v mnh m trong cy di
xc tc ca cc enzyme c hiu. Axit -Indol Axetic l loi auxin ph bin trong
cy, c tng hp t tryptophan bng con ng kh amin, cacboxyl v oxy ha.
Auxin c tng hp thng khng dng t do, m lin kt vi mt acid amin
(acid aspartic Pisum, acid glutamic cy c chua), hay glucid (AIA-glucoz, AIAthioglucosid, AIA-inositol). Cc dng lin kt ny khng c hot tnh auxin nhng d
dng phng thch auxin theo con ng enzim (bi s thu gii kim trong thc
nghim), l cc dng d tr (khng b ph hu bi AIA-oxidaz) v vn chuyn ca
auxin.
1.1.3. Tnh cht sinh l ca auxin
Auxin can thip vp nhiu hin tng sinh l, hot ng ca n tu thuc vo
nng v cc s h tng qua li ca chng vi cc cht iu ho khc. Mt s
hot ng chnh ca auxin:
1.1.3.1. Hot ng trong s ko di t bo
4

Auxin kch thch mnh s ko di t bo ngn chi. S ko di t bo l mt

qu trnh phc tp, kt hp nhiu hin tng: hp thu nc; dn di vch vi sc
trng; t cc hp cht mi ca vch gia cc mng vi si cellulos; sinh tng hp
protein v cc cht khc.
Vai tr ca auxin l gy nn s gim pH ca thnh t bo bng cch hot ha
bm proton ( H+) nm trn mng ngoi cht. Khi c mt ca auxin th bm proton
hot ng v bm H+ vo thnh t bo lm gim pH v hot ha enzyme xc tc
ct t cc cu ni ngang ca cc polysaccarit. Enzyme tham gia vo qu trnh ny
l pectinmetylesterase khi hot ng s metyl ha cc nhm cacboxyl v ngn
chn cu ni ion gia nhm cacboxyl vi canxi to nn pectat canxi, do m
cc si cenlulose tch ri nhau.

Ngoi ra, auxin cng kch thch s tng hp cc mRNA cc cht ribosome tham
gia vo s tng hp cc cht protein.

1.1.3.2. Hot ng trong s phn chia t bo

Kch thch s phn chia t bo tng tng
Auxin kch thch rt mnh s phn chia t bo tng tng (tng pht sinh
libe - mc), nhng hu nh khng tc ng trn m phn sinh s cp. Nh vy,
auxin tc ng trn s tng trng theo ng knh.
nng cao, auxin kch thch s to m so t cc t bo sng nh vo cht
histogene ( l cht to ra nhiu t bo ging nhau hon ton). y l c tnh tt
c p dng trong nui cy t bo.
Phn ho m dn
Auxin kch thch phn chia ca tng tng, ng thi gip s phn ho ca
cc m dn (libe v mch mc). Auxin c kh nng cm ng trc tip s phn ho
t bo nhu m thnh cc t chc m dn.
1.1.3.3. Hot ng trong s pht sinh hnh thi (r, chi, qu)
Kch thch pht trin chi
Auxin (phi hp vi cytokinin) gip s tng trng chi non v khi pht s
to m phn sinh ngn chi t nhu m. Tuy nhin, nng cao, auxin cn s
pht trin ca pht th chi va thnh lp hay chi nch: cc chi by gi vo trng
thi tim sinh.
Auxin gy hin tng u th ngn: Hin tng u th ngn l mt hin
tng ph bin trong cy. Khi chi ngn hoc r chnh sinh trng s c ch
sinh trng ca chi bn v r bn. y l mt s c ch tng quan v khi loi tr
u th ngn bng cch ct chi ngn v r chnh th cnh bn v r bn c gii
phng khi c ch v lp tc sinh trng. Hin tng ny c gii thch rng
auxin c tng hp ch yu ngn chnh v vn chuyn xung di lm cho cc
chi bn tch lu nhiu auxin nn c ch sinh trng. Khi ct ngn chnh, lng
auxin tch lu trong chi bn gim s kch thch chi bn sinh trng.
Kch thch pht trin r
Auxin nng cao kch thch s to s khi r (pht th non ca r),
nhng cn s tng trng ca cc s khi ny. c tnh ny c ng dng ph
bin trong gim cnh. S hnh thnh r ph trong gim cnh c th chia lm 3 giai
6

on: giai on u l phn phn ho t bo trc tng pht sinh, tip theo l xut
hin mm r v cui cng mm r sinh trng thnh r ph chc thng v v ra
ngoi. Giai on u cn hm lng auxin cao, giai on r sinh trng cn t
auxin v c khi khng cn c auxin.
Kch thch s hnh thnh, s sinh trng ca qu v to qu khng ht
T bo trng sau khi th tinh to nn hp t v sau pht trin thnh phi. Phi
ht l ngun tng hp auxin ni sinh quan trng, khuych tn vo bu v kch thch
s sinh trng ca bu hnh thnh qu. V vy qu ch c hnh thnh khi c
s th tinh. Nu khng c qu trnh th tinh th khng hnh thnh phi v hoa s b
rng. Vic x l auxin ngoi sinh cho hoa s thay th c ngun auxin ni sinh
vn c hnh thnh trong phi v do khng cn qu trnh th phn th tinh
nhng bu vn ln ln thnh qu nh auxin ngoi sinh. Trong trng hp ny qu
khng qua th tinh v do khng c ht.
Km hm s rng l, hoa, qu ca cy, v n c ch s hnh thnh tng ri
cung l, hoa, qu vn c cm ng bi cc cht ch sinh trng. V vy phun
auxin ngoi sinh c th gim s rng l, tng s u qu v hn ch rng n, qu
non lm tng nng sut. Cy tng hp lng auxin s c ch s rng hoa, qu,
l.

Auxin c hnh thnh lin tc trong nh sinh

trng ca thn v r cy... (nh: www.nsf.gov)

So snh qu trong t nhin (bn tri) vi

qu c x l bng cht kch thch tng
trng (bn phi). (nh: www.planthormones.info )

1.1.4. Cc cht auxin tng hp

IAA tng hp c s dng trong nui cy m nhng n d b bin tnh trong
mi trng nui cy v nhanh chng thoi bin trong m. Tuy nhin nhng c
tnh ny c th tr nn hu dng bi v trong cy, IAA (cng vi cytokinin) sau khi
cm ng hnh thnh m so s kch thch s to chi hoc phi khi hm lng ca
n trong m gim dn. IAA thng c s dung phi hp vi cc cht iu ho
sinh trng khc kch thch s pht sinh hnh thi trc tip (s to r ca cnh
gim in vitro) v trong nui cy nh sinh trng v chi. Tuy nhin, tu theo mc
ch th nghim m ngi ta c th s dng cc hp cht ging auxin khc c
tng hp v n c nhng hot ng hi khc vi auxin nh:

Trng
Tn cht
3-Indoleacetic acid

Vit tt

lng

IAA

phn t
175.2

Dung mi
1N NaOH

Nhit bo qun
Dng
Dng
bt

lng

0oC

0 oC
8

3-Indolebutyric acid
-Naphthaleneacetic acid
2,4-Dichlorophenoxyacetic
acid
2,4,5-Trichlorophenoxyacetic
acid
p-Chlorophenoxyacetic acid
2-Methyl-4chlorophenoxyacetic acid
-Naphthyloxyacetic acid
3,6-Dichloro-2methoxybenzoic acid
4-Amino-3,5,6trichloropicolinic acid
Phenylacetic acid
2,3,5-Triiodobenzoic acid

IBA
NAA

203.2
186.2

1N NaOH
1N NaOH

2-8oC
-

0 oC
2-8oC

2,4-D

221.0

Water

2-8oC

2,4,5-T

255.5

EtOH

2-8oC

4-CPA

158.1

EtOH

2-8oC

MPCA

NOA

202.2

1N NaOH

2-8oC

Dicamba

186.6

Picloram

241.5

DMSO

2-8oC

PAA
TIBA

136.2
499.8

EtOH
1N NaOH

0C

2-8oC
0 oC

2,4-D thng c s dng phi hp vi cytokinin cm ng to m so v

huyn ph t bo v n s c thay th bi IBA hay NAA kch thch s pht
sinh hnh thi. IBA v NAA l loi auxin thch hp trong nui cy chi.
2,4,5-T him khi c s dng trong nui cy m thc vt, hu nh ch c
s dng cm ng s to m so v s pht sinh gin tip ca cy mt l mm
nh Avena, Oryza, v Panicum. Heyser v cng s (1983) nhn thy Triticum
aestivum, mt s th to c m so c kh nng sinh phi vi 2,4-D, mt s
khc ch to m so vi 2,4,5-T.
Dicamba thng c hiu qu trong s to m so c kh nng sinh phi
nhm cy mt l mm.
Picloram thng c s dng cm ng v duy tr m so hoc huyn
ph t bo ca cc loi cy l rng hoc cm ng s to m so c kh nng
sinh phi. Picloram c hiu qu i vi bin cc kiu di truyn rng hn. Ch
trong mt s rt t trng hp c bit, loi auxin ny c dng trong nui cy nh
sinh trng v nui cy nt on thn v c dng vi nng rt thp (0.012
0.4 M) khi phi hp vi mt cytokinin.
1.2.

Gibberellin

1.2.1. Ngun gc

Gibberellin l nhm phytohormone

th hai c pht hin sau auxin. T
nhng nghin cu bnh l bnh la von
do loi nm k sinh cy la Gibberella
fujikuroi (nm Fusarium moniliforme giai
on dinh dng) gy nn. Nm 1926, nh
nghin cu bnh l thc vt Kurosawa
(Nht Bn) thnh cng trong th nghim gy bnh von nhn to cho la v ng.
Yabuta (1934-1938) tch c hai cht di dng tinh th t nm la von
gi l gibberellin A v B nhng cha xc nh
c bn cht ha hc ca chng.

10

Nm 1955 hai nhm nghin cu ca Anh v M pht hin ra axit gibberellic

cy la b bnh la von v xc nh c cng thc ha hc ca n l C 19H22O6.

Nm 1956, West, Phiney, Radley tch c gibberellin t cc thc vt bc

cao v xc nh rng y l phytohormone tn ti trong cc b phn ca cy. Hin
nay ngi ta pht hin ra trn 50 loi gibberellin v k hiu A 1, A2, A3,... A52.
Trong gibberellin A3 (GA3) l axit gibberellic c tc dng sinh l mnh nht.
Ngi ta tm c gibberellin nhiu ngun khc nhau nh cc loi nm,
thc vt bc thp v thc vt bc cao.

1.2.2. Sinh tng hp

Gibberelin l nhng terpenoid, c cu to t 4 n v isopren (C 5):

CH2=C(CH3)-CH=CH2. Cc n v ny t nhiu b bin i trong phn t gibberelin.
Theo l thuyt, cc gibberelin c 20C, nhng nhiu cht ch cn 19C (do mt CH 3
b oxi ho thnh COOH, v nhm ny c kh carboxyl).
Acid mevalonic (C6), c ngun gc t acetyl CoA trong con ng h hp, l
cht khi u ca cc sinh tng hp terpenoid. T acid mevalonic, cc isopren
c thnh lp v kt hp nhau qua nhiu giai on cho kauren (C 20), sn phm
chuyn bit u tin trong con ng sinh tng hp giberelin. Mi cht c hot tnh
giberelin u c nhn giberelan, khi u l GA 12-aldehyd.

11

Tm li, cc giai on chnh ca con ng sinh tng hp cc gibberelin l:

Acetil CoA acid mevalonic Kauren GA12-aldehid cc GA Trong s
cc gibberelin, GA1 l cht chnh kch thch s ko di thn thc vt. GA 3 t gp
thc vt, nhng l cht c hot tnh trong cc sinh trc nghim, v c xem nh
cht chun cho cc gibberelin.
Gibberellin c tng hp trong phi ang sinh trng, trong cc c quan
ang sinh trng khc nh l non, r non, qu non... v trong t bo th c tng
hp mnh trong lc lp. Gibberellin vn chuyn khng phn cc, c th hng
ngn v hng gc ty ni s dng. Gibberellin c vn chuyn trong h thng
mch dn vi vn tc t 5- 25 mm trong 12 gi. Gibberellin trong cy cng tn ti
dng t do v dng lin kt nh auxin, chng c th lin kt vi glucose v
protin.
Gibberelin lin kt vi cc cht ng: nhiu gibberelin-glycosid c tm
thy thc vt, nht l trong cc ht. Khi cc gibberelin c p dng vo thc
12

vt, mt phn gibberelin thng b glycosyl ho; ngc li, gibberelin-glycosid c

th c i thnh gibberelin t do.
1.2.3. Tnh cht sinh l ca gibberelin
S ko di t bo
Gibberelin kim sot hng t cc vi si celluloz ( va mi c tng hp
nh celluloz synthetaz) trong vch t bo, hng t ny li do hng t ca cc
vi ng ngoi vi t bo quyt nh. Gibberelin cm ng s t cc vi ng theo
hng ngang nhiu kiu t bo ( k c cc t bo m gibberelin khng kch thch
s ko di), tuy nhin s phi hp hot ng gia gibberelin v auxin trong s t
cc vi ng cha c bit.
Gibberelin h thp nng Ca2+ trong vch ( c l bng cch kch thch s
hp thu ion ny vo trong t bo), v do gip s ko dn vch, v Ca 2+ cn s
ko dn vch dicot (khng cn monocot). Trong hot ng ny, vch t bo
khng b acid ho bi giberelin ( khc vi hot ng nhanh ca auxin).
Gibberelin cn hot ng ca cc peroxidaz vch t bo, do lm chm s
ho cng ca vch, hin tng do s to lignin di tc dng ca cc peroxidaz.
S ko di ca thn
Hiu qu sinh l r rt nht ca gibberellin l kch thch mnh m s sinh
trng ko di ca thn, s vn di ca lng. Hiu qu ny c c l do ca
gibberellin kch thch mnh ln pha gin ca t bo theo chiu dc. V vy khi x l
ca gibberellin cho cy lm tng nhanh s sinh trng dinh dng nn lm tng
sinh khi ca cy. Di tc ng ca gibberellin lm cho thn cy tng chiu cao
rt mnh (u xanh, u tng thnh dy leo, cy ay cao gp 2-3 ln). N khng
nhng kch thch s sinh trng m cn thc y s phn chia t bo.
S ko di lng v tng trng l
Kch thch s ko di lng, va do s ko di va do s phn chia t bo
thn, l c tnh ni bt ca gibberelin. Gibberelin kch thch mnh s phn chia t
bo nhu m v v biu b. X l gibberelin lm tng nng sut ma cy v ng
(do kch thch s ko di lng).
Giberelin liu cao (hay phi hp vi citokinin) kch thch mnh s tng
trng l (din tch c th gp i bnh thng nh Trfle, Radis). Trn l yn
mch hay dip tiu la, giberelin ch c vai tr lm tng hiu ng auxin.
13

 S ny mm, ny chi
Gibberellin kch thch s ny mm, ny chi ca cc mm ng, ca ht v
c, do n c tc dng trong vic ph b trng thi ng ngh ca chng. Hm
lng gibberellin thng tng ln lc chi cy, c, cn hnh ht thi k ngh, lc ht
ny mm.Trong trng hp ny ca gibberellin kch thch s tng hp ca cc
enzyme amilaza v cc enzyme thu phn khc nh protease, photphatase... v
lm tng hot tnh ca cc enzyme ny, v vy m xc tin qu trnh phn hy tinh
bt thnh ng cng nh phn hy cc polime thnh monome khc, to iu kin
v nguyn liu v nng lng cho qu trnh ny mm. Trn c s , nu x l
gibberellin ngoi sinh th c th ph b trng thi ng ngh ca ht, c, cn hnh k
c trng thi ngh su.
S ra hoa, qu
Trong nhiu trng hp ca gibberellin kch thch s ra hoa r rt. nh
hng c trng ca s ra hoa ca gibberellin l kch thch s sinh trng ko di
v nhanh chng ca cm hoa. Gibberellin kch thch cy ngy di ra hoa trong iu
kin ngy ngn (Lang, 1956).
Gibberellin nh hng n s phn ha gii tnh ca hoa, c ch s pht
trin hoa ci v kch thch s pht trin hoa c.
Gibberellin c tc dng ging auxin l lm tng kch thc ca qu v to qu
khng ht. Hiu qu ny cng r rt khi phi hp tc dng vi auxin.

1.2.4. Cc cht khng-gibberelin

Vi cht lm chm tng trng, c dng to cy ln, l cc cht khnggibberelin. Th d: CCC (Chlorocholine chloride, trng lng phn t: 158.1), Amo1618 v phosfon cn s tng hp kauren; paclobutrazol ( tn thng mi: Bonzi)
cn giai on sau kauren, Acid abcisic khng cn chuyn bit

s tng hp

gibberelin, nhng hot ng i nghch vi gibberelin.

1.3. Cytokinin
1.3.1. Ngun gc

14

Vic pht hin ra cytokinin gn lin vi k thut nui cy m t bo thc vt.

Nm 1955 Miller v Skoog pht hin v chit xut t tinh dch c thu mt hp cht
c kh nng kch thch s phn chia t bo rt mnh m trong nui cy m gi l
kinetin (6-furfurryl-aminopurin: C10H9N5O).
Letham v Miller (1963) ln u tin tch c cytokinin t nhin dng
kt tinh t ht ng gi l zeatin v c hot tnh tng t kinetin. Sau ngi ta
pht hin cytokinin c trong tt c cc loi thc vt khc nhau v l mt nhm
phytohormone quan trng trong cy. Trong cc loi cytokinin th 3 loi sau y l
ph bin nht: Kinetin (6- furfuryl- aminopurin), 6-benzin- aminopurin v zeatin t
nhin.
Nc da (phi nh lng) t lu c dng trong nui cy (Van Overbeek et
al., 1941). Mi trng cha auxin v 10-20% nc da gip s phn chia ca t
bo thn phn ho (s to m so). Ngi ta tm cch xc nh bn cht ho
hc ca cht c trong nc da, nhng phi sau s khm ph cytokinin vi nm,
nc da mi c chng minh cha zeatin (Letham, 1974)
Sau zeatin, hn 30 cytokinin khc nhau c c lp. Ngy nay, ngi ta gi
cytokinin ch mt nhm cht thin nhin hay nhn to, c c tnh sinh l ging
nc da hay kinetin.
1.3.2. Cu trc v sinh tng hp
Zeatin t do dng trans trong phn ln thc vt, mc d c 2 dng cis v
trans u c hot tnh ca cytokinin. Nhiu cht tng hp c hot tnh cytokini,
chng u l cc aminopurin c thay th v tr 6, th d benzylaminopurin
(benzyl adenin, vit tt BAP hay BA) l cht c dng trong nng nghip. Ngoi
l, vi dn xut diphenilur c hot tnh cytokinin nhng yu.

15

Chui ngang ca cc cytokinin thin nhin c lin h v mt ho hc vi cao

su, carotenoid, gibberelin, axit abcisic, v vi hp cht bo v thc vt gi l
phytoalexin. Tt c cc hp cht ny, t ra l mt phn, c thnh lp bi cc n
v isopren c ngun gc t tin cht acid mevalonic. thc vt, cytokinin synthaz
l enzym xc tc s lin kt gia chui bn v adenosin monophosphat (AMP).
M phn sinh ngn r l ni tng hp ch yu cc cytokinin t do cho c c
th thc vt. T r, cytokinin di chuyn trong mch mc ti chi. Tuy nhin, cc
chi ( c chua) v phi cng l ni tng hp cytokinin.
Khc vi m phn sinh ngn chi, phi b tch khi cy vn tip tc tng
trng v pht trin bnh thng trn mi trng thiu hormon. Ngi ta khng
bit chnh xc khi no phi t lp v cytokinin, tuy nhin, c l phi qu non ( khng
c kh nng tng hp cytokinin) dng cytokinin hin din hm lng cao trong
phi nh.
Phn t tRNA (tham gia trong s tng hp protein trong cytosol hay trong dip
lp) khng ch cha 4 nucleotid to nn mi RNA, m cn vi nucleotid khng bnh
thng vi cc baz b bin i. Vi baz ny hot ng nh cytokinin khi tRNA b
16

thu gii (theo con ng enzym troong t bo). Nh vy, tRNA ca thc vt (v
hu nh ca mi sinh vt t vi khun n con ngi) cha zeatin, mc d dng
ng phn cis thay v trans nh cc cytokinin t do.
1.3.3. Cc loi cytokinin
Cc loi cytokinin t nhin: ngy nay, ngi ta cho rng kinetin khng phi l
cht t nhin m n c to thnh do s ti sp xp li cu trc ca mt cht
khc (Hecht, 1980), c t nht hai loi cytokinin t nhin c cu trc tng t nh
cu trc ca kinetin c xc nh, l nhgng hp cht t do hay nhng hp
cht c gn vi nhm glucoside hoc riboside (Entsch v cng s, 1980). Hai loi
cytokinin thng c s dng trong nui cy m l:
Zeatin:

4-hydroxy-3-methyl-trans-2-butenylaminopurine,

hoc

6-(4-hydro-3-

methylbut-2-enyl)-aminopurine hoc 2-methyl-4(-1H-purine-6-ylamino)-2-buten-1-ol)

2-iP (IPA): N6-(2-isopentyl)adenine hoc 6-(3-methyl-2-butenylamino)purine.
Dihydrozeatin: 6-(-hydroxy=3=methyl-trans-2-butenyl)aminopurine.

Cc loi cytokinin tng hp: cc loi cytokinie t nhin nh 2-iP v zeatin t

c s dng trong cc th nghim v gi thnh cao. Mt s hp cht tng hp
thuc nhm cytokinin thng c s dng trong cng tc nui cy l:
Kinetin: 6-furfurylaminopurine hoc N-(2-furanylmethyl)-1H-purine-6-amine
BAP (BA): 6-benzylaminopurine hoc benzyladenine .
TDZ: 1-phenyl-3(1,2,3 thiadiazol-5-yl)

17

 Trng lng phn t ca mt s cytokinin:

Tn cht
Adenine
6-Benzylaminopurine
Zeatin
6-(3-methyl-2butenylamino)purine

Vit
tt

Trng
lng

ADE
BAP
ZEA

phn t
135.1
225.3
219.2

2-iP

203.2

Dung mi

Nhit bo qun
Dng
Dng
bt

lng

1.0 HCI
1N NaOH
1N NaOH

0oC

2-8oC
2-8oC
0 oC

1N NaOH

0oC

0 oC

18

Kinetin
KIN
1-phenyl-3(1,2,3 thiadiazol-5-yl)
TDZ
1,3-Diphenylurea
DPU
1.3.4. Tnh cht sinh l ca cytokinin

215.2
220.2
212.3

1N NaOH
DMSO
DMSO

0oC
-

0 oC
2-8oC
2-8oC

Tnh cht c trng ca xytokinin l kch thch s phn chia t bo mnh m.

V vy ngi ta xem chng nh l cc cht hot ha s phn chia t bo, nguyn
nhn l do xytokinin hot ha mnh m qu trnh tng hp axit nucleic v protein
dn n kch s phn chia t bo.
Cytokinin nh hng r rt ln s hnh thnh v phn ha c quan ca thc
vt, c bit l s phn ha chi. Ngi ta chng minh rng s cn bng gia
t l auxin (phn ha r) v Cytokinin (phn ha chi) c ngha rt quyt nh
trong qu trnh pht sinh hnh thi ca m nui cy in vitro cng nh trn cy
nguyn vn. Nu t l auxin cao hn cytokinin th kch thch s ra r, cn t l
cytokinin cao hn auxin th kch thch ra chi. tng h s nhn ging, ngi ta
thng tng nng cytokinin trong mi trng nui cy giai on to chi.
trong cy r l c quan tng hp cytokinin ch yu nn r pht trin mnh th hnh
thnh nhiu cytokinin v kch thch chi trn mt t cng hnh thnh nhiu.
Cytokinin km hm qu trnh gi ha ca cc c quan v ca cy nguyn vn.
Nu nh l tch ri c x l cytokinin th duy tr c hm lng protein v
chlorophin trong thi gian lu hn v l tn ti mu xanh lu hn. Hiu qu km
hm s gi ha, ko di tui th ca cc c quan c th chng minh khi cnh dm
ra r th r tng hp cytokinin ni sinh v ko di thi gian sng ca l lu hn.
Hm lng cytokinin nhiu lm cho l xanh lu do n tng qu trnh vn chuyn
cht dinh dng v nui l. Trn cy nguyn vn khi b r sinh trng tt th lm
cho cy tr v sinh trng mnh, nu b r b tn thng th c quan trn mt t
chng gi.
Cytokinin trong mt s trng hp nh hng ln s ny mm ca ht v ca
c. V vy nu x l cytokinin c th ph b trng thi ng ngh ca ht, c v chi
ng.
Ngoi ra cytokinin cn c mi quan h tng tc vi auxin, cytokinin lm yu
hin tng u th ngn, lm phn cnh nhiu. Cytokinin cn nh hng ln cc

19

qu trnh trao i cht nh qu trnh tng hp axit nucleic, protein, chlorophin v v

vy nh hng n cc qu trnh sinh l ca cy.

II.

Cc cht c ch sinh trng thc vt:

Qu trnh sinh trng v pht trin ca cy c m bo bi hai tc nhn c

tc dng sinh l i lp nhau l tc nhn kch thch v tc nhn c ch. S cn
bng gia cc cht kch thch sinh trng v cc cht c ch sinh trng c
ngha quan trng trong vic iu ho s sinh trng, pht trin ca cy. Ln u
tin Lucuyn (Luckwil, 1952) tch c auxin v cht c ch sinh trng bng
phng php sc k trn giy. Ngy nay ngi ta pht hin ra nhiu cht c
ch sinh trng c hnh thnh trong cy v c gi l cc cht c ch sinh
trng t nhin. Cc cht c ch sinh trng t nhin phn b rng ri trong cc
b phn ca cy. Ngi ta pht hin chng khng nhng trong cc c quan dinh
dng nh thn, l, chi, r m cn trong cc c quan sinh sn nh ht, c, qu...
c bit khi cc c quan ny trng thi ng ngh.
c tnh chung ca cc cht c ch sinh trng t nhin l tch ly nhiu
trong cc m, cc c quan thi k ng ngh. c ch s ln ln ca t bo, c
ch s ny mm ca ht, s sinh trng ca chi. Km hm s hot ng ca cc
cht kch thch sinh trng.
Cn c vo bn cht ha hc v tc dng sinh l ngi ta chia cc cht c
ch sinh trng t nhin thnh ba nhm: Nhm cc cht c bn cht tecpenit m
i din l axit absxisic (AAB), etylen v nhm cc cht c bn cht phnol.

Axit abscisic (ABA)

2.1.1. S pht hin
Nm 1961, hai nh khoa hc ngi M Liu v Carn tch c mt cht
di dng tinh th t qu bng gi v khi x l cho cung l bng non gy ra
hin tng rng v gi cht l Abscisic I.
Nm 1963, Chkuma v Eddicott tch c mt cht t l gi cy u nga
v t tn l Abscisic II. Vo thi gian ny Wareing v cc cng s cng tch
c mt cht c ch c trong cc chi ang ng v t tn l min. Nm 1966,
20

dng phng php quang ph phn cc xc nh c bn cht ho hc ca

cht c ch ny. Nm 1967, hi ngh khoa hc quc t t tn cho cht c ch
sinh trng ny l axit

abscisic (AAB) v c cng

thc

C15H20O4.

ho

hc

2.1.2. Con ng sinh tng hp v s phn phi trong t bo

Acid abscisic l mt sesquiterpen (3 n v isopren). S nghin cu trn cc
Cercospora (nm tit nhng lng ln acid abcisic vo mi trng nui cy) cho
thy, ging gibberelin (cng l hp cht terpen), s tng hp acid abscisic bt u
t mevalonat, qua cc cht trung gian IPP (isopentenilpyrophsphat), DMAPP
(dimetilalilpyrophosphat), v farnesilpyrophosphat (sesquiterpen C 15). Tuy nhin,
thc vt bc cao, mevalonat- 14C gia nhp rt t vo acid abscisic, con ng sinh
tng hp acid abcisic tht s qua cc carotenoid (violaxantin hay neoxantin). S
peroxy-ho cc hp cht ny (nh lipoxygenaz) phng thch xantoxin (C 15).

21

Con ng sinh tng hp acid abscisic gin tip qua cc carotenoid c

xc nhn qua 2 s kin: s tng hp carotenoidxy ra ng thi vi s gia tng
hm lng acid abcisic trong m: vi t bin (bp, c chua, i mch,
Arabidopsis), cc xo trn trong sinh tng hp carotenoid dn ti s gim hm
lng acid abcisic.
Acid abscisic l mt acid yu, vt qua mng t bo d dng khi dng trung
tnh (gn proton). AAB hin din 70% trong dip lp,15% trong cytosol, 10% trong
khng bo v 5% trong apoplast. Dng lin kt vi glucoz ch gp trong khng bo.
mc biu b, acid abscisic c thy ch yu trong cc t bo kh khu.

2.1.3. Tnh cht sinh l ca acid abscisic

Axit absisic kch thch s xut hin ri phn cung, iu chnh s rng ca
cc c quan ca cy, v vy cc b phn gi sp rng cha nhiu axit absisic.
Trong cc c quan ang ng ngh, hm lng axit absisic tng gp 10 ln so
vi thi k sinh trng. S ng ngh ko di cho n khi no hm lng axit
absisic trong c quan ng ngh gim n mc ti thiu. Do vy t trng thi ng
ngh chuyn sang trng thi ny mm c s bin i t l gia axit absisic v
gibberellin trong cc c quan.
Axit absisic c chc nng iu chnh
s ng m ca kh khng. X l axit
absisic ngoi sinh cho l lm kh khng
ng li nhanh chng, v vy m lm gim
s thot hi nc ca l. Chc nng iu
khin s ng m kh khng c lin quan
n s vn ng nhanh chng ca ion K +.
Axit absisic gy cho t bo ng to nn
l thng K+, mt sc trng v kh
khng ng li. X l axit absixic ngoi sinh lm kh khng ng li hn ch s
thot hi nc qua kh khng, gim s mt nc ca l.
Axit absisic c xem l mt hormone ca Stress v khi gp cc iu kin
ngoi cnh bt li th hm lng ca n tng ln v tng tnh chng chu ca cy.
22

V d khi gp hn hm lng axit absixic trong l tng nhanh lm kh khng ng

li lm gim s thot hi nc ca cy. y l mt hnh thc thch nghi ca cy
trong iu kin kh hn.
Axit absisic cn c xem nh l mt hormone ca s gi ha, mc gi
ha ca c quan gn lin vi s tng lng axit absisic. Trong chu k sng, thi
k cy bt u ra hoa to qu, ht, c... hm lng axit absisic tng ln cho n
giai on cui. V vy, sau khi cy ra hoa th cy mau gi v rt ngn chu k sng
ca mnh.
Axit absisic c ch s tng hp axit nucleic trong t bo, c ch qu trnh
tng hp protein, t nh hng n qu trnh sinh trng pht trin ca cy,
lm cy mau gi v rt ngn chu k sng.

Etylen
S pht hin
Etylen l mt cht kh n gin kch thch s chn ca qu. Nm 1917, khi
nghin cu qu trnh chn ca qu thy c xut hin etylen. T nm 1933-1937
nhiu nghin cu khng nh n c sn xut trong mt s nguyn liu thc vt,
c bit l trong tht qu.
Nm 1935, Crocker v mt s cng s ngi M cho rng etylen l hormone
ca s chn. Sau bng cc phng php phn tch cc nhy c pht hin
ra etylen c trong tt c cc m ca cy v l mt sn phm t nhin ca qu trnh
trao i cht trong cy.

Con ng sinh tng hp

Con ng sinh tng hp etylen khi s t metionin, cht ny thng dng
hot ng SAM (S-adenoslmetionin), v qua ACC (acid 1-amino-cyclopropan-1carboxylic, tin cht ngay trc etylen.

23

AVG (aminoethoxyvinylglicin) v AOA (aminooxiacetic acid) cn s i SAM

thnh ACC. Cobalt cn bc sau cng ACC thnh etylen. Ag + (AgNO3 hay thiosulfat
Ag) cn mnh hot ng ca etylen, theo cch rt chuyn bit (cc ion khc khng
c hiu ng ny). Du hiu qu km hn Ag +, CO2 nng cao (5-10%) cn
nhiu hot ng ca etylen (hiu ng ny ca CO 2 c p dng gi tri cy
lu chn)
Auxin v cytokinin kch thch s sn xut etylen bng cch kch thch s thnh
lp ACC (vi hiu ng ca auxingin tip qua etylen: rng l, ra hoa cy thm).
Acid abcisic cn s thnh lp ACC.

Vai tr ca etylen
Etylen c tc dng lm qu mau chn. Nhiu nghin cu chng minh etylen
gy nn hai hiu qu sinh ha trong qu trnh chn ca qa: Gy nn s bin i
tnh thm ca mng trong cc t bo tht qu, dn n s gii phng cc enzyme
vn tch ri do mng ngn cch, c iu kin tip xc d dng v gy nn nhng
phn ng c lin quan n qu trnh chn nh enzyme h hp, enzyme bin i
chua, mm ca qu.... Mt khc etylen c nh hng hot ha ln s tng hp

24

cc enzyme mi gy nhng bin i trong qu trnh chn. Etylen l hormone xc

tin s chn qu, c sn sinh mnh trong qa trnh chn v rt ngn thi gian
chn ca qu.
Etylen cng tng tc vi axit absixic gy s rng ca l, hoa, qa. Etylen
hot ha s hnh thnh t bo tng ri cung ca cc b phn bng cch kch
thch s tng hp cc enzyme phn hy thnh t bo (xenlulase) v kim tra s
gii phng cc cenlulose ca thnh t bo. Etylen c tc dng sinh l i khng vi
auxin, v vy s rng ca cc c quan ph thuc vo t l auxin/etylen. Nu t l
ny cao th ngn nga s rng, cn t l ny thp th ngc li.
Etylen kch thch s ra hoa ca mt s thc vt, nu x l etylen hoc cc
cht c bn cht tng t nh etylen (axetylen) c tc dng kch thch da, xoi ra
hoa tri v, tng thm mt v thu hoch.
Etylen c tc dng i khng vi auxin. Trong t bo cc b phn ca cy,
nu t l auxin/etylen cao s lm cho cc b phn cy sinh trng tt, cy lu gi.
Etylen nh hng n s phn ha r bt nh ca cc cnh gim, cnh chit.
X l etylen kt hp vi auxin cho hiu qu cao hn vic x l auxin ring r.
Etylen cn gy hiu qu sinh l ln nhiu qu trnh sinh l khc nhau nh gy
nn tnh hng ca cy, c ch s sinh trng ca chi bn, xc tin s vn
chuyn ca auxin, tng tnh thm ca mng.

2.3. Nhm cc cht c bn cht phenol

Cc hp cht c bn cht phenol trong cy l sn phm trao i cht, c tc
dng c ch qu trnh trao i cht v c ch s sinh trng ca cy. Trong cy
chng thng dng lin kt vi gluxit to nn cc glucozit lm mt tc dng c
ch ca n. Khi trng thi t do chng c tc dng c ch cc qu trnh trao i
cht trong cy.
Nhm cc cht c bn cht phenol bao gm rt nhiu cht khc nhau. Cc i
din ca nhm ny gm cc cht nh: acid jasmonic, acid salisilic, cc
brassinosteroid v cc polyamin c vai tr trong s truyn mt du hiu bn ngoi
(stress, vt thng, sinh vt k sinh). Tuy nhin, s hiu bit v cc cht ny nh
hormon tng trng thc vt khc cn ri rc, cha y .
25

Vai tr sinh l ch yu ca cc hp cht c bn cht phenol : l hot

ha enzyme phn hy auxin AIA-oxidase lm gim hm lng auxin trong cy,
do km hm s gin ca t bo v c ch s sinh trng ca cc c quan b
phn trong cy; Xc tin hnh thnh lignin lm t bo ha g nhanh. Cng vi axit
abcisic cc cht c bn cht phenol nh hng n s ng ngh ca cy, c ch
s ny chi ca cy... Tuy nhin, vai tr km hm ca chng i vi s sinh trng
ca cy khng c ngha quyt nh.

26

III. Mt s nguyn tc khi s dng cht iu ho tng trng thc

vt:
Hin nay trong lnh vc ha hc nng nghip, vic s dng cc cht iu ha
sinh trng trong trng trt ang pht trin mnh m vi nhng mc ch khc
nhau. Cc cht iu ha sinh trng ca thc vt ngy nay v ang c s
dng rng ri trong trng trt nh l mt phng tin iu chnh ha hc quan
trng i vi s sinh trng, pht trin ca cy nhm tng nng sut ca cy trng,
nng cao hiu qu lao ng, tit kim cng sc v thi gian canh tc...
Khi s dng cc cht iu ha sinh trng trong trng trt cn lu cc
nguyn tc sau y.
3.1. Nng s dng: Hiu qu tc dng ca cc cht iu ha sinh trng
ph thuc vo nng . Nu s dng kch thch th dng nng thp, nu
dng c ch sinh trng hoc dit tr c th s dng nng cao. Mt khc cc
b phn khc nhau v tui ca cy khc nhau cm ng vi cc cht iu ha sinh
trng khng ging nhau, r v chi c cm ng mnh vi auxin hn thn cy.
Cy non c cm ng mnh hn cy gi. V vy mun s dng cc cht iu ha
sinh trng c hiu qu cn phi xc nh tng loi cy trng, thi k sinh trng
v cc cht kch thch sinh trng tng ng khc nhau.

3.2. Nguyn tc phi hp: Khi s dng cc cht iu ha sinh trng phi
tha mn c cc iu kin sinh thi v cc yu t dinh dng cho cy. V cc
cht iu ha sinh trng lm tng cng cc qu trnh trao i chtm khng
tham gia trc tip vo trao i cht, nn khng th dng cc cht thay th
cht dinh dng. V vy, mun s dng cht iu ha sinh trng c hiu qu cao
cn phi xc nh thi v v vng cy trng thch hp c cc iu kin sinh thi
ph hp nh yu t nhit , nh sng , m.... ng thi cn p ng y
nc v phn bn cho cy trng. Cng xut pht t ngi ta s dng bin php
phun hn hp cc cht iu ha sinh trng v cc nguyn t khong a lng v
vi lng nhm tng nng sut mt s loi cy trng . Nh vy r rng gia cc cht
iu ha sinh trng v phn bn c mi quan h khng kht. Phn bn lm tng
cng hiu qu kch thch ca cc cht iu ha sinh trng. Ngc li cc cht
27

kch thch lm tng hiu qu ca phn bn. V vy vic s dng phi hp gia phn
bn v cht iu ha sinh trng c ngha rt ln v cng l mt hng quan
trng trong nng nghip hin nay.

3.3. Nguyn tc i khng sinh l gia cc cht iu ha sinh trng ni

sinh v ngoi sinh: Khi s dng cht iu ha sinh trng cn ch nguyn tc
i khng gia cc nhm cht sau: Chng hn s i khng sinh l gia auxin x
l v etylen ni sinh trong vic ngn nga s rng l, hoa, qu; S i khng gia
gibberellin ngoi sinh v axit absisic ni sinh trong vic ph b trng thi ng ngh
ca cy; S i khng gia auxin v xytokinin trong s phn ha r v chi...

3.4. Nguyn tc chn lc: Nguyn tc ny thng p dng vi cc cht dit

tr c di. Cc cht dit tr c c tnh c chn lc cao. Mt cht dit c ch c tc
dng c i vi mt s loi cy nht nh m t hoc khng c i vi nhng loi
cy khc. Kh nng c chn lc ny c th ph thuc vo c trng gii phu c
kh nng ngn chn s xm nhp ca thuc hay kh nng phn hy nhanh trong
cy nh c cc enzyme c hiu... Do phi chn loi thuc dit c v khng c
cho cy trng, ng thi phi hp mt s thuc khc nhau dit ht cc i
tng c vn mn cm vi thuc rt ln. Chng hn cc dn xut ca axit
phenoxyaxetic ch dit c hai l mm m t c vi cy mt l mm nn c s
dng dit c trong rung cy ha tho nh la, ng.... Ngc li IPC
(Izopropinphenyl cacbamat) c i vi cy mt l mm m khng c vi cy hai
l mm. V vy dit c hn hp cn phi phi hp hai loi thuc ni trn.

28

IV. ng dng cc cht iu ho sinh trng trong trng trt :

Mt s ng dng ca cc cht iu ha sinh trng trong trng trt nh sau.
4.1. Kch thch sinh trng ca cy, tng chiu cao, tng sinh khi v tng
nng sut cy trng:
Trong sn xut nng nghip mc ch cui cng l nng cao sn lng c
quan thu hoch. Khi s dng cc cht iu ha sinh trng vi nng thp s c
tc dng kch thch s sinh trng, tng lng cht kh d tr , nn lm tng thu
hoch. Trong lnh vc ng dng ny c th s dng cc cht nh gibberellin (GA),
axit - naphtin axtic (-NAA). c bit s dng GA em li hiu qu cao i vi
nhng cy ly si, ly thn l v n c tc dng ln ton b c th cy lm tng
chiu cao cy v chiu di ca cc b phn ca cy. Phun dung dch GA nng
20 - 50 ppm cho cy ay c th lm tng chiu cao gp i m cht lng si ay
khng km hn. i vi cc cy rau vic tng sinh khi c ngha quan trng,
ngi ta thng phun GA cho bp ci, rau ci cc loi vi nng dao ng trong
khong 20 -100 ppm lm tng nng sut r rt. X l GA cho cy ch c tc dng
c tc dng lm tng s lng bp v s l ca ch, khi phun vi nng 0,01%
c th lm tng nng sut ch ln 2 ln, trong mt s trng hp c th tng nng
sut ln 5 ln.
4.2. Kch thch s hnh thnh r ca cnh gim, cnh chit.:
Phng php nhn ging v tnh i vi cc loi cy trng l mt phng
php nhn ging ph bin trong trng trt. Trong gim cnh v chit cnh ca cc
loi cy nh cy n qu, cy cng nghip, cy cnh, cy thuc thng s dng cc
cht kch thch sinh trng. Vic s dng mt s cc cht kch thch trng
nng cao hiu qu r rt v n kch thch s phn chia t bo ca m phn sinh
tng tng hnh thnh m so (callus) ri t hnh thnh r mi. x l ra r
ngi ta thng dng cc cht nh:Axit - indol axetic (IAA); Axit -indol butiric
(AIB); -NAA; 2,4-D; 2,4,5-T... Nng s dng ty thuc vo phng php ng
dng, i tng s dng v ma v.
Hin nay c 2 phng php chnh x l cho cnh gim v cnh chit.

29

- Phng php x l nng c hay phng php x l nhanh. Nng

cht kch thch dao ng t 1.000 - 10.000 ppm. Vi cnh dm th nhng phn gc
vo dung dch t 3-5 giy, ri cm vo gi th. Phng php x l nng c c
hiu qu cao hn c i vi hu ht cc i tng cnh gim v nng hiu qu
cho nhiu loi i tng l 4.000 - 6.000 ppm. Vi cnh chit th sau khi khoanh v,
tm bng bng dung dch cht kch thch c ri bi ln trn ch khoanh v, ni s
xut hin r bt nh. Sau b bu bng t m. Phng php ny c u im l
hiu qu cao v gy nn ci xc sinh l cn cho giai on u ca s xut hin r.
- X l nng long - x l chm. Nng cht kch thch s dng t 20 200 ppm ty thuc vo loi v mc kh ra r ca cnh gim. i vi cnh gim
th ngm phn gc ca cnh vo dung dch t 12 - 24 gi, sau cm vo gi th.
Vi phng php ny th nng hiu qu l 50 - 100 ppm. i vi cnh chit th
trn dung dch vo t b bu b bu cho cnh chit. V d c th dng 2,4D
chit nhn vi nng 20ppm v chit cam, qut vi nng 10 -15ppm cho kt
qu tt. Vic xc nh nng v thi gian x l thch hp tng loi cht iu ha
sinh trng trn tng loi cy trng trong vic gim, chit cnh cn c nghin
cu mt cch k lng mi cho kt qu tt. Thi v gim v chit cnh tt nht l
vo ma xun sang h (thng 3,4,5) v ma thu (thng 9,10).
4.3. Tng s u qu v to qu khng ht:
Sau qu trnh th phn, th tinh th qu bt u c hnh thnh v sinh
trng nhanh chng. S ln ln ca qu l do s phn chia t bo v c bit l
s gin nhanh ca t bo trong bu. S tng kch thc, th tch ca qu mt cch
nhanh chng l c trng s sinh trng ca qu. S sinh trng nhanh chng
nh vy l do c iu chnh bng phytohormone c sn sinh trong phi ht.
Ht c hnh thnh l do qu trnh th phn, th tinh xy ra. Nu chng ta x l
auxin v gibberellin ngoi sinh cho hoa trc khi th phn th tinh thay ngun
phytohormone ni sinh t phi th qu s c hnh thnh m khng cn th tinh,
trong trng hp ny qu s khng c ht. Ngi ta thng dng cc cht kch
thch nh -NAA, GA... phun cho hoa mi n th c th loi b c s th phn,
th tinh m qu vn ln c. V vy lm cho qu ln ln nhng khng c ht hoc
t ht, nng sut cao v phm cht tt. Nng s dng ty thuc vo cc cht

khc nhau v cc loi khc nhau. C th to ra qu khng ht i vi nhiu i

tng cy trng nh c chua, nho, cam, qut, t, da hu, da chut... Chng hn
phun -NAA nng 10 - 20 ppm cho c chua, phun GA cho nho hai ln trong thi
k ra hoa r v hnh thnh bu qu vi nng 0,01 - 0,02% (100 - 200 ppm) lm
tng kch thc v trng lng qu. Phun GA cho cy trng thuc h cam, chanh
trong giai on n hoa vi nng dung dch 0,025 - 0,1% lm tng nng sut v
phm cht qu (v mng, mu p, hm lng vitamin C tng). Vi to c th
dng GA nng 400 ppm hoc phi hp gia GA (250 ppm) vi auxin (10 ppm).
Vic x l to qu khng ht c ngha quan trng trong vic lm tng phm
cht ca qu, c bit l cc loi qu cha nhiu tht qu.
4.4. Ngn nga s rng n, hoa v qu:
tng nng sut cy trng, bn cnh bin php xc tin hnh thnh qu, cn
ngn nga hin tng rng n, hoa v qu non. Nguyn nhn ca hin tng ny
l khi qu sinh trng nhanh th hm lng auxin ni sinh t ht khng cung
cp cho qu ln. Nu gp mt s iu kin bt thun th s tng hp axit abxixic v
etylen tng nhanh lm cho s cn bng hormone thun li cho s rng, tng ri
xut hin nhanh chng.
ngn chn s hnh thnh tng ri th phi b sung thm auxin ngoi sinh.
Ngi ta s dng cc cht iu ha sinh trng nh -NAA, GA, SADH cho cy.
Nng x l thch hp ph thuc vo tng loi cht v loi cy trng. ngn
chn giai on rng qu non ngi ta phun ln hoa hoc qu non ca nho dung
dch GA vi nng t 1- 20 ppm. i vi l phun -NAA vi nng 10 ppm hoc
SADH 1000 ppm u c hiu qu tt trong vic ngn chn s rng ca qu trc
v lc thu hoch. i vi to x l -NAA nng 20 ppm vo lc qu c biu hin
bt u rng th ko di thi gian tn ti ca qu trn cy thm mt s ngy na.
4.5. iu chnh thi gian ng ngh ca cc loi c, ht:
S ng ngh thng xy ra vi cc loi ht sau khi chn, cc loi c, cn hnh
cng nh cc chi ng. Nguyn nhn quyt nh s ng ngh l do cc cht c ch
sinh trng. Trong ht, c, chi ang ng ngh tch ly mt lng ln cht c ch
sinh trng m ch yu l axit abxixic, ng thi hm lng cht kch thch sinh
trng gim n mc ti thiu, c bit l gibberellin.

 ph b trng thi ng ngh, ngi ta s dng ch yu GA 3. GA3 khi xm

nhp vo cc c quan ang ng ngh s lm lch cn bng hormone thun li cho
s ny mm. Khi ht ny mm th qu trnh tng hp gibberellin din ra mnh,
gibberellin hot ha tng hp cc loi enzyme thy phn cn thit cho qu trnh ny
mm. V vy mun ht ny mm th tng hm lng gibberellin trong chng.
ph b trng thi ng ngh cho khoai ty thu hoch v ng trng v
xun bng cch x l GA 3 nng 2 ppm cho khoai ty mi thu hoch kt hp vi
xng hi hn hp rindit hoc CS 2 trong hm t kn s kch thch ny mm trong
thi gian t 5 - 7 ngy. Ngoi ra nu kt hp x l GA 3 vi x l nhit thp (4 10OC) th c kh nng ph b s ng ngh ca nhiu i tng khc nhau.
Trong kho bo qun, nhiu trng hp phi ko di thi gian ng ngh.
ko di thi gian ng ngh c khoai ty, ngi ta thng phun MH vi nng 200
- 500 ppm trc thu hoch. chng tp v chng ny mm ca cc loi c hnh,
ti trong bo qun, ngi ta c th x l IPCC (Izo - Propyl - Cloro - Carbamat) vi
nng 500 - 2000 ppm.
5.6. iu chnh s ra hoa ca cy:
Vic s dng cc cht iu ha sinh trng kch thch s ra hoa sm cng
l mt trong nhng ng dng ph bin v c hiu qu trong trng trt.
cho da ra hoa tri v lm tng thm mt v thu hoch, ngi ta phun NAA vi nng 25 ppm hoc b 1g t n (CaC2) ln nn da, khi gp ma
hoc ti nc t n s tc dng vi nc gii phng axetylen kch thch da ra
hoa. To, l, hng khi x l ADHS (Acid Dimetyl Hydrazid Sucxinic) nng 500 5.000 ppm c tc dng kch thch ra hoa sm v lm tng nng sut qu. i vi
u phun axit benzotiazon axetic nng 30 - 50 ppm s ra hoa nhiu, tng nng
sut qu. X l GA3 cho cy hai nm c th lm cho cy ra hoa vo nm u (x l
cho su ho, bp ci, x lch).
X l cc cht iu ha sinh trng tng s lng hoa v rt ngn thi
gian ra hoa ca mt s loi hoa v cy cnh. V d x l GA 3 cho cy hoa loa kn
vi nng 10 - 30 ppm lm cho cy ra hoa sm.
5.7. iu chnh gii tnh ca hoa:

Nhiu nghin cu cho thy vic s dng auxin s lm thay i t l gia

hoa c v hoa ci ca mt s loi cy. Nu s dng gibberellin s kch thch s
hnh thnh hoa c, s pht trin ca bao phn v ht phn. Cn nu s dng
xytokinin v ethrel s kch thch hnh thnh hoa ci. cy h bu b v cc cy n
tnh khc: s dng ethrel 50 - 250 ppm s to nn 100% hoa ci nn lm tng
nng sut ca cc cy h bu b. Trong vic sn xut ht lai F1 ca bu b, ngi
ta phun GA3 to cy mang hon ton hoa c v trng cy ch mang hoa ci
cnh cy hoa c v s to qu cho ht lai.
5.8. iu chnh s chn ca qu:
Trong thc tin sn xut, vic lm qu chn nhanh v chn ng lot thu
hoch c gii c ngha rt quan trng. Mt s cc loi qu khc nh chui, c
chua...thng thu hoch xanh d vn chuyn v bo qun c lu, v vy vic
iu khin qu chn ng lot, c mu sc p l cn thit. Cht c s dng ph
bin hin nay iu chnh s chn ca qu l ethrel dng dung dch, khi xm
nhp vo qu s b thy phn v gii phng ra etylen. Phun ethrel cho qu trc
khi thu hoch hai tun vi nng 500 - 5000 ppm s kch thch qu chn ng
lot. S dng ADHS 5000 ppm cng c hiu qu r rt ln s chn ca qu. X l
ethrel kch thch s chn ca nho vi nng 500 - 1000 ppm. Phun ethrel vi
nng 100 - 500 ppm cho h tiu vo thi k qu bt u chn s lm cho qu
chn nhanh. Phun ethrel vi nng 700 - 1400 ppm lm qu c ph chn sm hn
2 - 4 tun so vi khng x l. S dng ADHS vi nng 1000 - 5000 ppm xc
tin nhanh s chn ca qu o v anh o.
5.9. Nui cy m t bo:
Trong k thut nui cy m t bo th vic ng dng cc cht iu ha sinh
trng l ht sc quan trng. Hai nhm cht c s dng nhiu nht l auxin v
xytokinin. nhn nhanh invitro, trong giai on u cn phi iu khin m nui
cy pht sinh nhiu chi tng h s nhn. V vy ngi ta tng nng
xytokinin trong mi trng nui cy. to cy hon chnh ngi ta tch chi vo
cy trong mi trng c hm lng auxin cao kch thch ra r nhanh. Nh vy,
s cn bng auxin/xytokinin trong mi trng nui cy quy nh s pht sinh r hay
chi.

Cc cht thuc nhm auxin c s dng l IAA, -NAA v cc cht thuc

nhm xytokinin l kinetin, axit benzoic hoc ly t dung dch hu c nh nc da,
dch chit nm men...Ngoi cc cht kch thch sinh trng v dch hu c, cn b
sung thm cc hp cht nh ng, axt amin, lipt, mt s vitamin, cc nguyn t
a v vi lng vo mi trng nui cy.
Nng v t l ca cc cht kch thch ph thuc vo cc loi khc nhau,
cc giai on nui cy khc nhau...T l auxin/xytokinin cao th kch thch s ra r,
thp th kch thch s ra chi v trung bnh th hnh thnh m so (callus).
4.10. Cc cht iu ha sinh trng vi mc ch dit tr c di
(herbicid):
Cc cht diu ha sinh trng khi s dng vi nng rt cao cng c th
gy nn s hy dit. Cc cht nh 2,4D; 2,4,5T; MH... cng c s dng kh ph
bin vo mc ch dit c. Nguyn tc c bn khi s dng thuc tr c l phi quan
tm tnh chn lc ca thuc l ch dit cc loi c di m khng m khng nh
hng xu n cy trng.
Thuc phng tr c di c th chia lm hai nhm v c v hu c. Nhm hu
c li chia thnh hai nhm nh: nhm cc cht khng cha nit v nhm cc cht
cha nit.
Nhm cc cht khng cha nit thng l dn xut Cl ca axit phenoxyaxetic,
axit -phenoxypropionic, axit -phenoxybutyric, axit -phenoxyethyl. Cc i din
ca nhm ny nh: 2,4-D; 2,4,5-T; ACMP... Cc dn xut ca axit benzoic, axit
phenylaxetic nh: 2,3,6-ATB; 2,3,5,6- ATB... Nhm cc cht ny c nh hng
nghim trng ln qu trnh trao i chttrong cy v c ch hot tnh ca cc
enzyme lm cho qu trnh phn chia t bo trong m phn sinh v s sinh trng
gin ca t bo b ngng...
Nhm cc cht cha nit nh cc amit, cc dn xut ca urea, thicacbamat,
dithiocacbamat, dinitrophenol... Cc amit km hm enzyme cha nhm -SH, cc
dn xut ca urea km hm s c nh CO 2 v thi O2 ngoi sng, cc hp cht
cha nit khc cng vi phm n qu trnh quang hp v h hp ca cy. Cc loi
thuc tr c c s dng rt thnh cng cho mt s loi cy trng nh la m, la
mch, la go, cao lng, bng, c chua, u tng, c ci ng...

Ngi ta dng 2,4-D dit c hai l mm trong rung ng, phun trc hoc
sau khi c xut hin vi cc liu lng nh sau:
1- Mui Na ca 2,4-D:1,5 - 2 kg/ha.- Mui amn ca 2,4-D: 1,0 - 1,5 kg/ha
- Este ca 2,4-D: 0,8 - 1,0 kg/ha.
C th kt hp phun 2,4-D v 2,4,5-T vi mt s loi thuc tr c khc c
cha nit dit c g v cc loi c khc trong rung ng m khng nh hng
n cy trng.
Trong mt s trng hp vic duy tr tui th v hnh dng ca c li rt c
ngha. V d trong lnh vc trang tr, duy tr cc thm c trang tr cng vin
ngi ta thng phun cc dung dch km hm sinh trng. c bit l dng MH vi
liu lng 3-6 kg/ha lm km hm sinh trng ca c, duy tr thm c bn lu,
cng xn m li nng cao cht lng trang tr.

T I LIU THAM KHO

11. Nguyn Nh Khanh, 1996, Sinh l hc sinh trng v pht trin thc vt.
NXB Gio dc H Ni.
22. Phm nh Thi, Nguyn Duy Minh, Nguyn Lng Hng, 1987. Sinh l hc
thc vt, NXB Gio dc H Ni.

33. V Vn V, V Thanh Tm, Hong Minh Tn, 1999. Sinh l hc thc vt,
NXB Gio dc H Ni.
44. Bi Trang Vit, 1998, Sinh l thc vt i cng, NXB H quc gia Tp. H
Ch Minh.
55. Nguyn c Lng, L Th Thu Thu, 2006, Cng ngh t bo, NXB H
quc gia Tp. H Ch Minh.
66. TS. Dng Tn Nht, 2007, Cng ngh sinh hc thc vt -tp 1, NXB.Nng
Nghip.
77. Dng Cng Kin, 2002, Nui cy m thc vt, NXB H quc gia Tp. H
Ch Minh.

8 http://www.sigmaaldrich.com/
9 http://www.hoalanvietnam.com.vn/
10 http://www.plant-hormones.info