SINH L HC THC VT

NGUYN DU SANH

Ti liu tham kho chnh

1) Bi Trang Vit 2000 - Sinh L Thc Vt i Cng, Phn II. Pht trin. Nh xut bn i hc Quc gia Tp. H Ch minh, 349 trang 2) Bi Trang Vit 2002 - Sinh L Thc Vt i Cng, Phn I. Dinh dng. Nh xut bn i hc Quc gia Tp. H Ch minh, 333 trang 3) Campbell, N. A. (1996) Biology, 4th ed. Benjamin/Cummings, Menlo Park, CA. 4) Hopkins G., 1995 - Introduction to Plant Physiology. John Wiley & Sons Inc., New York, 464 p. 5) Lea, P.J and Leegood R.C., 1999 - Plant Biopchemestry and Molecular Biology. John Wiley & Sons Inc., New York, 363 p. 6) Leegood, R.C., Sharkey, T.D., and Susanne von Caemmerer., 2000- Photosynthesis: Physiology and Metabolism. Kluwer Academic Publishers, Dordrecht 7) Sage, R. F., and Monson, R.K., 1999 C4 Plant Biology. Academic Press 8) Frank B. Salisbury, F.B and Ross, C.W. 1992 - Plant Physiology. Wadsworth Publising Company, California 1992. 9) Taiz, L., and Zeiger, E., 1991 - Plant Physiology. The Benjamin/Cumming Publishing Company, Inc. California, 565p. 10) V Vn V, Hong c C, V Thanh Tm v Trn Vn Li 1993- Sinh l hc Thc Vt. Gio trnh Cao hc Nng nghip Sinh hc. NXB KHKT H Ni. 11) Plant Physiology online (http://www.plantphys.net/article.php?ch=e&id=282) 2 12) General Plant Biology (http://hcs.osu.edu/hcs300/)

M U

S sng khi u sau 1 t nm. Sinh vt u tin ch gm c protein v acid nucleic.

Ngy nay th gii sinh vt chia thnh 5 gii (kingdom): 1) Monera (gii vi khun = prokaryot) xut hin khong 3,5 t nm. 2) Protista (sinh vt n bo nhn tht : eukaryot) xut hin cch nay 1,5 t nm. 3) Fungi (gii nm) 4) Plantae (gii TV) 5) Animalia (gii ng vt) Tt c mi sinh vt u c chung 4 c trng c bn (trao i cht v nng lng, sinh trng v pht trin, sinh sn, cm ng v vn ng)

PHN LOI THC VT

Rong: sng trong nc a y: do rong v nm hp li i Thc vt (ru): sng ch m, c c quan sinh bo t r Khuyt TV: c r tht (cy), c mch, cha c c quan hoa (quyn b, mc tc, rng: dng x , ). Cy c hoa: Ht trn (lo t: thng, thin tu, tng bch) Ht kn (b t: xai, la, ... ) V TR PHN LOI Gii (Plantae) Ngnh (-phyta) Lp : (song t dip: Dicotyledonae hai l mm; n t dip: Monocotyledonae mt l mm) B (-ales) H (-aceae) Ging (Chi): 4 Loi: (thng hai t v vit in nghing)

CC KHI NIM C: B phn dinh dng di t v ph ra . THN: B phn mang l.

L: Do mt cung ( petiole) mang ly mt phin (limbe)

HOA Hoa: gm:L i, cnh hoa, nhy c (nh), nhy ci (nhy Pht hoa (hoa t): Nhiu hoa mc ghp li vi nhau. C nhiu kiu pht hoa: chm, gi, tn, tn phng, hoa u () , t tn. TRI Gm nhiu loi: qu mp (ph qu), qu nhn cng, tri kp, qu kh (gm b qu hay nang) HT: Cha phi v cht d tr nui phi.
5

I TNG NGHIN CU
Thc vt bc cao gm hai nhm: Song t dip (hai l mm- Dicotyledons):khong 200.000 loi + Gn l phn nhnh, hnh mng. + B mch thn sp xp trn mt vng (libe nm trn b mc) v c tng tng libe-mc + Thng c r cc (r ci taproot). + Hoa t hay ng phn. Ht cha phi vi 2 l mm

n t dip (mt l mm -Monocotyledon): khong 50.000 loi. + Gn l song song. + B mch thn sp xp trn nhiu vng ng tm, khng c tng tng libe-mc. + Thng c r chm (r si fibrous roots). 6 + Hoa tam phn. Ht cha phi vi mt l mm.

Sinh l thc vt l mn hc nghin cu v bn cht ca sinh hc thc vt.

Nghin cu sinh l thc vt l tm hiu nhng c ch bn trong c th (ni sinh) thc vt di tc ng ca cc yu t bn ngoi (ngoi sinh = mi trng sng) t chc li c th nhm tn ti v pht trin. Vic nghin cu th hin cc mc c th, c quan, t bo v di t bo. Thc vt c 3 hot ng c bn S hp thu v vn chuyn cc cht S bin dng S pht trin Nghin cu sinh l thc vt l nghin cu y c ba qu trnh v chng lin quan cht ch vi nhau.

CC MN HC KHC C LIN QUAN :

Hnh thi gii phu thc vt Sinh ha hc thc vt Sinh hc phn t Sinh thi hc thc vt Sinh l sinh thi thc vt Di truyn hc thc vt Trong thc t cc mn khoa hc ny rt gn gi, lin h cht ch v b sung cho nhau, kh tch bch.

NGHA CA VIC HC SINH L THC VT:

Hiu r hn v thc vt l sinh vt c tm quyt nh cho cuc sng trn tri t (sinh vt t dng). Hiu v vai tr ca thc vt trong chu trnh vt cht (sinh a ha) v chu trnh nng lng m bo cho s sng trn tri t. L c s cho vic trng trt, cng ngh sinh hc: tng nng sut, cht lng.
8

S LC V T CHC C TH THC VT
T bo thc vt v cc bo quan: T bo l ni hot ng bin dng chnh, nhn l trung tm iu khin. Mc t chc t: T bo m c quan c th.

PHN DINH DNG

CHNG I: S HP THU V VN CHUYN NC I- Nc v cc c tnh ca nc .Phn t nc Cu ni O-H to vi nhau mt gc 105. Cc trng thi ca nc
S ngng t

Hi
Lng

S bay hi
S ng bng

lng
rn

S tan chy
S thng hoa

Rn

S hnh thnh sng

hi
10

Cc c tnh ca nc Nc l dung mi hon ho Phn t nc nh v hu cc nn nc c th ha tan mt s ln cc cht, c bit l cc ion v cc cht tch in bng cch bao quanh v lm gim tng tc tnh in gia cc cht ny. Nc gip thc vt gim bt cc dao ng nhit V nc cn mt nhit lng tng i ln (t nhit v n nhit bc hi) ph v cu ni hydrogen v bc hi nn s thot hi nc l l cch tt nht lm gim dao ng nhit thc vt (lm mt). Nh ni hydrogen, nc c kh nng kt v bm rt mnh ct nc. Nc c sc cng b mt rt ln Nc to p sut thy tnh: Do sc cng trong nc kh ln nn ta phi cn mt lc ko kh mnh ph v ct nc (120 atm)
11

VAI TR CA NC TRONG I SNG THC VT

Nc l nhn t quan trng bc nht i vi tt c cc sinh vt sng. mc t bo: + To sc trng + Tham gia phn ng bin dng + Tham gia vo s tng trng v c ng t bo + L dung mi ha tan cc cht khong

mc c th: + Nc gip vn chuyn cc cht + Nc cn lm mt l

HM LNG NC V NHU CU NC CA CY Nhu cu nc ca cy rt ln v ty thuc vo c im sinh l, sinh thi, ty vo loi cy khc nhau. 12

Th nc
.Phng trnh th nc Phng trnh th nc: = - + P : p sut thm thu ca dung dch P: p sut thy tnh
.Trng thi nc ca t bo thc vt Nc t do & nc dnh. Nc di chuyn theo th nc, l qu trnh th ng. .Trng thi co nguyn sinh v trng nc ca t bo

Mi trng u trng (t bo co nguyn sinh)

Mi trng nhc trng (t bo phng ln)

13

Cc qu trnh vn chuyn nc trong thc vt

Mao dn Chiu cao ty thuc: lc ko ct nc i ln v trng lc. Khuych tn Khuych tn l s chuyn ng ca cc phn t theo khuynh nng (s chnh lch v nng ). Dng khi (c ch quan trng nht) Dng khi l qu trnh vn chuyn nc theo khuynh p sut. Thm thu L qu trnh khuych tn ca nc qua mt mng thm chn lc.

Mi trng u trng

Mi trng nhc trng

14 Mi trng ng trng

C ch hp thu v vn chuyn nc thc vt Nc t dch t ti b mt r

S di chuyn ca nc trong t ch yu theo c ch dng khi.

Cc con ng hp thu nc bi r : 3 con ng

-Con ng apoplast -Con ng symplast - Con ng qua mng (thng k chung vo symplast)
Cau lien bao Mang te bao

Bieu b

Vo

Noi b

Tru b

Vach te bao

Khung caspary
15

C ch hp thu v vn chuyn nc thc vt S di chuyn ct nc ln cao: do - Sc y ca r - S thot hi nc l (quan trng nht) - Lc kt hp gia cc phn t nc S thot nc l - S thot hi nc qua kh khu (QT) - S tit nc git qua thy khu * ngha ca s thot hi nc + M khu trao i CO2. + Vn chuyn cc cht dinh dng. + Lm mt l. + Lm t bo ng thng v tng trng
C CH NG M KH KHU

16

CHNG II: S HP THU V VN CHUYN CHT HA TAN

Cc nguyn t trong c th thc vt
I- Nhu cu v dinh dng khong
Trong cy hu nh c hu ht cc nguyn t. Ty vo hm lng trong cy, cc cht khang c xp thnh hai nhm: a v vi lng Cht khong l thut ng ch cc cht v c, dng ion, hin din trong thin nhin, thng sn sng trong t cho cy s dng.
Nhu cu v dinh dng khong
Tang trng thieu toi hao oc

Nong o chat khoang

17

II- Cc nguyn t cn thit v cc triu chng thiu khong Khi nim v nguyn t cn thit
- Cn thit cho s pht trin bnh thng ca vi loi thc vt. - Khng th thay th c. - Gy cc triu chng thiu c bit. Chn nguyn t a lng: C, H, O,N, S, P, K, Mg, Ca, By nguyn t vi lng: Fe, Mn, Cu, Zn, Mo, B, Cl

Vai tr tng qut ca cc nguyn t cn thit

Cc nguyn t a lng + to thnh cc hp cht hu c + To th thm thu cho t bo + D tr v trao i nng lng + Hot ha enzym Cc nguyn t vi lng L thnh phn ca enzyme hay coenzyme.
18

TRIU CHNG THIU NGUYN T CN THIT

N (Nitrogen) L b hong ha (thng l gi) hay c mu , cy chm tng trng, thn mnh v thng ha g. P (Phosphor) Cy non gim tng trng, l c mu xanh thm , cy km u tri, chm chn. Nng sut c thp. K (Potassium) L hong ha ( ngn v mp l), thn mnh v yu t vi nhng lng ngn bt thng. Nng sut c thp. S (Sulfur) Cy c triu chng nh thiu nitrogen. S hong ha xy ra trc cc l non. Ca (Calcium) C biu hin ca s thiu st (a vng), m b mm nhn, l non hp v cong xung. Mg (Magnesium) S hong ho xy ra trc l gi. Nng sut c thp.
19

Fe (st) S hong ha bt u cc l non. Cu (ng) L c mu lc sm, c th b xon hay bin dng. L non c cc vt hoi m (bt u t cht v lan xung mp l), v c th rng. B (bo) S phn chia t bo b cn, s hoi m xy ra l non, n hay c, tri v r ph to, cy mt u tnh ngn v phn nhnh nhiu. Mn (mangan) C s hong ha v s pht trin ca cc vt hoi m nh. Zn (km) Tng trng lng gim, l nh, vn vo, ba l nhn, hong ha l gi. Mo (molypden) C s hong ha v hoi m cc l gi, hoa rng sm hay khng thnh lp c. Cl (chlor) S hong ha v hoi m xy ra, ko theo s ho ca ngn l, l c mu ng 20 v tng trng chm, r dy ln vng gn ngn.

III- S cung cp cc cht dinh dng cho thc vt

- Cy c th hp thu cc cht khang qua thn, l, r ch yu l qua r. - Thc vt ly cht khang t mi trng di dng cc cht ha tan. Phi cung cp v ng lng cng nh lai nguyn t ty thuc vo mi giai an pht trin ca cy. Cn ch n pH ca dung dch ha tan.
IV- Cc kiu vn chuyn cht ha tan S vn chuyn n mch mc Theo ba con ng: apoplast, symplast v qua mng (symplast). S vn chuyn cc ion qua mng t bo C ch vn chuyn ch ng (cn nng lng) C ch vn chuyn th ng (khng cn nng lng)

V- C ch hp thu v vn chuyn trong mch mc - S phng thch ion vo m mc - S vn chuyn i ln ca nha nguyn trong mch mc - S thu nhn ion khong t dch mc
21

 Nhu cu N 1-3% nhng c ngha quan trng bc nht.

Thc vt ly m t t dng NH4+, dng cc nitrat (NO3-). Dng nit cung cp nh 6 ngun: - Qu trnh tng hp ha hc (rt t t sm st) -Qu trnh c nh nit ca vi khun, vi khun lam sng t do - Qu trnh c nh nit ca cc vi khun, to cng sinh - Ngun nit hu c t xc sinh vt - Ngun nit dng phn bn - Ngun m ng vt sng (t ch mt s loi) S hp thu N qua 2 dng: NO3- (nitrat) v NH4+ (ammon) Qu trnh amin ha: NO3- NH4+ Qu trnh ng ha nit amon trong thc vt Sinh tng hp protein S tng hp protein trong t bo c thc hin trn cc ribosome ca t bo 22 cht.

VI- S bin dng nit

CHNG III: QUANG HP

I- Cu trc b my quang hp v cc sc t Dip lp
Dip lp hnh trng, di 3-10 m, rng 1-4 m. II- C ch quang hp nh ngha quang hp 6 CO2 + 6 NADPH + H+ v 12ATP C6H12O6 + 6 H2O Quang hp c 2 giai on chnh: Pha sng: qu trnh quang gii nc. Pha ti: kh CO2 thnh glucid. H thng thu nh sng PS II (P680) v PS I (P700) Gm cp phn t dip lc t a phn t sc t ph phc hp thu nh sng (phc hp anten).

23

HAI PHA CA QU TRNH QUANG HP Pha sng: trn mng thylakoid S quang gii nc 4H2O 4OH- + 4H+ 4OH- 2H2O + O2 + 4eSau , c s chuyn in t qua 2 con ng: ng khng vng v ng vng Pha ti : trong stroma Kt np CO2 (chu trnh C3, PCR, Chu trnh Calvin) Gm 3 giai on chnh: - Carboxyl ha ribulose-1,5-biphosphat (RuBP) to phosphoglycerat (APG) - Kh APG - Ti sinh RuBP

24

III- Cc yu t nh hng n qu trnh quang hp nh sng Nhit Nng CO2 :ti ho khong 0,1%; >3% c ch Nc Dinh dng khong Cc yu t khc: tng nng O2, s hin din ca cc cht cn bin dng,
cc cht gy m (chloroform, ether), cht tr c (Diuron) lm cn mnh s quang hp.

25

IV- Bin dng carbon (cc kiu quang hp thc vt)

Chu trnh C3 (PCR - Photosynthesis Carbon Reaction) Quang h hp (photorespiration) L hin tng thoi bin ta CO2 nh h hp nhng xy ra trong cc t bo cha dip lc t di nh sng. Quang h hp gn lin vi chc nng c nh O2 (oxygenase) ca Rubisco khi kh quyn c nng O2 cao v nng CO2 thp (<50ppm) (im b = im b tr CO2). Chu trnh C4 CO2 c ng ha thnh cc acid C4 bi PEP carboxylase trong t bo tht l. Cc acid ny khuych tn vo t bo vng bao, kh carboxyl v phng thch CO2.
Chu trnh CAM Cc cy theo c ch ny thc hin m khu v nhn CO2 vo bui ti. Ban ngy, CO2 c nh t cc cht nhn v tham gia vo chu trnh C3PCR.
26

So snh chu trnh CAM vi chu trnh C4 Hai c ch rt ging nhau nhng c khc bit quan trng: cy C4, s thnh lp cc acid C4 v c nh CO2 bi chu trnh C3PCR tch bit nhau theo khng gian. S tch bit ny do cu trc c bit ca cy C4. cy CAM, hai qu trnh trn tch bit nhau theo thi gian.

HNH THI GII PHU L CY C3, C4

27

ac iem Hnh thai Phan bo Thng gap Ho Cau truc La Loai luc lap Cau truc Kranz (vong bao bo mach : BS) Chat nhan CO2 au tien San pham au tien Enzym thc hien Thu nap CO2 Nhiet o(oC) Anh sang bao hoa Quang ho hap iem bu CO2 Hieu suat quang hp (mg CO2/dm2/gi) Nhu cau nc (gr nc /gr chat kho) Kha nang tao chat kho Tai phan bo chat onng hoa

Thc vat C3 a dang Khap ni (80%) Nhieu Ho Cha nhieu te bao M 1 (diep mo: M) Khong RuPB (+ CO2) Acid C3 (PGA) RubisCO Ban ngay 10 25 trung bnh co 30-50 ppm 10-30 500 600 trung bnh trung bnh

Thc vat C4 Cay bui, co Nong am (13%) Hoa ban, Lac Te bao M xep 1-2 vong ong tam vi BS 2: M va BS Co PEP (+ HCO3) Acid C4 (OAA, Malic, Aspartatic) PEPC Ban ngay 25-45 khong co khong <10 ppm 40-60 250 300 cao nhanh

Thc vat CAM Cay mong nc Nong kho (7%)

Song i, Xng rong, Lan

Cha khong bao to 1 (diep mo: M) Khong PEP (+ HCO3) Acid Malic PEPC Ban em 30-45 rat cao khong < 5ppm 10-25 50 thap thay oi

28

S tng hp sucrose v tinh bt

Tng hp sucrose (saccharose)

Dihydroxyaceton-phosphat (t chu trnh C3PCR) trng hp to thnh fructose 1,6-diphosphat kh phosphoryl ha thnh fructose-6-phosphat isomer ha thnh glucose-1-phosphat. Glucose-1-phosphat (G-1-P) c hot ha bi UTP G-1-P + UTP UDP-G + Ppi(Pyrophosphat) UDP-G + fructose UDP + saccharose ( khng bo hay TBC)

Tng hp tinh bt
Trong dip lp = Xy ra trong stroma, qua fructose 6-phosphat (F-6-P) isomer ha thnh G-1-P c hot ha bi ATP. ATP + G-1-P ADPG + PPi [Glucose]n + ADPG [glucose]n+1 + ADP Enzyme tng hp tinh bt c hot ha bi K+, c cm ng bi cytokinin . Trong bt lp = cc t bo tch tr (sink cell) 29

CHNG IV: S VN CHUYN TRONG M LIBE

I- Khi nim v ngun (source) b (sink) Ngun (source ni cho) l ni sn xut cc cht ng ha nhiu hn l nhu cu s dng ca ni ny (l (trng thnh) l ngun ch yu) B (sink ni nhn) c hai lai b: tiu th & tch tr

II- Mt s khi nim lin quan n libe

Libe v cc t bo lin h Libe (t bo: km, nhu m) T bo truyn Tia

III- Tnh cht ca s chuyn v trong libe C bn tnh cht cn bn: - C t chc: xy ra trong libe. - Chn lc: chuyn cc cht hu c c ngun gc t quang hp. - nh hng: t vng cho n vng nhn. - Trt t: khng th b xo trn hay o ngc.

30

IV- S vn chuyn t nhu m vo mch sng

S chuyn triose-phosphat t dip lp vo t bo cht

Nh cc protein vn chuyn. S chuyn saccharose t t bo tht l ti yu t sng Ch yu theo con ng symplast (qua cu lin bo). S np vo libe (nhp sng = phloem loading) Saccharose c chuyn vo yu t sng theo m hnh ng vn chuyn. Cc cht khc nh acid amin v hormone thc vt vo ng sng bng cch khuych tn. V- S vn chuyn trong mch sng S vn chuyn trong mch sng l c ch th ng, c dn bi khuynh p sut gia vng cho v vng nhn. VI- S chuyn t mch sng ra nhu m (tho sng = phloem unloading) S chuyn saccharose t mch sng vo cc t bo nhn xy ra theo con ng symplast hay apoplast (ch yu). Trong qu trnh chuyn, saccharose 31 c ct thnh glucose v fructose nh invertase.

CHNG V: S H HP
I- nh ngha v vai tr ca h hp t bo
H hp t bo c ngha l trao i kh gia sinh vt v mi trng. Vai tr: 2 vai tr quan trng: - Phng thch nng lng. -Cung cp cc cht cn thit (sn carbon). Ba giai on lin tip ca s h hp t bo xy ra : - Cytosol (t bo cht): qu trnh glyco gii. - Matrix ca ty th: chu trnh Krebs (hay chu trnh acid citric; chu trnh TCA tricarboxylic acid) Mng trong ty th: s vn chuyn e- ti oxy ca khng kh.

II- Cu trc ti th Ti th l mt bo quan c dng hnh cu, hnh que hay hnh si di; c ng knh 0,5-1m, chiu di 1-5m.

32

III- S Glyco gii (con ng EMP Embden-Meyerhoff-Parnas)

2ADP 2ATP

Glucose
2NAD+

2 acid pyruvic
2NADH

IV- Chu trnh Krebs Pyruvat chuyn qua mng trong ti th Trong matrix, pyruvat gii phng CO2 AcetylCoA + cht nhn oxaloacetic (OAA) acid citric chu trnh Krebs. Qua mt lot phn ng trung gian, cht nhn (OAA) c ti sinh. 1 glucose 4 phn t ATP (2 do glyco gii, 2 do succinyl CoA) + 10 NADH + 2 FADH2. IV- Chui chuyn din t v sinh tng hp ATP Bt u t NADH, in t s ln lt i qua cc cht nhn n oxygen l cht nhn cui cng to nc.
33

V- Con ng pentose phosphat (PPP)

Hin din

Con ng pentose phosphat cng tham gia oxy ha glucose. Gia tng khi t bo thc vt chuyn t trng thi m phn sinh n trng thi m phn ha hn. Din tin 2 giai on: s oxy ha trc tip glucose-6-phosphat v ti sinh G-6-P. Vai tr Vai tr trong s bin dng thc vt: + NADPH sinh ra c dng cho cc phn ng kh trong cytosol, tham gia vo s bin dng nng lng t bo (to ATP). + Ribulose-5-P l tin cht ca ribose v deoxyribose.

34

CHNG VI: BIN DNG LIPID

I.1 S tn to ng VI.2 -oxyha VI.3 S ng ha cc cht dinh dng

35

PHN SINH TRNG V PHT TRIN

36

CHNG VI: PHT TRIN V CC BIU HIN CA S PHT TRIN

I- Chu trnh pht trin ca thc vt c ht
Chu trnh pht trin gm nhiu giai on k tip nhau. Ht cy mm cy con cy trng thnh hoa, tri, ht hoc c quan d tr lo suy. Chu trnh pht trin c th n bo (t bo) bt u t s phn bo ca t bo m v kt thc khi hai t bo con c to ra. Lo suy (senescene): Bao gm mt chui s kin bnh thng khng th o ngc s ph hy t chc t bo s cht ca thc vt. Lo ha (aging, vieillissement) l s thay i theo thi gian (khng lin quan g n s gi ci v s cht). Ranh gii gia chng khng r rng: Lo suy l trng thi sinh l sau cng ca s lo ha.
37

II- PHT TRIN V CC BIU HIN CA S PHT TRIN 1)- Pht trin (Development) L thut ng c dng ch nhng thay i ca c th thc vt theo thi gian hon thnh chu trnh pht trin ca n.
2)- Cc biu hin ca s pht trin: Pht trin = Phn chia (sinh sn) + gia tng kch thc + phn ha a)- S sinh sn: Ch yu xy ra cc vng sinh m (m phn sinh = meristem) * Ty theo v tr trong c th sinh m c phn chia thnh 3 loi: + Sinh m ngn + Sinh m lng + Sinh m bn * Theo thi gian xut hin, cc sinh m chia lm hai loi: + Sinh m s cp + Sinh m th cp Cc t bo trong vng sinh m c vch mng, nhn to, khng bo nh, kch 38 thc u (ng knh) lun trng thi phn chia.

b)- S gia tng kch thc: Chng lin quan n cu trc vch t bo (cu to bi pecto-cellulosid). S hnh thnh vch ring qua hai giai on: Vch s cp: Cc vi si cellulose to mt mng li c bao bi mt cht bt nho gm hemicellulose v pectic. Bt nho ny tng i mm nn vch s cp co gin c. S tng trng l do s tng hp thm cht vch. Vch th cp: cc lp mi s chng ln nhau k tip trn cc lp c. S tng trng vch th cp l do s dn thm vo (apposition) t bo khng ln thm, kch thc c nh. S gia tng kch thc t bo c hai kiu: # Tng trng nh # Tng trng khuch tn xy ra cc m c t bo ng nht c)- S phn ha: L qu trnh to nn cc c tnh chuyn bit v cu trc v chc nng, cc mc khc nhau.
39

CHNG VII: NG HC CA S TNG TRNG (C TH)

I- ng hc ca s tng trng
Ngi ta o s tng trng bng cc cch: 1. Gia tng bi hnh thi (di, rng, din tch, th tch) (khi tch) 2. Gia tng trng lng kh ti (khi lng) 3. L s gia tng nguyn sinh cht 4. L s phn chia s t bo 5. L s gia tng khi tch lin tc theo thi gian Thng thng, s gia tng v trng lng hoc gia tng v chiu di c dng ni v s tng trng. II- ng cong tng trng
S tang trng

Thi gian

+ Giai on u ca chu trnh di hay ngn ty thuc vo i sng ca thc vt. + Giai on ly tha. S tng trng cng t l vi s t bo ang phn chia hay ang ko di. + giai on cui c mt s chm tng40 trng

III- S pht trin c quan sinh dng:

1. R: R thng c phn chia thnh bn vng: vng chp r, vng sinh m cht, vng ko di, vng lng ht hay vng trng thnh. S sinh r (Rhizogense) khi s t mt s kh phn ha cc t bo ni ti, tip theo l s ti hot ng ging nh sinh m. 2. N (chi): Sinh m chi c hai vng: + Tunica ngoi phn chia thng gc (anticlinal) bao ph b mt s khi l v cc m nm ngoi mt. + Corpus pha trong, phn chia theo mi hng gip tng trng th tch to phn tr (li) ca thn hay nhnh. 3. Thn chnh v cnh ph: Thn chnh xut pht t n cht, cn cnh ph (nhnh) bt u t n nch hay n bt nh. Vng ko di l vng pha di n (p cht 4. L: l c mt s tng trng b mt bi mt vng khng c ranh gii r nh 41 nhnh. Cung l tng trng ging nh nhnh.

IV- S PHT TRIN C TH

mc ton c th, pht trin thng ch thc vt vo giai on trng thnh biu hin bng s to hoa. 1. S tng hoa v nhp pht trin: S tng hoa l mt hin tng c bit phc tp chu nh hng ca cc yu t bn ngoi v bn trong c th. Cu trc ca mt n hoa xut pht t mt vng sinh m (sinh m ch). 2. Nhp iu pht trin: S hin hin ca hoa c th chia thc vt thnh 3 nhm ln: + Cy nht nin (mt nm, hng nin): + Cy lng nin (hai nm): + Cy a nin:

42

CHNG VIII: CC YU T NH HNG LN S PHT TRIN

I- Cc yu t nh hng ln s pht trin

1. Nhit Mi loi thc vt c mt nhit giai ring tng trng v pht trin. Nhit nh hng ln c nng cc qu trnh quang hp, h hp, phn ng bin dng, dinh dng nc, mui khong, thot hi nc, di chuyn nha nh hng ln tng trng ng biu din s nh hng ca nhit tng t nh cc hot ng sng khc. Nhit k tnh (thermoperiodism) (Went 1964) L tnh nhy cm ca thc vt i vi nh k ca nhit trong ngy m hay trong nm v nhng phn ng m nh k ny to ra. Went ghi nhn nhit m thp hn nhit ngy th c li cho s tng trng.

43

2. nh sng nh hng ln qu trnh quang hp v ln s pht trin ca thc vt. Quang k tnh l tnh p ng li cc nh k chiu sng thay i trong ngy (ngy v m) hay trong nm. thc vt, quang k tnh c nh du r bng hin tng to hoa. Quang k tnh cn nh hng trn s dinh dng. 3. Nc nh hng ca nc cn ty thuc vo giai on pht trin ca thc vt. Nc nh hng trn s tng trng nhiu hn trn s pht trin. 4. Kh Cc kh c nh HCN, H2S, CO2, khi thuc l. nh hng ln s h hp dn n ri lon bin dng v lm cht thc vt. 5. Thc phm (m, hu c, khong) Cn thit tng hp mau l cc protein trong t bo sinh m.
44

II- MI TRNG NG HNG (ISOTROPE) S p ng vi cc yu t mi trng lm cho thc vt c 1 nhp iu. 1. Nhp iu sinh hc (nhp ngoi sinh) Nhp iu sinh hc l kt qu ca cc thay i trong c th nh k to ra do mi trng nh hng ln s tng trng nhp ngoi sinh. Nhp ngoi sinh thng thy c nh hng r bi hai nhp iu: - Nhp iu ngy: vi mt nh k 24 gi, lun phin ngy v m. -Nhp iu nm: vi mt nh k 4 ma (hoc 2 ma).

2. Nhp ni sinh L cc thay i do cc yu t bn trong gy ra. Gi l s iu ha sinh hc. C th thc vt c sn mt ng h sinh hc (cy c tr nh).

K1 A B

K2 C
45

3. Nhp iu tng trng Nhp iu hng nm: r cc x c 4 ma. * i vi cy nht nin: tng trng mnh cho n khi tr hoa chm ngng tng trng. * i vi cy lng nin: Cy tng trng mnh vo ma xun to b phn dinh dng, gim tng trng vo ma h, ti lp tng trng ma thu to cht d tr thn, r, c v ngng tng trng vo ma ng. * i vi cy a nin: Tng i phc tp. Nhp iu trong thi gian ngn (nhp ngy) Khng hin r nh nhp iu hng nm cn phi o v ghi nhn k. Thng thng trong ngy: Thc vt c dip lc t: Cy quang hp c mt nhp ngoi sinh ti a vo ban ngy, ti thiu vo ban m.
46

4. Pht sinh hnh thi Nhp iu tng th (Rhythmes morphogntiques): Nhp iu iu khin s thnh lp c quan mi. Nhp iu ny c bit rt t. 5. Tng quan tng th Trong mt c th, s pht trin ca mt c quan khng nhng ty thuc vo iu kin ngoi cnh hay nhng tim lc bn trong cc m m cn ty thuc vo hot ng ca cc c quan khc. gi l s tng quan ca hai c quan. S tng quan tng th do hai yu t chnh nh hng: - Mt nhm c bn cht dinh dng: - Mt nhm c bn cht hormon (cht iu ha)

47

III MI TRNG KHNG NG HNG (ANISOHOPE) C mt s tc ng khng ng u ln thc vt ca mt yu t no . 1. Hng ng (tropism) Hng ng l phn ng i hng ca c quan di nh hng ca mi trng khng ng hng. C ng ny khng hi phc. 2. ng ng: L nhng thay i hng ca c quan do s thay i cu trc gii phu ca c quan nui dng n ch khng phi do mi trng bt ng hng. Hin tng ny c hi phc.

48

CHNG IX: CC HNG NG V C NG CA THC VT

Thc vt cng c cc c ng v cc c ng ny kh nhn bit hn ng vt. Trong t nhin, thc vt c 2 loi c ng (Movement): Hng ng (tropism) l mt c ng m hng kch thch ca mi trng xc nh hng ca s vn ng. y l c ng khng hi phc ng ng (nastic movement) thng c khi mo bi mt kch thch ngoi sinh. Hng ca s kch thch khng xc nh hng ca s vn ng. C ng ny hi phc li khi mt tc nhn cm ng. Mt c ng ca thc vt tun theo c ch p ng ba bc: .Nhn tin (perception): .Truyn tin (transduction): .p tin (response):
49

I- Hng ng
Khi c s chiu sng khng ng u + l c quang hng ng nghing (plagiotropism). + r c hin tng quang hng ng nghch c tnh: S cong do quang hng l mt qa trnh tng trng khng u hai bn mt c th, vng ko di di cht. 2. a hng ng: L s cong ca c quan di tc dng ca trng lc. R c a hng ng dng thn, l c hin tng a hng ng m 3. Cc hng ng khc Ha hng ng 1. Quang hng ng (phototropism) Xc hng ng
Kt lun: Cc hng ng gip thc vt i ph vi mi trng rt hu hiu mc d chng khng di chuyn c nh ng vt. Ngoi ra, cc 50 hng ng gip cho thc vt c cu trc ring.

II- C NG NI BO C ng ni bo do s di chuyn ca protein cu trc (l nhng vi si hay vi ng) gi l tubulin. Cc protein ny co gin do s polymer v kh polymer y l s c ng cn nng lng. III- NG NG( NASTIC MOVEMENT) L v l th cp ( l kp) thng c c ng ng ng. 1.C ng thc ng ( c ng ban m - nyctinasty): y l s c ng theo mt nhp iu c kim sot bi yu t mi trng v ng h sinh hc. L kp l c quan d hc hin tng ny. C ch: lin quan n nng K+ bn trong v nng H+ trn mng cc t bo co v dui. 2. ng ng kh m (thy ng ng-hydronasty) Thy ng ng c lin h n s cun li hay khp li ca l do m thp, l cun li gim thot hi nc. Do s mt i p sut trng ca t bo vch mng (t bo hnh bt bulliform 51 cell) thng gp n t dip.

3. Xc ng ng (ng ng tip xc thigmonasty) c bit gp vi thnh vin trong h ph mc c (Leguminosae) ca h u. C ch: Do s mt nc ca t bo th gi (c lin h n s vn chuyn K+). S truyn tn hiu lm khp l xy ra do hai c ch: c ch in (electrical mechanism) v c ch ha hc (chemical mechanism). * Cy bt rui ( The Venus fly trap) C ch: c ch acid (bm H+ mng t bo). 4. C ng cun ngoi (revolutif) Cht ngn ca dy leo c mt c ng cun ngoi to mt vng xon trong khng gian tm vt ch. C ch ca c ng cun ngoi: Do nhp ni sinh qua trung gian ca a hng ng C s thot nc do bm proton t di theo vch. C gi thuyt l auxin kch thch tng trng chi ngn ko di.
52

5. Nhng c ng khc: C ng ca dip lp: C s sp xp ty theo s chiu sng Cc bin i trng nc Do c ng ca nc t khng bo lm ng m khu, h xung hoc ging ln ca mt vi c quan (nh c, l)

Bin i do lng nc cha trong vch

Bin i ny theo m ca mi trng. To cc bin hnh c hc ca t bo cht nh lm m nang bo t, m ti phn hoc s nt ca cc qu.

Di chuyn c th, cm xc ng ng
Xy ra cc c th n bo.

53

CHNG X: KIM SOT S TNG TRNG BI CC HORMONE

THC VT
I- HORMONE THC VT (CHT IU HA SINH TRNG THC VT) V LCH S PHT HIN Duhamel du Monceau (gia TK 18) Sachs (1880) : cc thng tin ha hc (chemical mesengers) cc Hormon thc vt hay cht iu ha sinh trng thc vt.

Ni yu t l cc cht do thc vt t tng hp nn chng c vai tr iu ha s pht trin. Cc ni yu t c th k nh: + Cc sc t + Cc cht ph tr cho s tng trng: sorbitol, inositol, manitol + Cc vitamin + Cc cht iu ha sinh trng (hormone)

54

LCH S CA CHT IU HA SINH TRNG Tip theo ghi nhn ca Sachs: - 1880, Darwin nghin cu quang hng ng ca Phalaris canariensis. - Boysen Jensen (Thy in) nm 1913: chm mt ming gelatin gia ngn v phn di dip tiu, khi chiu sng, dip tiu cong. - Paal, 1919: s cong l do cht ha hc di chuyn khng u t ngn - Went 1926: ly trch c cht ny. ng gi cht ny l auxin (ting Hy Lp auxin c ngha l s tng trng). - Nm 1948, Thimann pht biu Thc vt cng c kch thch t nh ng vt. Kch thch t thc vt l mt cht hu c tng hp mt ni trong c th sng, c nh hng n mt ni khc cch xa ni n c to ra. Cc hormone khc ln lt c pht hin sau : Gibberellin (Kurosawa 1926), Cytokinin (Skoog et al. 1950). l cc cht kch thch tng trng . Bn cnh , trong thc vt cng gp nhiu cht c ch tng trng. Cc cht ny thng thuc nhm phenol (vnari nm1949), Acid Abscisic (Addicott v csv. 1961), Ethylene (Cousins 1910).
55

- nh ngha Kch thch t thc vt l nhng cht hu c do thc vt tng hp mt ni ri di chuyn n ni khc, chng hot ng vi lng rt nh. Chng c chia lm hai nhm: + Kch thch t tng trng (iu kin sinh trng) + Kch thch t tr hoa (iu kin tr hoa) Sau ny gi tn l Cht iu ha sinh trng thc vt l nhng cht hu c, c th km hm hoc lm thay i mt trong cc bc ca chu trnh pht trin, tc dng vi mt lng rt nh, c th l cht iu ha sinh trng hoc iu ha tr hoa. Nm 1984 hi ngh ti Pratislava Tip Khc b sung nh ngha. Cht iu ha sinh trng thc vt l mt cht hu c cn thit cho qu trnh pht trin thc vt. Chng hot ng vi mt lng rt nh. Chng nh hng trn cc bc ca qu trnh pht trin, chng l cht cn thit nhng khng phi l cht dinh dng Hin nay: chia lm nm nhm: Auxin, Giberelin, Cytokinin, Acid Abcisic, 56 Ethylen

II- CC NHM CHT IU HA SINH TRNG THC VT 1. Auxin CH2 - COOH Lch s Do Went tm ra Auxin c tc dng ko di t bo ngn thc vt. N l acid indol acetic (AIA, IAA). H Hin din Auxin hin din hu ht thc vt bc cao, hin din nhiu c quan ang tng trng. Auxin hin din hai dng: + T do hay dng hot ng +Dng buc, dnh, kt hp Ly trch, o auxin Thng ly trch auxin trong dung mi hu c Auxin khuych tn qua lp agar. Nhc im: lm h ni ct v cht khng auxin cng c th khuych tn qua.
57

- Xc nh hot tnh ca auxin: Sinh trc nghim: + o cong dip tiu: + o s tng trng trc tip: Phng php ha hc Salkowski: Do FeCl3 to mu hng vi nhn Indol, xc nh mt quang bng quang ph k. Ngoi ra cn c th dng phng php sc k lng cao p khi ph (HPLCMS) hay phng php min dch hc Sinh tng hp T trypophan acid indol-3-pyruvic indol-3-acetaldehyde AIA + S chuyn amin ha thc hin bi transaminase + Kh CO2 ca Indol pyruvic thnh Indol acetaldehyde + Oxy ha bi NAD+ iu kin: nh sng yu v c ion Zn2+ .

58

Di chuyn Di chuyn t ni tng hp (sinh m ngn) n ni nhn. Auxin di chuyn theo hai li:(1) hot ng (cn nng lng) v (2) th ng (khng cn nng lng).

Thoi ha Auxin rt d b oxyt ha. Trong c th c AIA-oxydase (enzym peroxydase) l enzym ch yu phn hy auxin. Tc dng ca auxin: * mc c th: Auxin tc dng trn s ko di t bo cc m cn non. * mc c quan - thn v chi: auxin nh hng trn vng p cht (vng ko di). - l: phin l song t dip b c ch cn thn, cung l c kch thch. - Tng trng tng pht sinh.
59

- Tng trng qu b - Trn cc hng ng: - Trn s rng (l, hoa , tri): Auxin lm ngn cn s rng. Mt khi vng rng thnh lp, auxin kch thch s rng xy ra nhanh hn. - Kch thch to r bn v r bt nh: * mc t bo -Lm tng tnh dn ca vch t bo - Trn s tng di v tng rng t bo

60

Cch s dng: Thng s dng auxin dng dung dch, dng mui. pH c nh hng trn ha tan v trn hot tnh ca auxin, thng dng dung dch m c pH 4,5 - 6,5. Auxin tc ng vi mt lng rt thp tnh bng phn triu (ppm) thng thm cht ph gia vo phn phi u. Thng x l auxin bng cch phun, ngm, chch hay bi.

Phi s dng auxin ng liu lng, ng thi gian, ng giai an. Auxin c s dng trong nhn ging thc vt (nui cy m, gim cnh, trong s ra hoa khm, kch thch m cao su, s tng trng tri, chng rng hoa v l... c bit c s dng nhiu nht dit c di trong nng nghip.

61

GIBERELINS ( Gb) Lch s: - Nm 1926, Kurosawa li trch Gibberellin t nm. - Nm 1930, ghi nhn cht trch t nm khng nhng ch kch thch tng di la m cn trn nhng thc vt khc. - Nm 1936 Yabuta li trch c tinh th gi l Giberelin A. - Nm 1954 Brian Cross (Anh) v Stodola (M) tm ra cng thc n l mt acid: Acid Gibberellic (GA) (slide). - 1954, Takahashi v Tamura phn lp c 3 giberelin t Giberelin A. Chng c t tn l Giberelin A1(GA1), GA2, GA3. GA3 l acid giberelic. - Nhng nm gn y tm thy c 126 cht ging giberelin Trong c th thc vt cng lc hin din nhiu Gb, nhng khng phi Gb no cng c hot tnh. Gb kch thch ko di t bo. V cu trc c hai nhm Gb: nhm c 20 C v c 19 C. Cc Gb khc nhau ch yu do s thay i nhm OH C2, C3 hoc C5 v v tr gn ca chng dng hay .
62

Hin din Gb c tm thy c thc vt bc thp ln thc vt bc cao, gn y cng ghi nhn vi khun. Trong c th, ty theo c quan v giai on pht trin m Gb phn b vi nhiu loi liu lng khc nhau. Gb cng c th hai dng t do v dnh (lin kt). n nm 1992 (Salisbury &Ross) ghi nhn c 84 GA c pht hin Li trch o Ly trch bng cc dung mi hu c . iu kin ly trch t nghim nht hn auxin. Sinh trc nghim Dng cc thc vt ln do t bin lm mt gen sinh tng hp Gb nh bp ln, u ln, la ln..., hay trn hot tnh amylase trong s ny mm ht m cc, hay trn s tng di cy mm da chut, cy mm ci salad... sinh trc nghim hot tnh Gb.
63

Sinh tng hp Gb l nhng terpenoid c 20 carbon l polymer ca 4 n v isopren (5C). Xut pht t Acetyl CoA acid mevalonic (6C) phosphoryl ha v kh carboxyl isopentenyl pyrophosphat (5C) trng hp to gerany pyrophosphat (10C) hay geranyl-geranyl pyrophosphat (20C). Geranyl-geranyl pyrophosphat ng vng to ent- kauren nhm methyl C19 c oxy-ha thnh carboxylic, vng B co rt li cn 5C GA12 aldehid cc Gb khc nhau. Tt c cc ha cht lm ln hay lm chm tng trng u c tc dng ngn chn sinh tng hp Gb. Tt c Gb u c sn ent-giberelan.

thc vt con ng bin i gia cc Gb c nghi nh sau GA12 aldehid (20C) oxy ho C7 GA12 OH ha C3 hay C13 hoc c hai. t bo, s tng hp sn kauren t acid mevalonic c th xy ra trong cc lp, c bit l hong lp (etioplaste).
64

Di chuyn Gb di chuyn khp mi ni trong c th qua mch mc, libe v t bo. Thng di chuyn th ng theo cht ng ha, hay theo dng nha nguyn. Gb c th thot ra ngoi qua hin tng nc git. Thoi ha Cha c nghin cu k, c th kt hp vi cht khc tr thnh trng thi khng hot ng.

65

Tc dng sinh l * mc c quan + Kch thch lng ko di. + Gb kch thch tng trng l. + Gb cng kch thch tng trng tri + Gb gi tr hoa cho nhng thc vt cn th hn hay nhng thc vt ngy di cn to hoa trong iu kin ngy ngn. + Gb cng bin i hoa lng tnh thnh hoa c, kch thch to hoa c + Gb gip ny mm, ph v min trng. + Gb kch thch tng trng r * mc t bo + Gb lm tng lng auxin ca cc m bng cch tc ng trn s tng hp protease phn gii protein c cha tryptophan l tin cht ca auxin. Mt khc, Gb gip tng hp acid polyhydroxy cinnamic, cht ny cn hot ng ca enzym AIA -oxydaz phn hy auxin. + Gb iu ha biu hin gen
66

CYTOKININ

Lch s - Haberlant (1913) ghi nhn c mt cht di chuyn trong m libe, cm ng s phn chia t bo trong nhu m khoai ty. - 1939, Chiball ghi nhn trong qu trnh tng hp protein c mt cht xut pht t r lm cho l khng gi ( nhnh c r mc, l c gi lu) - Skoog (1949) vi phng php nui cy m thn cy thuc l ghi nhn + ty thn khng tng trng c khi khng c AIA + Khi c AIA, t bo tng rng nhng khng phn chia + T bo ch phn chia sau khi m mch c thnh lp v tip xc vi nhu m ty. Nh vy, m mch phi c cht g cm ng phn chia t bo. - Nm 1954, Miller ly trch t tinh trng c mi mt cht kch thch phn chia t bo v xc nh cu trc ca n. Sau , khi ly trch trong nhiu thc vt v cht ny c tnh cht phn chia t bo (cytokinesis) nn gi l cytokinin.
67

.Hin din Trong tt c cc thc vt v ni ang c s phn bo mnh Cytokinin hai dng: dng t do v dng dnh vi ARNt. .Ly trch v o Dung mi hu c (alcool, aceton, butanol) vi pH thch hp .Sinh tng hp Ni tng hp : cht r (vng sinh m). .Di chuyn Theo hai chiu c mch mc v libe nhng nhiu nht l qua m mc. C s di chuyn th ng v hot ng (cn O2) .Thoi ha c bit rt t.
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.Tc dng * mc c quan + G min trng chi, ht + To n mi, cn s lo ha, cn s rng l + Lm tng dy l (kch thch phn chia t bo) + Kch thch s thnh lp c, gip tch tr tinh bt c. * mc t bo Cytokinin kch thch s phn chia t bo v phn ha t bo. * mc phn t Cha c bit r. C vai tr trong s nhn bit codon thch hp, iu ha s sinh tng hp protein. .Tng hp Do cng thc tng i t phc tp nn tng hp c nhiu loi .ng dng Trong nui cy m, trong gim cnh, kch thch to hoa ci. Trong tng trng tri, lm tri lu chn, chng rng

69

ACID ABCISIC (ABA) Cht gy rng l non (Abscission) .Lch s - Addicott (1961) chng minh c mt cht lm rng l v ng t tn l Abscisine. - Dukuma (1965) ly trch cht , o v tm c cng thc. - Nhm ca Wareing (1964) cng ly trch c vi hot tnh cao nht mt cht c chi min trng ca cy Acer v t tn l Dormin (t dormancy). - Sau , nhiu tc gi cng ghi nhn l gp nhiu thc vt khc. ABA (acid 3 methyl trans pentadienoic)
CH3 CH3 CH OH O CH3 CH CH3 C COOH

CH

70

.Hin din Gp mi loi thc vt , ABA hin din nhiu m lo v trng thnh hoc hoa tri non, ht. Lng trung bnh 0,01 1ppm Thng hai dng t do v dnh vi glucoid.

.Ly trch o Nh cht auxin: dng sc k kh, trc nghim min dch, sinh trc nghim ( dip tiu, cy mm) .Sinh tng hp Ni tng hp: l trng thnh Tin cht: acid mevalonic qua trung gian carotenoid. .Di chuyn Di chuyn theo hai hng mch mc v mch libe.
.Thoi ha Cha r, c th dng dnh.

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.Tc dng: * Mc c th: - Gy s lo, lm hoa, tri, n b rng - Lm mt mu xanh (bin i dip lc t lm 4 nhn pyrol khng ng vng c) - Gy min trng chi, ht -Lm chm s di chuyn qua mch libe. -Lm ng khu, gip thc vt chng chu hn, cn s thot hi nc. * mc t bo L mt anti Giberelin c tc dng cn phn chia t bo. * mc phn t Cha r. C l ABA c ch tng hp ADN, ARN.

.Tng quan - c ch tng hp ethylen - Cn phn bo, cn ko di t bo, cn s vn chuyn cc cht d tr . Ap dng: (cha nhiu)
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.Lch s c bit t lu. - Giardin (1864) nhn thy cy b rng l do t - Werner ghi nhn trn cy mm u khng theo quang hng ng thun khi ny mm. - Nm 1901, ethylen c coi nh l mt yu t ca hi c. - Nelsubov (1913) ghi nhn chnh cy mm u khng mc thng ng l do kh hi trong phng th nghim (khi ma ng dng l si). Nu kh ny c kh c bi oxyt ng s khng c mm u cong. Ong chng minh acetylen, ethylen vi nng 0,06 ppm lm cho mm u mc ngang. - Sau , trong qu trnh vn chuyn tri (cam, chanh), ngi ta pht hin c nhng l chn nhanh. Quan st k th hin tng ny do nm mc to ethylen thc hi s chn (ghi nhn California). - Game (1934) gi c ethylen v xc nh c lng Ethylen to ra tri to.
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ETHYLEN (C2H4)

.Hin din Ethylen hin din mi thc vt, mi ni trong c th v c nng tng i thp (50l/kg). Thng gp khi c vt thng, hoa rng, tri chn, c quan lo

.Ly trch v o Ly trch dng sc k kh. L mt cht hi nn kh o, dng php sc k kh khi ph .Sinh tng hp Ni tng hp: ty th gi, c quan lo.
.Di chuyn L mt cht hi, s di chuyn khng t ra. Ethylen thot ra b c ch bi nhit thp v nng oxy thp. .Thoi ha To thnh CO2

74

.Tc dng * mc c th - Thc y s chn tri, s lo lm rng tri, l, hoa. - Lm mt tnh hng ng - G min trng chi, ht - Lm tri mm, i mu -Trong ti, Ethylen cn s ko di nhng kch thch ph to ra. - Kch thch to hoa, mt vi cy kch thch to hoa ci. * mc t bo Ethylen nh hng trn tnh thm ca mng, cc cu t d b phn hy. * mc phn t Cha r v y l mt cht hi, n gin. Ethylen gip sinh tng hp cellulase, polygalacturonase, mARN.

75

.Tng hp Thng dng acetylen (C2H2), propylen c hot tnh ko di hn Ethrel (acid 2-chloro- ethan phosphoric) nh ethylen t t. .Tng quan (auxin giberelin cytokinin ABA) Ethylen c lin quan n auxin v cytokinin (trong s tng hp). Ethylen khi to ra lm hi s chuyn ch auxin. Kch thch tng hp ABA. nh hng trn s phn bo, khng cn tnh bn thm mng. . ng dng: Kch thch to m cao su Ra bng ng lot mt s loi TV Chn tri. Cn ethylen bng cch gim nng oxy, bo qun nhit thp.

76

CC NI YU T KHC: Vitamin l mt hp cht hu c c thc vt tng hp thc hin cc phn ng sinh ha trong c th thc vt. V d: vitamin B1, B6, B5 cn cho r Acid amin l mt nhu cu khng th thiu i vi ng vt. thc vt c th t tng hp acid amin.

77

CHNG XI: KIM SOT S RA HOA BI CC HORMONE THC

VT
S chuyn hng t sinh dng sang pht trin l s to hoa. S to hoa bao gm s tng v n hoa.

I- Hin tng hnh thi

Khi c s cm ng: cc t bo vng cht ngn hot ng, chi dinh dng tr thnh chi hoa; sinh m ch hot ng. Lp tunicar bin i to: + tin sinh m bo t nhy v bu non. + tin sau cnh hoa phin hoa, cnh hoa. Sinh m sn hot ng mnh to s ko di trc hoa (cung hoa).
78

II- Nhng yu t ca s tng hoa

1. Ngoi yu t: Yu t dinh dng + Vic bn phn phi nghin cu theo i tng. + T l C/N c lin quan n cht to s tng hoa. + Yu t no lm ngn cn s tng trng nhnh cui giai on u nin th cht s thc hi s to hoa. NC Cn thit cho dinh dng v pht trin ca thc vt. NHIT + nhit thp, c s th hn tr hoa. + C mt cht xut pht t chi di chuyn qua vng thp kch thch gy cm ng tr hoa. Cht ny c ly trch v gi l vernalin. iu kin th hn * Nhit x l: 0-10C cho la m, 9-17C cho a s thc vt khc Thi gian x l: thay i ty theo loi t 4 ngy n 8 tun. * Cn Oxy ( hot ng h hp) 79 * Khng c nhit cao qu sau th hn.

NH SNG Quang k: l thi gian chiu sng trong mt ngy. Lch s: Nm 1920, Garner v Allard xng ra nguyn tc quang k. chia thc vt lm ba loi cy: cy ngy di, cy ngy ngn v cy phim nh. Yu cu ca quang k Tui ca thc vt: Thc vt phi tri qua mt giai on u nin mi cm ng c. S quang k cm ng Quang k cm ng l quang k kch thch tr hoa, khi cy tr hoa ri th khng cn duy tr na. Cy phi c gi trong cc iu kin quang k c nh trong mt hoc nhiu chu l lin tip mi tr hoa c. Quang k ty thuc thc vt iu kin: D k phi lin tc (nghim nht).
80

Cht nhn nh sng - Hendricks v Bordrick (1932) ghi nhn tia 660nm v 730nm c tc ng ti a. - Nm 1932, Flink v Alister cng ghi nhn ht salad mun ln mm phi chiu tia 660nm cn tia 730 nm cn ln mm. - Nm 1946, Hendricks ghi nhn trn Xanthium cng c hin tng tng t: 730 nm kch thch tr hoa, 660 nm cn tr hoa. - Nm 1966, ngi ta ly trch c cht nhn tia sng, l phytochrom (P). Phytocrom hai dng: + Pr thu nhn tia R (660 nm) v i ngay thnh Pfr + Pfr thu nhn tia FR (730 nm) v i thnh Pr vi vn tc chm hn. Pr : dng thm lng (bt hot) Pfr: dng hot ng (cn tr hoa) Phn ng 1 (Pr Pfr) l phn ng quang ha nghim nht (khng b nh hng bi nhit ) Phn ng 2 (Pfr Pr) xy ra chm, c l l nh enzyme.
81

Ni yu t (cht iu ha sinh trng thc vt) Qua cc th nghim, nhn thy cht to hoa l mt cht c th di chuyn c, i qua c quan c quan (l ln thn) v qua c th (cy qua cy). Bn cht ca cht v tm quan trng ca n cha c bit r. Bn cht ca cht iu ha Chailakhyan nu gi thuyt rng bn cht ca cht iu ha sinh trng cm ng to hoa c hai thnh phn l Giberelin v Anthesin. Cht ny c gi chung l Florigen. Theo thuyt hormone ra hoa: Florigen gm hai thnh phn: + Giberelin tng hp trong iu kin ngy di + Anthesin tng hp trong iu kin ngy ngn Ngc li, ngy ngn cn tr s tng hp Giberelin cn ngy di cn tr tng hp Anthesin. Vai tr ca Giberelin l ko di trc hoa m khng c nh hng trn s thnh lp hoa nhng cy ngy ngn; trong khi iu kin ngy di li tr hoa. Vai tr ca Anthesin kch thch s phn ha n hoa.
82

Ti sao cy u nin khng ra hoa? Gi thit 1: do cy khng Florigen trong thnh phn. Gi thit 2: tuy c Florigen nhng sinh m ch cha cm ng. Gii p: * i vi gi thit 1: cy n nin khng c s to thnh Florigen. Khi ly trch Florigen cho vo, khng nhn c cm ng. Khi cy gn giai on trng thnh, kh nng hp thu cao, cy mi tr hoa c. * i vi gi thit 2: Nui cy trn mi trng dinh dng, ngi ta ghi nhn: Cy trng thnh: Nng saccharose 0,7% cy to 100% n dinh dng Nng saccharose 3,5% cy to 17% n dinh dng, 83% n hoa Cy con: bt k nng dinh dng no, cy cng to 100% n dinh dng. Nh vy, cy cn giai on u nin cht cn thit cho s bin i to hoa.

83

.Cht cn ra hoa nhiu loi thc vt, ngoi vic cm ng quang k, nu b l cng s to hoa nhanh; cn nu l s khng to hoa. V d: mai Nh vy, iu kin quang k khng thch hp, cy s to ra mt cht cn tr s to hoa. Cht ny theo dng cht hu c n n v cn tr to hoa. Khi c quang k thch hp, cht kch thch s c to ra di chuyn t l n n. y c mt s cn bng nghing v cht kch thch, cy s to hoa. Kt lun: S pht trin ca thc vt do gen kim sot. iu kin mi trng thch hp s cm ng ADN to ARN v gii m ra Protein to thnh enzyme xc tc cc phn ng. Qu trnh ny c kim sot bi cc cht iu ha sinh trng thc vt. Cc cht ny lun tc ng ln nhau t n mt t l cn bng gip thc vt pht trin theo hng nh sn.

84

CHNG XI:

S TO TRI V HI

-Khi hoa tng trng n mt lc no hoa s n: Cnh hoa m ra a ra hai c quan sinh dc c (b nh) v ci (nhy). y l hai b phn quan trng nht ca hoa. m nhim chc nng sinh sn qua qu trnh gim phn: -Nh: s cho cc giao t c (n) = ht phn -Nhy: s cho cc giao t ci (n) = nan (trng) -Ht phn rt trn nm nhy s kch thch bu nhy pht trin = TRI -Ht phn ny mm to ng phn i n bu nan, sau i vo ti phi cho hin tng th tinh kp (mt tinh t phi hp vi nan, mt tinh t phi hp vi nhn cc = nhn ph) ri pht trin to HT.

85

86

NGUN GC TRI
Cc khi nim tri Tri pht trin t b nhy, nhng nhiu c quan khc cng tham gia vo s thnh lp tri: Cnh hoa (du tm), (du ty), l bc (thm), c hnh thnh bi cc c quan hoa kt hp vi (Pyrus malus), hay bao quanh trc phc hoa (sung). Nhng c quan khc b nhy tham gia vo s hnh thnh tri th gi l tri gi (gi qu). Thng tri pht trin sau s th tinh, nhng cc tri chui (Musa), cam qut (Citrus), nho (Vitis) pht trin m khng c s hnh thnh ht. Hin tng ny c gi l tri khng ht. mt s thc vt nh u phng (Arachis hypogaea), tri ch pht trin sau khi cung nhy xuyn vo trong t mang theo l non vi bu non th tinh (Zamski and Ziv, 1976).
Cu to tri: V qu ngai (ngai qu b): c mu xanh do vch bu nhy pht trin V qu gia (trung qu b): do nhu m sinh ra. V qu trong: do biu b trong to ra, c vch dy. T bo c th di thnh si hoc c lng nh tri cam. tri cam v trong l thnh phn tch ri khi tri d dng khi trng thnh. 87

CAU TAO TRAI

88

Phn loi tri (qu)

Tri (qu) c phn thnh ba nhm Qu n: khi qu c thnh lp t mt bu non do mt tm b hoc do nhiu tm b dnh nhau (u c 1 tm b, chm rut c 3 tm b). C 2 loi qu n: qu mp v qu kh. * Qu mp: khi qu b dy v mm nh tch tr nhiu dng liu. Tu theo cu to ca ni qu b ta c hai nhm qu mp: ph qu v qu nhn cng. + Ph qu: khi ni qu b ln vi trung qu b v dnh st vo ht (i, c chua, u , bu b) + Qu nhn cng: khi ni qu b ho mc t tr nn cng bao bc ly ht bn trong (xoi, chm rut, cc). *Qu kh: khi qu b chn b tm mc t tr nn cng. C hai 89 nhm: qu kh t khai v bt khai.

+ Qu kh t khai: khi chn qu kh t m ra phng thch ht. Tu theo cch m, ta c cc loi qu sau: - Manh nang: qu kh do bu non c mt tm b lc chn khai theo 1 ng dc thai to (lp bp, trm). - Gip qu: qu kh do bu non c mt tm b nhng khai theo hai ng dc: 1 ng theo thai to, 1 ng theo sng lng (u xanh). - Gic qu: qu kh nt theo 4 ng dc dc theo hai bn thai to (mng mng). - Qu hp: qu kh nt theo mt ng vng ngang cho ra np bn trn v bnh cha bn di (mng g, rau sam, mi gi). - Nang: qu kh do bu non c t hai tm b tr ln to thnh. + Qu kh bt khai: qu kh khi chn khng nt phng thch ht ra ngoi, ta c cc loi qu sau: - B qu: qu kh bt khai c ht khng dnh qu b (sen, u)
90

Dnh qu: qu kh bt khai c ht dnh st qu b (la). Dc qu (qu c cnh): l loi b qu c qu b mc di ra thnh cnh bay theo gi (sao, du) Qu kp: khi qu do nhiu tm b ri to nn. Mi tm b cho mt qu. Cc qu ny phi hp vi nhau thnh mt qu duy nht (mng cu). Ngai ra cn c: Gi qu: qu khng do tm b to nn m do cc b phn khc ca hoa to nn. *Qu do hoa to nn (qu du ty) *Qu do cung hoa to nn (o ln ht) *Qu do pht hoa to nn (qu sung): y l mt phc hoa c hnh ci bu mang cc hoa rt nh trong. ca pht hoa ph to. *Qu do bao hoa to nn (cy hoa (bng) phn): sau khi th phn bao hoa rng i ch cn li phn y pht trin bao ly tri bn trong. Khi chn tri c mt lp v do bao hoa to nn bn ngoi. *Qu mt: do pht hoa (mt gi rt to) mang rt nhiu hoa c l i 91 nhng khng c cnh hoa pht trin. Mi tri l mt mi mt.

HT
ICu trc ht Ht gm: Phi cha: r mm thn mm chi mm t dip Phi nh cn hay c tiu ha V ca ht IINgun gc Non sau khi th tinh ht. Hp t ph 3n (1n t tinh t v 1 nhn ph-2n) phi nh . Hp t chnh (1 tinh t v 1 non cu) to trng (2n) phi. Phi tri qua bn giai on: + giai on hnh cu + giai on hnh tri tim (t dip bt u hnh thnh) + giai on hnh c ui (t dip hon thnh h thng mch xut hin) + giai on t dip hon chnh (h thng mch hnh thnh) Khi phi trng thnh (tch tr cht d tr, gim hp thu nc) v i vo min trng to nn ht trng thnh. 92 V non bin thnh v ht.

CU TRC HT (2 L MM)

93

CU TRC HT (MT L MM)

94

III-

Thnh phn ha hc

a. Nc: Nc chim mt lng ti thiu khang t 6 n 15% trng lng kh (TLK) ty lai ht. . b. Cht khong chim 1- 3% TLK ty lai ht
c. Cht hu c : carbohydrat (tinh bt) la, mt ; du (lipid): da, u phng, m v protid (m): u: xanh, nnh

d. Vitamin: nhiu lai vitamin rt cn thit cho con ngi: A (gc); B1 (la),
e.Cht iu ha sinh trng thc vt Trong ht c cha lng auxin t; cytokinin v giberelin nhiu. c bit c ABA, polyphenol khi ht trng thnh. Cc hp cht va k trn u c tng hp cc phn khc trong c 95 th thc vt (hay ngay ti ht) v di chuyn n tch tr ht.

IVi sng ca ht i sng ca ht l khong thi gian t khi phi c thnh lp y n khi ht ny mm thnh cy con. y l giai on sng chm. i sng ngn: cao su, thu du c i sng ca ht nh hn mt thng; u phng, m ht c i sng t 6 thng n 1 nm. i sng trung bnh: c thi gian sng t 3-10 nm. V d: da chut, c chc, a s ht ca cy n tri i sng di: > 10 nm: g, sen. i vi cc loi ht c i sng ngn, trong ht c cha du. Ngc li i vi cc ht c i sng di, ht thng cha tinh b, v ht dy, xp kht nhau lm gim s trao i vi mi trng.

96

VS ln mm ca ht Ln mm l s ti lp tng trng ca phi a r mm ra ngoi v. R mm chui ra non khu v m thng vo t. C hai kiu ln mm: Ln mm thng a: tr h dip tng trng a t dip (ht) ln trn mt t. T dip va c nhim v d tr (d tr l ch yu) va c nhim v ng ha v c dip lc nhn nh sng. Ln mm h a: tr thng dip tng trng nn ht (phi nh) nm li trong t. T dip ch l c quan d tr. a s ht c phi nh u c kiu ln mm h a.

97

VI- Hin tng sinh l Quan st hin tng ny bng cch tch ring Phi v phi nh nui ring Phi v phi nh nui chung Ch nui Phi vi mi trng dinh dng a. S hp thu nc Phi hp thu nc bng cch thm thu v c ch ch ng n bo ha. b. H hp Tng qu trnh h hp v thi nhit. Ban u phi c s h hp k kh c. Thay i cht d tr Cht d tr cha trong phi c chiu hng gim.

98

d. S bin i cc enzyme Enzyme c rt t ht kh nhng c tng hp mnh ht ngm nc. ht ny mm, c mt lng ln enzyme to thnh. S tng hp cc enzyme chu nh hng ca nhit , lng nc trong cc c quan. Phi thng tng hp enzyme nhiu nht ri n t dip v phi nh. e. Acid nucleic Tng rt mnh, nht l cc ARNm, ARNt, ARNr f. Sinh t (vitamin) Sinh t c nh hng n s pht trin ca cy mm. Nu trong thnh phn ca ht thiu sinh t th n s t tng hp thm cht ny khi ny mm. g. Cht iu ha sinh trng thc vt Ht tng tng hp cc cht kch thch (cytokinin, auxin, giberelin) v loi b cht cn (acid abcisic, polyphenol,.).

99

VII- iu kin cho s ln mm a. Ngoi yu t

Nc
L yu t quan trng nht. Nc c ht hp thu cho n khi bo ha. Oxy L cht cn cho qu trnh h hp. Nng oxy cn thit ph thuc vo tng loi ht. Nhit Nhit ma hoc nhit ngy c nh hng ln s ny mm ca ht. kch thch s ln mm ca ht, ngi ta thng s dng nc kt hp vi nhit thp. nh sng y l iu kin khng nghim nht, ty theo loi. Cht thu nhn nh sng ca ht l phytochrome, gm c hai loi: Pr thu nhn tia Pfr thu nhn tia xa Pr khi thu nhn tia s chuyn thnh Pfr kch thch s ln mm. Ngc li, t 100 Pfr chuyn thnh Pr s cn ny mm.

S cng sinh
i vi vi loi thc vt, cng sinh c tc dng tch cc ln s ny mm ca ht. V d: i vi ht lan, khi cng sinh vi vi khun Rhizotonia kch thch ht ny mm. Nguyn nhn: vi khun cng sinh cung cp cht dinh dng phi nh pht trin y . Khi , ht mi ln mm.

b. Ni yu t:
Ht phi trng thnh: C s pht trin y cc c quan ca phi Cha cc cht d tr cn thit gip cy mm pht trin Ht khng cn trng thi min trng: V ca ht thm nc v oxy Cc cht cn (nu c) c lai b Tng hp y cc cht thuc nhm kch thch
101

I SNG CHM
Trong qu trnh pht trin (chu k sng), c lc cc hot ng gim n mc ti thiu. gi l trng thi sng tim sinh. V d: ht, bo t

c tnh - Hot ng bin dng gim n mc ti thiu - C th s tr li i sng bnh thng nu iu kin thch hp. - S sng chm c tnh hon nghch.
Y ngha ca i sng chm: Chng chi li vi ngoi cnh.

102

A. Min trng ca ht
ICu trc ht Ht gm phi- r mm thn mm chi mm t dip Phi nh V ca ht. IINgun gc Non sau khi th tinh ht. Hp t ph 3n (1n t tinh t v 1 nhn ph-2n) phi nh . Hp t chnh (1 tinh t v 1 non cu) to trng (2n) phi. Phi tri qua bn giai on: + giai on hnh cu + giai on hnh tri tim + giai on hnh c ui) + giai on t dip hon chnh Khi phi trng thnh (tch tr cht d tr, gim hp thu nc) v i vo min trng to nn ht trng thnh. V non bin thnh v ht.
103

III- Thnh phn ha hc Nc (ti thiu), cht khong, cht hu c, vitamin, cht iu ha sinh trng thc vt. IVi sng ca ht i sng ca ht l khong thi gian t khi phi c thnh lp y n khi ht ny mm thnh cy con. Chia lm 3 loi: i sng ngn: t 6 thng n 1 nm. Thng cha du. i sng trung bnh: t 3-10 nm. i sng di: ht c cha tinh b, v ht dy, xp kht nhau VS ln mm ca ht Ln mm l s ti lp tng trng ca phi a r mm ra ngoi v. R mm chui ra non khu v m thng vo t. C hai kiu ln mm: + Ln mm thng a: 104 + Ln mm h a:

VI- Hin tng sinh l a. S hp thu nc Phi hp thu nc bng cch thm thu v c ch ch ng n bo ha. b. H hp Tng qu trnh h hp v thi nhit. Ban u phi c s h hp k kh c. Thay i cht d tr Cht d tr cha trong phi c chiu hng gim. d. S bin i cc enzyme Enzyme c rt t ht kh nhng c tng hp mnh ht ngm nc. e. Acid nucleic Tng rt mnh, nht l cc ARNm, ARNt, ARNr f. Sinh t (vitamin) Sinh t c nh hng n s pht trin ca cy mm. Nu trong thnh phn ca ht thiu sinh t th n s t tng hp thm cht nay khi ny mm. g. Cht iu ha sinh trng thc vt Ht tng tng hp cc cht kch thch (cytokinin, auxin, giberelin) v loi b cht cn (acid abcisic).
105

VII- iu kin cho s ln mm a. Ngoi yu t Nc L yu t quan trng nht. Nc c ht hp thu cho n khi bo ha. Oxy L cht cn cho qu trnh h hp. Ph thuc vo tng loi ht. Nhit Nhit ma hoc nhit ngy c nh hng ln s ny mm ca ht. nh sng y l iu kin khng nghim nht, ty theo loi. Cht thu nhn nh sng ca ht l phytochrome: Pr v Pfr. S cng sinh i vi vi loi thc vt, cng sinh c tc dng tch cc ln s ny mm ca ht. b. Ni yu t
106

B- S min trng ht
D cc iu kin ny mm (ngoi yu t) nhng ht vn khng ln mm min trng l do cc yu t bn trong (ni yu t). Min trng thay i ty theo loi ht. I- nh ngha Ht min trng l ht cha c kh nng ln mm d ht vn sng. l v ht cn phi c thi gian tng hp cc cht iu ha sinh trng thc vt, bin i cc cht cn trong ht ht ny mm. II- Nguyn nhn a. Do ht cha trng thnh sinh l b. Trng lng ht Ht nng cn, pht trin u n s d ln mm hn. c. V ca ht d. Min trng do c quan Do chi, r mm khng tng trng bnh thng do s phn phi nc trong ht km. 107 Do vi khun tiu hy auxin lm cho l mm qun queo.

e. S thiu cht dinh dng Thiu acid amin nh cystein, thioure; thiu cht iu ha sinh trng: cytokinin, giberelin; d cht cn nh acid abcisic f. Cc cht cn Cht cn s ln mm gm: Acid hu c: tri mp. Acid v c: HCN. V d: h hoa hng (rosaceae) Tinh du: khng cc ht cy khc ln mm, ch cho ht ca nhng cy c tinh du pht trin. V d: s, khuynh dip, trm Lacton: coumarin Alkaloid: cafein, nicotin Ethylen t tri chn a s cht khng cn s hp thu O2, nc nh hng ln s h hp. Thng th cht cn s gim dn trong qu trnh bo qun, ngm hoc phi ht do cc cht ny b oxy ha, quang oxy ha hoc nhit phn hy.
108

C- Hu min chi
I- iu kin mi trng Trong nhiu loi cy g, c bit l thc vt c ht vng n i, min trng chi rt ph bin. S to chi ngh (min trng) c kim sot bi quang k. S min trng chi cn ko theo s rng l, s gim hot ng ca tng tng v s gia tng chng chu lnh.
II- Ni nhn cm ng L l ni nhn bit s thay i ca mi trng. III- C ch Quang k nh hng trn min trng chi. Phytochrome trn l l ni nhn nh sng (theo Vince-Prue, 1985). S min trng chi c hai kiu: - Kim sot bi m di: c kim sot nghim nht - m di khng bt buc.
109

IV- Cht cm ng gy min trng a. ABA (acid abcisic) C mt s lin h gia nhm cht cn tng trng v s min trng. iu ny c chng minh bng sinh trc nghim. Tuy nhin, khng phi ABA l cht duy nht gy min trng. b. Giberelin C lin quan n min trng r rt nht. cy trong iu kin ngy ngn, lng giberelin gim xung rt thp. Khi cc chi min trng hoc tng trng chm, lng giberelin cng gim theo. c. Cytokinin Khng nh hng trc tip ln min trng chi. Nhng cy vo min trng, lng cytokinin gim n ti thiu v chng s gia tng tr li khi ph v min trng. V- Ph v min trng chi - Thc vt phi tri qua mt thi k lnh. - X l cc cht kch thch tng trng nh giberelin - Ko di di ngy cng cn min trng.

110

Kt lun
Thc vt qua cc cch trao i nc v cht khong vi mi trng s l ngun nguyn liu tng hp cht hu c. Qua c ch quang hp v h hp, thc vt to thnh cc hp cht bin dng v nng lng cn thit tng trng v pht trin. Trong qu trnh sng thc vt phi chu tc ng bi cc yu t mi trng. Cc yu t tc ng ln c th thc vt t thc vt s t chc li c th (cu trc v hat ng bin dng) phn ng vi mi trng mt cch tt nht tn ti v pht trin. Cc hat ng c xem xt cc mc khc nhau: c th, c quan, m, t bo v di t bo.

111