Sinh trưởng thực vật

8/13/2019 Sinh trng thc vt

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Vn 2: Sinh trng thc vt: khi nim, c ssinh hc phn t vngun n, c ch trnh thn tin inh hc

hm h n nh-i trn hc Page 1

Mc lc

Trang

I- Khi nim3 - 4

II- Cssinh hc phn tv ngun gen4 19

1. Kch thc, tchc vtnh phc tp ca bgen (genome)5

2. Sbiu hin gen vsinh hc pht tr in5 19

a) Sbiu hin gen cthtin nhn (Prokaryote)..5 9

b) Sbiu hin gen cthc nhn (Eukaryote).10 19

III- Cchv qu trnh thng tin sinh hc...20 251. Nhng khi nim cbn vnh nh.20 21

2. Cc con ng truyn tn hiu vcc cht nhn cthtin nhn.21- 25

a) Struyn tn hiu trong cthtin nhn21

b) Cc cchcbn.21

c) Vi khun sdng cc hthng iu tit hai thnh phn cm nhn cc tn

hiu ngoi bo.21 22

d) Phlan truyn ca cc trli.22 - 23

e) Vi khun sdng hthng hai thnh phn pht hin thm thu ca mitrng23 - 25


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3. Tn hiu thhai............................................................................25 - 31

a) Quan nim vtn hiu thhai..25

b) Mt si din tn hiu thhai phbin.26 - 31

4. Struyn tn hiu trong cthc nhn.31 - 36

a) Hai lp tn hiu xc nh hai lp cht nhn..32

b) Cc cht nhn steroid hot ng nhl cc tc nhn phin m.32 - 33

c) Cht nhn bmt tbo c thtng tc vi G-protein33 - 34

d) Chu trnh G- protein dtam phn l cu dao phn tca cc dng hot tnh vbt hot.34 - 35

e) Hot ha adenylat xiclaza.35

g) Shot haphotpholipaza C35

h) IP3mcc knh canxi vph LNSC vpha mng khng bo.35 - 36

i) Cc hthng hai thnh phn trong cthc nhn..36

5. Spht tn hiu..36 - 37

Ti liu tham kho.38


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I- C th thc vt pht trin nh hai kiu in i lin quan cht ch vi nhau: in i

nh lng v in i nh tnh (v cht).

- Nhng bin i vlng nhsgia tng khng thun nghch ca kch thc (chiudi, bmt, thtch) hay khi lng.

Khc vi qu trnh bin i nh lng ca vt thv sinh, chng hn, mt thanh kimloi di ra khi nhit mi trng tng, nhng ri co li khi gp lnh, hay ct thch nhtrongcc hang ng cao, to thm theo thi gian nhng khng c sto mi vcht, ngc li mt

ht khi ht nc sny mm, trong thi gian ny mm mc du khi lng kh ca ngim xung do cht dinh dng trong ht btiu ph cy mm xut hin v ln ln, mttrng thi mi vi nhng cu trc mi (tbo; m; cquan nhr, thn, l c hnh thnh(hnh 1.1)). Khi cy c rht c cc cht dinh dng v l ha lc quang hp c,cy mm chuyn sang cy non vi kiu sng tdng, lc ny kch thc, bmt l v khilng cy tng nhanh. Nh vy, sinh trng l mt khi nim hon ton sinh hc. Sinhtrng ca thc vt l qu trnh tng khng thun nghch vkch thc(chiu di, bmt,thtch) v khi lng km theo sto mi cc thnh phn cu trc (tbo, m, cquan)ca cth.

- Sbin i nh tnh c thc hin nhshnh thnh mi vmt hnh thi v chcnng lm xut hin nhng sai khc vcht gia cc tbo, m v cquan c gi l s

phn ha.

Phn ha l khi nim thuc phm tr pht trin sinh hc.Pht trin l nhng bin ivcht trong cu trc, hot tnh chc nng ca ton bcthv ca cc bphn cu thnhn (cquan, m, tbo) trong tin trnh pht sinh c th.

Pht sinh c th(ontogenesis) hay chu trnh sng l tng thnhng bin i chc v

hnh thi do di truyn gy nn trong cththc vt bt u thp thay mm sinh dngn ci cht tnhin trong iu kin bnh thng ca ngoi cnh (hnh 1.1).


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Hnh 1.1. Chu trnh sng ca thc vt Hai l mm

(Ngun: Sinh hc pht trin thc vt - Nguyn NhKhanh)

II- Cssinh hc phn tv ngun genNh ni trn, sbin i vcht trong chc nng v hnh thi (kiu hnh) ca c

thl kt quca sbiu hin gen. Cc tn hiu bn trong l rt cn phi hp nhp nhngsbiu hin gen trong chu trnh pht trin v cy c thphn ng i vi tn hiu tmitrng ngoi.


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Trc khi nghin cu cc qu trnh pht trin mc tbo v cthchng ta cn cnhng thng tin csngn gn vsbiu hin gen v s truyn tn hiu trong c th tinnhn (vi khun), trong nm men v ng vt tham kho. Nhng hnh mu biu hin gen

trn i tng y scung cp cho ta kin thc khungi vi nhng thnh tu mi nht trongnghin cu spht trin ca thc vt cc chng tip theo.

1. Kch thc, tchc v tnh phc tp ca bgen (genome)

Kch thc b gen c mi lin quan xc nh i vi mc phc tp ca c th.Chng hn bgen caE. Coli cha 47.106cp bazo (Cb), bgen ca rui gim (Drosophila)l 2.108Cb trong tbo n bi, chs ca con ngi l 3.109Cb. Kch thc ca bgenthc vt cng bin i nhiu, t1,5.108Cb trong cy Arabidopsisn 1.1011Cb trong cyMt l mm Trillium. B gen thc vt cha khong 25000 gen, trong khi b gen ca

Drosophilacha khong 12000 gen. Tuy nhin, kch thc ca bgen trong tbo cthcnhn (Eukaryote) l chs cha ng tin cy vphc tp ca n v khng phi ton bchui ADN u m ha gen. Chng hn, ngy nay sau khi phn lp bn gen ngi, thyrng khong hn 95% chiu di ca xon kp ADN khng cha gen no. S lng gen mha protein trong bgen ngi l khong 30000, chgp hai ln so vi giun hoc rui gim.

Trong cc c th tin nhn (Prokaryote) hu nh tt c ADN u cha cc trnh tnucleotit m ha cc protein hay cc phn tARN chc nng. Cc nhim sc th(NST) trongcthc nhn cha slng ln cc ADN khng m ha. Hnh nhiu c lin quan vi

chc nng tchc v cu trc ca nhim sc th(Lincoln Taiz, Eduardo Zeiger, 1998). Slng cc ADN khng m ha bao gm cc trnh tnucleotit c nhiu bn sao c gi lADN lp li (repetive DNA). Phn cn li ca ADN khng m ha c to thnh t cctrnh t nucleotit t sao chp n c gi l ADN chm. Kt hp ADN lp li v ADNchm c thto thnh phn ln ca bgen tng thtrong mt scthc nhn. Chng hn,trong cthcon ngi chc khong di 5% ca ton bADN cha gen c ngha l nhngtrnh tduy nht c m ha cho stng hp ADN v protein

2. Sbiu hin gen v Sinh hc pht trin

a) Sbiu hin gen cthtin nhn (Prokaryote)

- Protein lin kt vi ADN iu tit sphin m trong cthtin nhn

Trong c thvi khun cc gen c sp xp vo cc operon. Operon l b(tp hp)cc gen ktip nhau gm ccgen cu trcv ccgen iu tit. V d, lactozo (lac)operon

E. Coli c Francois Jacob v Jacques Monod vin Pasteur Paris m t ln u tin vo


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nm 1961. Lac operon l mt v dvoperon cm ng.E. Coli sng trong rut ngi. Nubui sang ta n bnh gato pht bth vi khun schm trong ng glucozo v fructozo vtiu ha thc n bo. Nu sau ta ung sa th mi trng sng ca E. Coli s thay i

mnh. Sau khi ta ung sa, mt trong nhng cht dinh dng ch yu trong sa l nglactozo. Khi lactozo cha y trong rut,E. Coli c thto ra enzim sdng lactozo. Jacobv J. Monod xut m hnh gii thch bng cch no tboE. Coli c thsn xut ra ccenzim ca n trong phn ng tr li nhng bin i thng xuyn xy ra trong mi trng

bao quanh n.

M hnh ca Jacob v Monod (hnh 1.2A v 1.2B) gii thch lm thno gen m hacc enzim cm ng sdng lactozo c nghay ngtphthuc vo iu kin c lactozohay khng.

Trong thi gian thch ng vi lactozo, trong cthE. Coli cm ng ng thi 3 enzim:- galactosidazac tc dng phn gii lactozo thnh ng glucozo v galactozo; permeazacn cho svn chuyn lactozo qua mng vo bn trong tbo; transacetylazakhng tham giavo schuyn ha lactozo.

E. Coli sdng 3 enzim thu nhn v bt u trao i lactozo, lc cc gen m hacng ng vai tr nhl n viu tit. Nhn vo hnh 1.2A, ta thy r rang 3 gen m hacc enzim ktip nhau sdng lactozo. l 3 gen cu trc(gen sdng lactozo). St knhm cc gen cu trc c cc on ngn ADN gip khng chchng. Mt trnh tc

hiu cc nucleotit, mtgen khi u(promoter), nh du im bt u phin m i vi ttc3 gen cu trc. gia gen khi u v nhm 3 gen cu trc l mt trnh tcc nucleotitc gi lgen chhuy(gen iu khin- operator). Gen ny hot ng nhci cu dao ng,ngt in. Gen chhuy quy nh thi im enzim phin m, ARN- polimeraza, c thlin ktvo gen khi u v y cc gen di chuyn vpha trc.

Mt cm cc gen cu trc nhvy vi nhng chc nng lin quan, cng vi gen khiu v gen chhuy c gi l mt operon. Operon ch tn ti trong cthtin nhn.

Hnh 1.2A trnh din lac operon trong m hnh ngt chra trng thi khi khng

c ng lactozo trong mi trng bao quanh tbo. Sphin m bngt (bphong ta) biphn tprotein c gi l cht c ch. l mt protein hot ng bng cch lin kt vigen chhuy v phong ta khng cho ARN- polimeraza tip xc vi gen chhuy. Gen iu ha(hnh 1.2A) nm pha ngoi operon; gen iu ha m ha cht c ch.

Sbiu hin ca gen iu ha xy ra lin tc, do vy tbo lun c cung cp ccphn tcht c ch.


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Lm sao cc operon c th ng (hot ng) nu cht c ch lun xut hin? Hnh1.2B (lac operon ng) cho thy lactozo cn trshot ng ca cht c chbng cch linkt vi cht c chhnh thnh nn phc hlactozo- cht c chlm bin i cu hnh ca n

(cht c ch). Vi cu hnh mi, cht c chkhng c khnng lin kt vi gen chhuy vgen ny vn duy tr trng thi ng. trng thi ny, enzim ARN- polimeraza c thlin ktvi gen khi u v t n bt u trt ngc dng t5 n 3 theo cc gen ca operon.mARN c sinh ra nhl mt phn tn mang trnh tm ha ca tt c3 enzim cncho shp thv sdng lactozo. Tbo c thdch thng tin thnh cc polipeptit tch bitv rng mARN c cha cc m tn hiu khi u v kt thc ca qu trnh dch m.

(Ngun:http://withfriendship.com/images/c/13181/3-operon-lac-operon.gif)

Lac operon ch l mt trong nhiu kiu operon trong c th vi khun. Nhng kiuoperon khc nhau cng c gen khi u, gen ch huy v mt s gen cu trc, nhng ccoperon khc nhau c cc phng thc khng ging nhay trong vic kim tra sng, ngtgen chhuy. Trn hnh 1.3 trnh by hai kiu cc operon kim sot cht c ch. Cht c chca lac operon trng thi hot tnh khi tdo v bmt hot tnh khi lin kt vi lactozo.Trn hnh 1.3 cng gii thiu kiu operon th hai, kiu trp operon iu khin theo cchngc li. Trp operon c iu khin bi cht c chm khi trng thi tdo th bmt
http://withfriendship.com/images/c/13181/3-operon-lac-operon.gifhttp://withfriendship.com/images/c/13181/3-operon-lac-operon.gif


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hot tnh. c hot tnh, kiu cht c chny phi lin kt vi cht phn tb c hiu.Trong v dtrn hnh 1.3, cht c chphn tb l axit amin tryptophan.E. Colic thsn ra axit amin tryptophan nu n cn, bng cch sdng cc enzim c m ha trong trp

operon. Nhng n sngng sn sinh ra tryptophan v hp thn tmi trng xung quanhkhi c th. Khi c sn tryptophan, cht ny lin kt vi cht c chca trp operon, hot hacht c chca trp operon lm cho n ngt operon. Nhvy, kiu operon ny cho php vikhun ngng tng hp mt scc cht chyu khi cc cht ny c sn trong mi trnggip tit kim nguyn liu v nng lng cho tbo.


Ngoi hai kiu operon va nu cn c kiu operon thba. l kiu operon sdngcc cht hot ha. Cc cht hot ha l cc protein ng operon bng cch lin kt vi cc

ADN. Bng cch no cc protein y lm cho cc ARN- polimeraza ddng lin kt vigen khi u hn l phong ta enzim ARN- polimeraza, nhtrng hp cc cht c chtcng. Ngoi ra cn cc kiu operon khc na.

Vi vic trang bcc kiu operon khc nhau c iu tit theo cch sdng cc chtc chhay cc cht hot ha,E. Coli v cc cthtin nhn khc c thpht trin c trongnhng mi trng lun bin ng.


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b) Sbiu hin gen cthc nhn (Eukaryote)

Trong c th tin nhn, s dch m (translation) lin kt vi s phin m(transcription) v mARN phin m ko di ra, gn vi cc riboxom v tng hp protein (dchm). Tuy nhin, trong cthc nhn, mng nhn tch bgen ra khi bmy dch m. Trcht, trong cthc nhn, mng nhn tch bgen ra khi bmy dch m. Trc ht cc sn

phm phin m c vn chuyn ra tbo cht, phi c bsung cc tuyn (mc ) kimtra khc.

- Sng gi ADN trong nhim sc th ca tbo c nhn nh hng n sbiuhin gen

V d, tng ADN trong 46 nhim sc thca tbo ngi nu cng ra sdi 3 mt

(Neil Campbell et al., 1997) trong lc ng knh ca nhn in hnh chl 5m. Tt cADN trong mi tbo u nm gn trong nhn chc hthng xp gp (ng gi) ADN tmnhiu bc trong mi mt nhim sc th.

Vai tr quyt nh ca sng gi l mi lin kt ADN vi protein phn t b chiu c tn l histon ch pht hin c cthc nhn (vi khun cng c protein tng t,nhng chng khng c sxp gp nhiu bc ADN nhtrongEukaryote). Trn cc nh hinvi in tphc hADN- histon c hin ra nhl cc ht gn trn si dy. Mi ht c tngi l nucleoxom (thnhn), nucleoxom gm c ADN qun quanh protein c tm phn thiston. Nucleoxom c thgip kim sot sbiu hin gen bng cch hn chcc enzim phinm xm nhp vo ADN. Phn ln cc nucleoxom cng cha cc protein khng histon. Tboc nhn in hnh cha khong 1000 cc protein nhim sc thkhng histon khc nhau m

phn nhiu trong chng c ththam gia iu tit gen.

mc ng gi tip theo ca ADN, dy c gn ht li c gi vo si xon kp cht.Sau si xon kp li c qun tip vo cun di hn vi ng knh khong 200 nm.Tip theo cc cun ADN li c tht thnh vng v ng gi cht hn na nh ta c ththy nhim sc thk gia (metaphase) trong qu trnh nguyn phn (mitosis). Sxp gpnhiu bc ktip nhau trong sng gi ADN lm cho mi nhim sc thcha c lng

khng lADN. Vai tr ca sng gi ADN trong skim sot biu hin gen hu nhchabit y ngoi trmt st trng hp c bit hp dn. Mt trng hp nhvy l thBarr (Barr body) c tm thy trong cc tbo c thci ca ng vt c v. ThBarr ldng nhim sc thX c ng gi rt cht. Hu ht cc gn trong nhim sc thX l bmthot tnh. Khi con ci sinh ra, mt trong hai nhim sc thX ca cc tbo xoma l thBarr(hnh 1.4) v v vy mt hot tnh. Nhim sc thX trong tbo no bmt hot tnh l


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hon ton ngu nhin. Kt qul trong dhp tci i vi cc gen trn nhim sc thX cacc tbo khc nhau biu hin ra thnh cc alen khc nhau.


Mt v dgy n tng su sc vnh hng n kiu hnh ca thBarr l mo calico.

-iu tit sbiu hin gen mc phin m

iu tit sbiu hin gen mc phin m trong tbo c nhn thng din ra nhtrong cthProkaryota. Kin thc phn ny cho n nay chyu l da vo cc thnh tunghin cu trn i tng ng vt. i tng c nghin cu tmvmt di truyn l ruigim (Drosophila melanogaster). Sbiu hin mc phin m c pht hin cc t

bo tuyn nc bt ca u trng rui gim l sminh chng v c chiu tit qu trnh

phin m trong cc tbo ca cthc nhn. i vi thc vt, m hnh biu hin gen angc nghin cu cyArabidopsis thaliana(ci soong).

Thc nghim cho thy siu tit hot tnh biu hin gen lin quan n cc tn hiuha hc. Cc hoocmon l tn hiu ha hc. Chng hn hoocomon steroid iu tit sphinm ca mt sgen xc nh bng cch lin kt vi protein iu ha v hot ha n. Trong


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trng hp ny, protein iu ha l mt cht hot ha (activator). Cht ny hot ha cc genbng cch lin kt cc trnh tnucleotit (cc gen c hiu).

Ngi ta nh r c s lng ln cc protein hot ha trong t bo c nhn.Ging vi cc protein iu ha trong cthtin nhn, nhiu protein hot ha trong cthcnhn hot ng theo cch gn vo ADN v bng cch no nh hng n khnng caARN- polimeraza bt u phin m.

- Skhc bit trong tchc bgen ca tbo c nhn

(Ngun: Sinh hc pht trin thc vt - Nguyn NhKhanh)Trong sln cthc nhn, mi gen m ha mt polipeptit n. Bgen trong nhn

ca c th c nhn khng cha operon tr mt sngoi l (khong 25% sgen trong congiun trn Caenorhabitis eleganstrong cc operon. Operon cc ARNphin mtrthnh ccmARN c th thc hin m ha cc polipeptit n (monocistronic mARNs) bng cch phihp sphn ct, poliadenin ha v tch intron ra khi ARN (Kuersten v CS., 1997). Ngoi


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ra, cc gen trong cthc nhn c phn thnh cc min m ha gi l exonv cc minkhng cha m thng tin gi l intron(hnh 1.5).

- Siu tit sau phin m

Sau khi phin m tADN thnh ARN gi l tin mARN (pre- mARN) hay ARN phinm (RNA transcript). ARN ny cn cha cc intron, do cha c ththc hin sdch m(tng hp protein) tip ngay nhtrng hp vi khun c. Tin ARN ny phi tri qua qutrnh chin (qu trnh xl sau phin m) bao gm cng on ct bcc intron v ni cc exonthnh ARN gi l ARN ni (RNA splicing), tip theo cn phi thc hin qu trnh to m(capping) v qu trnh ni ui poly- A. Khi hon tt cc cng on x l , tin ARN(pre- ARN) chin v gi l ARN ni (mARN), mARN ny phi ri khi nhn i vo tbocht khi u sdch m tng hp ra cc protein tng ng (hnh 1.5).

+ Qu trnh ct bintron v lin kt exon thnh ARN nic thxy ra theo hai cch:1. Qu trnh c phc hprotein v cc ARN phn tb xc tc; 2. Trong nhng trnghp xc nh khc, sni kt cc exon xy ra hon ton khng c protein hay ARN phn t

b tham gi, ARN phin m (tin ARN) txc tc qu trnh. Ni cch khc, ARN hot ngnhmt enzim.

(Ngun:http://thuviensinhhoc.violet.vn/entry/show/entry_id/1838862)
http://thuviensinhhoc.violet.vn/entry/show/entry_id/1838862http://thuviensinhhoc.violet.vn/entry/show/entry_id/1838862http://thuviensinhhoc.violet.vn/entry/show/entry_id/1838862


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+ Chc nng sinh hc ca intron l g? Hin nay cha c cu tr li y . Tuynhin, cc nh sinh hc c nhng nhn xt vvai tr ca intron nh sau: Mt sADNkhng c intron khng cha nhng trnh tnucleotit bng cch no iu tit hot tnh gen.

Qu trnh ct bintron c thgip kim sot c dng mARN tnhn n tbo cht. Hnna, trong mt s trng hp, tbo c th thc hin sct bintron v ni kt exon theocch khc nhau v do c th to ra nhng phn tmARN khc nhau t cng mt tinmARN (hnh 1.6).

(Ngun: Sinh hc pht trin thc vt - Nguyn NhKhanh)Vi cch nhvy, mi cthc thsn ra nhiu kiu polipeptit tmt gen n. S

phn ha gii tnh cng l do skhc bit ln trong hnh mu phn ha ca qu trnh toARN ni (Neil A. Campbell v CS., 1997).

Cc intron cng c thgp phn vo tnh a dng trong pht trin. Chng hn, straoi cho trong gim nhim l mt trong nhng ngun chyu ca sbin ddi truyn trongcthc nhn. Sc mt ca intron lm cho ADN di ra bng cch tng slng cc im,ni c thdin ra strao i cho gia hai alen ca mt gen. Trong tnh a dng di truyn

do sc mt ca cc intron c thc ngha quan trng trong stin ha ca cthc nhn.


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(Ngun:http://trithucsangtao.vn/i485-ctnnsh-bai-6--tu-dna-sang-rna-su-phien-ma.aspx)

Trc khi ct b intron, tin mARN c chp m bng cch thm mmetylguanilat vo u 5. mARN phin m c gn m ngay sau khi bt u tng hpmARN. Mt trong cc chc nng ca m5 l bo vARN ang c hnh thnh khi benzim ARN- aza phn gii. Ti mt bc vsau, trong qu trnh tng hp sn phm phin mu tin (ARN phin m) ti mt v tr c hiu, u 3 phn ct v c gn thm ui

poly- A (hnh 1.7) thng gm t100 n 200 gc adenilic.


ui poly- A c mt s chc nng: 1. Bo vARN khi ARN- aza v lm tng tnh nnh ca phn tmARN trong tbo cht; 2. ui poly- A v mti 5 cn cho schuynARN qua lnhn; 3. Tng hiu sut phin m trn riboxom.
http://trithucsangtao.vn/i485-ctnnsh-bai-6--tu-dna-sang-rna-su-phien-ma.aspxhttp://trithucsangtao.vn/i485-ctnnsh-bai-6--tu-dna-sang-rna-su-phien-ma.aspx


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Cc tc nhn phin m chung (general transcript factors) gn vo phc h khi uphin m (transcription inititionx complex) ti hp TATA (TATA box) ca trnh tkhi unm trong 100 Cb ti vtr bt u phin m ca gen.

(Ngun:http://murj.mit.edu/features/31)

Cc trnh tiu ha dng cis bsungnhhp CAAT (CAAT box) hay hp GC (GCbox) lin kt vi cc tc nhn phin m c tc dng tng s biu hin gen. Cc trnh tnucleotit iu ha txaphn bxa hn im ngc dng (upstream) lin kt vi cc tc nhn

phin m khc c gi l cc cht hot ha (activator) hay cc cht c ch (repressor).Nhiu gen thc vt cng c cc gen tng cng (enhancer), nhng trnh tiu ha dngtnh txa, iu khin.

Mc du phn tn sut khp bgen, nhiu gen trong tbo c nhn u l gen cmng v cng c iu tit. Cc gen c iu tit nhp nhng c cc trnh tiu ha dng

cis chung trong gen khi u ca chng. Hu ht cc nhn tphin m trong thc vt chanhp cha kha cbn. (bZIP).

Trong c th thc vt c nhm quan trng cc tc nhn phin m: cc gen pht sinhng ngun (gen ng dng) cha min MADS (chci u tin ca nm thnh phn u ca
http://murj.mit.edu/features/31http://murj.mit.edu/features/31


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lp gen: gen nm men MCM- 1 m ha tc nhn phin m, cc gen ng dng ca cquanhoa AGAMOUS v DEFICIENS, tc nhn trli huyt thanh ca th, mammalian serum).

Qu trnh tng hp hu ht protein trong tbo c nhn c iu tit mc phinm. Sphc tp trong phin m ca tbo c nhn so vi tbo tin nhn c biu hintrong cc khc bit sau:

c im cu trc cbn l sbin ng ca nhp kha loxin c cha nhng axit aminght nc thay thloxin v min lin kt ADN cha cc axit amin. V dhai protein cha rtrong cy ng, cc tc nhn hp G cha cc gen iu ha phitocrom, cc tc nhn phin mlin kt cc yu ttrli lin kt axit abxixic (ABA).

Thnht, tn ti ba ARN- polimeraza khc nhau trong tbo c nhn: I, II, III. ARN-

polimeraza I nh ctrong nhncon v c chc nng tng hp hu ht ARN riboxom,ARN-polimeraza II cng phn b trong c cht nhn (nucleoplasm) c chc nng tng hp tinmARN, ARN- polimeraza III cng phn b trong c cht nhn chu trch nhim tng hpnhng ARN b nhtARN v rARN 5s.

Thhai, ARN- polimeraza tbo c nhn i hi cc protein bsung gi l cc tcnhn phin m chung c chnh xc vo v tr khi u (initition site). Cn ARN-

polimeraza trong t bo tin nhn cng i hi cc polipeptit ph c tn gi l tc nhnim (). Cc polipeptit ny l cc bphn di n vca ARN- polimeraza. Ngc li,cc tc nhn sau phin m chung to thnh phc hkhi u phin m nhiu kin trc din v (multisubunit). V d, by tc nhn phin m chung cu thnh phc h khi u(initition complex) ca ARN- polimeraza II, mi mt tc nhn phi c bsung theo mttrt txc nh trong thi gian tp hp.

Thbal skhc bit trong tnh phc tp ca gen khi u, trnh tngc dng (5)ca v tr khi u iu tit sphin m. C thchia cu trc ca on khi u thnh hai

phn: Phn li (core) hay phn khi u ti thiu (minimum promoter) gm trnh tngcdng ti thiu (minimum upstream sequence) cn cho sbiu hin gen v cc trnh tiuha bsung thc hin skim tra hot ng ca phn hot ng li.

Mi mt trong ba ARN- polimeraza c kiu cc on khi u khc nhau.

Cc trnh tADN tchng hot ng c gi l trnh tdng cisv chng st kvi cc n vphin m m chng iu ha. Cc tc nhn phin m gn vi trnh tdng cisgi l cc tc nhn dng transv cc gen m ha chng phn b ti u trong b gen.

Nhiu cc trnh tdng cis phn bxa im ngc dng ca cc trnh ton khi u gn


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tm c thkim tra dng hay m i vi cc phn khi u trong tbo c nhn. Cc trnhty c gi l cc trnh tiu ha txav chng thng phn btrong 1000 Cb ca vtr bt u phin m. Cc tc nhn phin m (protein) dng dng tnh lin kt vo cc vtr

y c gi l cc cht hot ha, cn nhng tc nhn c chsphin m gi l cht c ch.

Siu tit biu hin gen do cc hoocmon thc vt lin quan vi hin tng mt hottnh ca cc protein c ch. Cc trnh t dng cis lin quan vi c ch iu tit gen do

phitohoocmon hay do nhng tc nhn ha hc khc gi l thnh t tr li (responseelements). Cc trnh t iu ha dng tnh phn b xa gi l gen tng cng. Gen tngcng c thphn bhoc ngc dng hoc xui dng so vi gen khi u. Nhiu thnh ttrli iu tit sbiu hin gen c nhn din trong cththc vt.

-iu tit sbiu hin gen mc trong v sau dch m

cthc nhn, sau khi xl ARN phin m thnh mARN chin ( loi bintron,gn mv gn ui), mARN ri nhn vo tbo cht. Xut hin chi iu tit biu hingen.

+ Thi gian sng ca phn tmARN l tc nhn iu tit biu hin gen mc dch m

Ni chung mARN sng lu c thtng hp c nhiu protein hn so vi mARN ci sng ngn. mARN trong cthProkaryotec thi gian sng rt ngn, chng bcc enzim

phn gii chsau t pht. l mt trong cc nguyn nhn lm cho vi khun c ththay i

nhanh protein ca chng trong phn ng trli i vi mi trng bin i.

Ngc li, trong c thEukaryotemARN c thi gian sng hang gihoc thm chhng tun l. V dn tng nht l thi gian sng lu ca mARN trong cc tbo hng cuca ng vt c xng sng n mt thng Chim v cn lu hn B st, ch nhi, C.

Trong cthc nhn tn ti nhng cchiu tit sau dch m:

Cchiu tit sau dch mbao gm ct cc polipeptit thnh cc sn phm cui cngb hn v hot tnh hn.

Nhp iu n nh hay phn gii cc mARNkhc bit gia cc m phthuc vo trngthi sinh l ca cy. V d, cy u (Vicia faba) snhim bnh nm gy nn sphn giimARN vn m ha protein giu prolin vi vch tbo cy u. V dkhc vsiu tit

biu hin gen ca mt trong cc gen i vi cu trc di n vb ca cht rubico trong rcy bo tm (Lemma gibba). Rbo tm quang hp v v vy n biu hin gen i vi cutrc di n vrubico, nhng sbiu hin ca mt trong cc gen trong rthp hn trong


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l. Jane Silverthorne v cng s cho thy mc biu hin gen thp nhvy bo tm l donhp iu phn gii cao ca mARN ca gen trong nhn (Peters and Silverthorne, 1995).

Tnh n nh trong dch mca cc phn tmARN cng bin ng. Chng hn, sxpgp ARN thnh cc phn tc cu trc bc hai, bc ba bin i cng gy nh hng n khnng t n riboxom ca m khi u dch m (trnh tAUG u tin).

Vic sdng m cng l mt tc nhn khc c thnh hng n khnng dch mca mt mARN. Sdtha trong m bba vn c hiu vi mi axit amin trong dch m vmi mt tbo c nhp iu ring c trng cho mi tARN mang axit amin c gi l lch m (codon bias). Cui cng, vtr ca tbo, ni din ra sdch m cng nh hng nnhp iu ca qu trnh biu hin gen. Nhng polixom tdo c thdch mARN vi nhng tcrt khc nhau ty thuc vo mi lin kt polixom vi mng li ni sinh cht, thm ch

bn trong li ni sinh cht cng c thtn ti nhng nhp iu dch m khc nhau. C thtm tt qu trnh iu tit biu hin gen trong tbo c nhn nhtrn hnh 1.8.



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III-Cchv qu trnh thng tin sinh hc1. Nhng khi nim cbn v ga

- Tbo phn ngi vi ngoi cnh bng cch ti tchc cu trc ca n, iu tithot tnh protein v thay i hnh mu biu hin gen. Skch thch cho nhng phn ng trli nhvy c gi l tn hiuv tn hiu c thl phn tb, i phn thay tc nhn vtl nhnh sangTn hiu tng tc vi tbo thng qua cc phn tcht nhn.

- Ccphn tbthng tc ng nhl cc tn hiu khuch tn. Trong cth, cc tnhiu khuch tn c thl tngoi cnh hoc ttbo khc (chng hn, kiu pheromone canm men, cAMP trong Dictyostelium). Trong ng vt a bo, tn hiu c th tcc tbo

bn cnh v khuch tn trong khong cch gn (tn hiu paracrin), hay xut pht tcc tboxa c truyn qua hdn (tn hiu ni tit). Cc tn hiu i phn tthng gn kt vi ccht ni bo hay nm trn b mt ca cc t bo ln cn (tn hiu ln cn juxtacrinesignaling). Tn hiu phn tlin kt vi cc cht nhn c gi lphi t(ligand).

- Cc tn hiu c thdin ra trong ba con ng. Mt stn hiu ha hc c ththmqua mng sinh cht ca tbo v tng tc vi cht nhn ni bo (chng hn nhsteroid).

Tuy nhin, phn ln cc tn hiu l cc phn ta nc, li bn ngoi tbo. Nhng tnhiu ny tng tc vi cc cht nhn xuyn mng (cu xuyn mng) hay vi cc cht nhnlin kt vi mng v gy ra sbin i cu trc ca cht nhn gip cht ny thm c vo

bn trong tbo (qua c mng nhbin i cu dng ca cht nhn, nhto ra cc l, vd, trong trng hp knh ion hay cht nhn xm nhp vo tbo bng cch thc bo). S

bin i cu dng cht nhn c th cm ng hot tnh enzim bn trong tbo mun to rahiu ng xui dng (theo gradient) trong khi phi t li bn ngoi (truyn tn hiu-signal transduction). Kch thch vt l c thtng tc vi cht nhn hay c thbng cch tcng trc tip. nh sang kch thch G- protein lin kt vi cht nhn rhodopsin v opsin t

bo hnh nn khi photon (lng t) lm cho phn tnhy cm nh sang c tip hp 11- cisretinal bin i tt ccu dng trans. Tri li, phn ng vi sc nhit v stress tng tctc ng trc tip nhgia tng protein bin tnh trong tbo.

- Struyn tn hiubao gm nhng con ng hot ha theo trnh tcc enzim v cciu bin mc cc phn tb gi l cc tn hiu thhai. iu cho php khuch i tn


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hiu gc. Con ng truyn tn hiu trc tip c thl hi thay phn hngcho php tngkch thch ti sinh nhng phn ng tng tv cc tn hiu ring bit to ra cc phnng trli khc nhau vi thi gian v cng kch thch, v stng hp c hiu tbo ca

cc cht nhn khc nhau v cc thnh phn ca tn hiu. Cc con ng truyn tn hiu lchc nng ca mng li iu tit phc tp, v phn ng trli phthuc vo scn bngca cc lc i khng trong tbo.

- Sphn btn hiubao gm shot ha hay c chcc tc nhn phin m enzim vcc thnh phn cu trc ca tbo, thng l do sbin i trng thi photphorin ha cachng.

2. Cc con ng truyn tn hiu v cc cht nhn cthtin nhn

a) Struyn tn hiu trong cthtin nhn

Tbo vi khun khng c thsng st hng tnm tin ha nu khng c khnngpht trin tuyt vi gic quan cm thmi trng. Vi khun phn ng i vi shin dinca cc cht dinh dng bng cch tng hp ra nhng protein lin quan n shp thvtrao i cht dinh dng . Vi khun cng c th phn ng i vi tn hiu khng dinhdng cvmt vt l v ha hc. Vi khun c thhiu chnh svn ng ca n ty thucvo gradient ang tn ti vpha nh sng, oxy, thm thu, nhit v ha cht c ctrong mi trng.

b) Cc cchcbn lm cho vi khun cm nhn v tr li i vi mi trng baoquanh l chung cho tt ccc h thng th cm ca tbo v bao gm skch thch phthin, khuch i tn hiu v cc trli tn hiu thch hp. Nhiu con ng truyn tn hiu vi khun cho thy s tn ti nhng n viu bin gi l cht truyn (transmitters)vcht nhn (receivers). Nhng cht iu bin y to nn csca cc hthng iu tit haithnh phn.

c) Vi khun sdng cc hthng iu tit hai thnh phn cm nhn cc tn hiungoi bo

Vi khun cm nhn ha cht c trong mi trng nhmt hnhcc cht nhn bmttbo, mi mt cht nhn lin quan vi phn ng trli i vi mt nhm cc ha cht xcnh (sau y sc ni n nhl cc phi t). Protein trong mng sinh cht ca vi khunlin kt trc tip vo phi thay vi protein ha tan vn gn vo phi ttrong khng gianngoi bin ca tbo cht v vch tbo. Nhvo slin kt, protein mng chu sbin i


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cu dng lan truyn qua mng n min tbo cht ca protein cht nhn. Sbin i cudng nhvy khi u con ng tn hiu dn n phn ng trli.

d) Phlan truy

n c

a cc tr

l

ivi khun gm c siu tit thm thu, tnh hng

ha v to bo tu do cc hthng hai thnh phn iu tit. Hthng iu tit hai thnhphn gm protein thcm v protein iu tit trli (hnh 1.9) (Parkinson, 1993).


Protein cm th tm thy kch thch min tn hiu vo v truyn tn hiu cho mintruyn bng cch bin i cu hnh (mi tn ---> u tin). Min truyn ca protein cm thsau lin kt vi cht iu tit trli bng cch photphorin ha protein ca min nhn. S

photphorin ha min nhn cm ng ra sbin i cu hnh (mi tn ---> thhai) lm hot

ha min li ra v i hng phn ng trli ca tbo (Parkinson, 1993).Chc nng cm thl nhn tn hiu v gi tn hiu n cht iu tit phn ng trli

c tc dng lm bin i hng phn ng ca tbo, c bit l sbiu hin gen. Cc proteincm thc hai min: min tn hiu vo v min truyn tn hiu. Min tn hiu vo thu tn hiutmi trng; min truyn tn hiu gi tn hiu n cht iu tit phn ng trli. Cht iutit phn ng trli cng c hai min: min nhn, nhn tn hiu tmin truyn tn hiu ca

protein cm thv min tn hiu ranhmin lin kt ADN vn lm i hng phn ng trli.

Tn hiu xut pht t min truyn n min nhn bng con ng photphorin haprotein. Min truyn c khnng tphotphorin ha sdng ATP ti gc histidin c hiu cnh nhm amin u cui cng. V vy, protein cm thcha min truyn c gi l kinazahistidin tphotphorin ha. Nhng protein y thng hot ng nhcc nhphn vn l nixc tc ca cu trc di n vphotphorin ha vtr cht nhn trn mt cht khc.


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A-Min truyn tn hiu ca protein cm thcha histidin (H) ti N - u cui cng can, cn min nhn ca cht tr li cha aspactat (D).B- Min truyn tphotphorin ha tihistidin v truyn photphat cho aspactat ca cht phn ng trli. Sau cht phn ng trli chu sbin i cu dng dn n trli (theo Parkinson, 1993).

Ngay sau min truyn trnn tphotphorin ha ti gc histidin cnh trung tmca min nhn ca protein iu tit trli, kt qul gc aspactat c hiu ca cht iu tittrli c photphorin ha (hnh 1.10). Sphotphorin ha gc aspactat gy nn sbini cu dng ca cht iu tit tr li thhin ra chn (cht iu tit tr li) c hot

ha.

e) Vi khun sdng hthng hai thnh phn pht hin thm thu ca mitrng

Hthng hai thnh phn tng i gin n vi khun l hthng tn hiu lin quann scm ththm thu E. Coli. E. Coli l vi khun Gram m v nhvy n c hai


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mng tbo, mng trong v mng ngoi tch bit nhau bi vch tbo. Mng trong l mtvt chn thm thnht ca tbo. Mng ngoi cha nhng l ln gm hai kiu protein l,OmpF v OmpC. Cc lc protein OmpF to nn ln hn so vcc lc protein OmpC

hnh thnh.

Khi m E. Coli buc phi chu thm thu cao trong mi trng, n tng hp nnnhiu proten OmpC so vi protein OmpF lm xut hin cc lnhtrn mng ngoi. Cc lnhny lc thi ra nhng cht tan tvng tbo cht ngoi bin c tc dng bo vmngtrong khi tc ng ca nng cht tan cao trong mi trng bn ngoi. Khi cho vi khunvo mi trng c thm thu thp, protein OmpF c tng hp nhiu hn v kch thctrung bnh ca cc ltng ln.

Sbiu hin gen m ha 2 protein lc hthng hai thnh phn iu tit trnh by

trn hnh 1.11, protein cm thEnvZ nh ctrn mng trong. Protein c min tn hiu voN- u cui cng vng sinh cht ngoi bin. l protein pht hin sthay i thmthy trong mi trng nm gia hay phn mnh mng bc ngang qua v min truyn tbocht C- min cui cng.

Khi thm thu ca mi trng cao, protein cm thtrn mng, EnvZ (dng nhphn) tc ng nhl kinaza tphotphorin ha histidin. EnvZ c photphorin ha sau photphorin ha cht iu tit tr li OmpR vn c min lin kt ADN. OmpR cphotphorin ha gn vo gen khi u ca hai gen lOmpC v OmpF tng cng sbiu hin

dng biu hin cht c chvsau. Khi thm thu ca mi trng thp, EnvZ hot ngnhl protein photphataza thay vo kinaza v loi photpho ca OmpR lin kt vi gen khiu ca hai gen l. Sbiu hin OmpC bc chcn sbiu hin OmpF c kch thch(theo Parkinson, 1993).

Khi thm thu ca mi trng tng ln, min tn hiu vo chu sbin i cu dngv c truyn qua mng n min truyn. Sau min truyn tphotphorin ha gc histidinca n. Gc photphat nhanh chng c dn n gc aspactat ca min nhn ca cht iutit trli OmpR. N- u cui ca OmpR cha min lin kt ADN. Khi c hot ha nh

photphorin ha, min ny tng tc vi ARN- polimeraza ti gen khi u ca cc gen l(poringen) lm tng sbiu hin gen ca OmpC v c chsbiu hin ca OmpR, kch thchsbiu hin OmpF v c chsbiu hin OmpC. Theo cch , kch thch thm thu ctruyn n cc gen.


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3. Tn hiu thhai

a) Quan nim vtn hiu thhai

Cc t bo tr li i vi hng lot tn hiu khc nhau v i hi phi c s lngkhng lcc cht nhn. Tuy nhin, vng phn bca cc phn ng trli th t hn nhiu.

Nhiu tn hiu t n bmt tbo lm cho tbo hoc phn chia hoc ri khi chu trnh tbo. Nhng tn hiu khc cm ng sbiu hin ca nhm gen c trng vn bo v tbokhi sc. V vy, nhng con ng tn hiu sm hi tvo mt scc mng tn hiu ni bov iu cho php t bo chuyn i thng tin phc tp n c b mt t bo thnhnhng tn hiu ha sinh n gin trong tbo cht. Nhng phn tlin quan n qu trnh gi l cc tn hiu thhai.


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b) Mt si din tn hiu thhai phbin (hnh 1.12)

- Cc tn hiu thhai - nuleotit mch vng

l 3, 5 AMP vng (3, 5-cAMP) v 3, 5-GMP vng (3, 5-cGMP). Nhng tnhiu thhai ny c pht hin sm nht. Hm lng cc phn tny trong tbo c iutit bi hot tnh ngc chiu ca cc nucleotidylat xiclaza vn xc tc phn ng NMPcNMP (v dAMP cAMP) v photphatdiesteraza nucleotit vng (PDEs), enzim ny xctc phn ng ngc li.


* cAMP (Cyclic adenosine monophosphate)

(Ngun:http://en.wikipedia.org/wiki/Cyclic_adenosine_monophosphate)
http://en.wikipedia.org/wiki/Cyclic_adenosine_monophosphatehttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search?q=cAMP&hl=vi&biw=1517&bih=701&prmd=imvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/File:Cyclic-adenosine-monophosphate-3D-balls.png&usg=ALkJrhjBnALGflT_RMieH-LU-zV8B5j4JQhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search?q=cAMP&hl=vi&biw=1517&bih=701&prmd=imvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/File:Cyclic-adenosine-monophosphate-2D-skeletal.png&usg=ALkJrhiOAvP0sstUz7VXae2GcM2-mzM6TQhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search?q=cAMP&hl=vi&biw=1517&bih=701&prmd=imvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/File:Cyclic-adenosine-monophosphate-3D-balls.png&usg=ALkJrhjBnALGflT_RMieH-LU-zV8B5j4JQhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search?q=cAMP&hl=vi&biw=1517&bih=701&prmd=imvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/File:Cyclic-adenosine-monophosphate-2D-skeletal.png&usg=ALkJrhiOAvP0sstUz7VXae2GcM2-mzM6TQhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search?q=cAMP&hl=vi&biw=1517&bih=701&prmd=imvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/File:Cyclic-adenosine-monophosphate-3D-balls.png&usg=ALkJrhjBnALGflT_RMieH-LU-zV8B5j4JQhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search?q=cAMP&hl=vi&biw=1517&bih=701&prmd=imvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/File:Cyclic-adenosine-monophosphate-2D-skeletal.png&usg=ALkJrhiOAvP0sstUz7VXae2GcM2-mzM6TQhttp://en.wikipedia.org/wiki/Cyclic_adenosine_monophosphate


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Adenosine monophosphate vng (cAMP, AMP vng hoc 3'-5'-cyclicadenosinemonophosphate ) l mt tn hi th haiquan trng trong nhiu qu trnh sinh hc.cAMPc t ngun tadenosine triphosphate (ATP) v c s dng chotruyn tn hiuni o

nhiu t chc khc nhau, truyn t conng ph thuc cAMP. Chng hn nhchuyngiao vo cc tbo nh hng cakch thch tnhglucagon vadrenaline , m khng thiqua mng tbo.N c tham gia vo vic kch hot ccprotein kinase v iu chnh cctc ng ca adrenaline v glucagon. cAMP cng lin kt v quy nh v chc nng caccknh ion

cAMP c tng hp t ATPadenylyl cyclase nm pha n trong ca mng otng.Adenylyl cyclase c kch hot i mt lot cc phn t tn hiu thng qua kch hotcc kch thch cyclase adenylyl G (G )-protein-coupled th th v c ch i cht ch vnca adenylyl cyclase c ch G (Gi)-protein-coupledth th.Gan adenylyl cyclase phn ng mnh

m hn na glucagon, v c adenylyl cyclase phn ng mnh m hn na adrenaline.cAMP phn hy thnhAMP c xc tc i enzymephosphodiesterase .

cAMP v kinase lin quan n chc nng ca n trong cc qu trnh sinh ha, ao gmc cc quy nh caglycogen ,ng, vchuyn halipid .

sinh vt nhn chun, AMP vng hot ng ng cch kch hot protein kinase A(PKA,protein kinase ph thuc cAMP.) PKA l nh thng khng hot ng nh l mttetramericholoenzyme , ao gm haicht xc tcv hai n v qun l (C2R 2),vi cc nv quy nh ngn chn cc trung tm xc tc ca cc n v xc tc. AMP vng lin kt ncc a im c th v cc n v qun lca cc protein kinase, v gy phn ly gia cc tiu

n v qun l v xc tc, do kch hot cc n v xc tc v cho php h phosphorylateprotein cht nn.

Cc tiu n v hot ng xc tc chuyn giao phosphate t ATP serine hoc d lngc th threonine ca cht nn protein.Cc protein c phosphoryl ha c th hnh ng trctip trn cc knh ion ca t o, hoc c th kch hot hoc c ch enzym. Protein kinaseA cng c th phosphorylate protein c th lin kt vi vng promoter ca DNA, gy ra sgia tng ca gen c th.Khng phi tt c cc protein kinase p ng cAMP. Mt s cc lpca kinase protein, ao gm protein kinase C, khng ph thuc cAMP.

nh hng hn na ch yu ph thuc vo protein kinase ph thuc cAMP, thay i

da trn cc loi t o.Tuy nhin, c mt s chc nng c lp PKA nh ca cAMP, v d nh, kch hot ca

ccknh canxi , cung cp mt con ng nh mtng trng hormone gii phnghormone nguyn nhn pht hnh cahormone tng trng[1]

Tuy nhin, quan im rng phn ln nhng tc ng ca cAMP c kim sot biPKA l li thi.Nm 1998, mt gia nh ca cAMP-protein nhy cm viyu ttrao i
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guanine nucleotide(GEF) hot ng c pht hin. y c gi l protein Exchangekch hot bng cch cAMP (Epac) v gia nh ao gmEpac1 vEpac2 . C chkch hottng tnh ca PKA: min GEF thng c che y bi khu vc N-thit bu cui c

cha cc tn min cAMP rng buc. Khi cAMP lin kt, phn ly min v a ra hin nay hotng GEF min, cho php Epac kch hot nhRas ging nh protein GTPase,

Trongvi khun, mc ca cAMP thay i ty thuc vo phng tin c s dngcho s tng trng.c it, cAMP l thp khi glucose l ngun caron.iu ny xy rathng qua vic c ch enzyme cAMP sn xut, adenylyl cyclase, nh l mt tc dng ph cavn chuyn glucose vo t o.Cc yu t phin mprotein th th cAMP(CRP) cng gi lCAP (cataolite gen hot ha protein) to thnh mt phc tp vi cAMP v do c kchhot lin kt vi DNA.CRP-cAMP lm tng iu hin ca mt s lng ln gen, ao gmc mt s m hacc enzyme c th cung cp nng lng c lp ca glucose.

cAMP, v d, c tham gia vo cc quy nh tch cc caoperon lac . Trong mt mitrng c nng glucose thp, cAMP tch t v lin kt vi cc trang weallosteric trnCRP (cAMP th protein), mt phin m kch hot protein.Protein gi nh hnh dng hotng ca n v lin kt vi mt trang we c th thng ngun ca promoter lac, lm cho nd dng hn cho RNA polymerase rng uc cc promoter ln cn t u phin mca operon lac, tng t l sao chp lac operon.Vi nng glucose cao, nng cAMPgim, v CRP disengages t operon lac.
http://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/Guanine_nucleotide_exchange_factor&usg=ALkJrhigsCNzkn0QYs42DeS598jyMAtu6Qhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/RAPGEF3&usg=ALkJrhjXNZ8mawE4mj9WscEgEEQKSIdPQwhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/RAPGEF4&usg=ALkJrhjBlM7vAH6QhGQJ7Cs_4IQKCBdxoghttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/Bacteria&usg=ALkJrhg_FYHlT9tc-_yce1J41NduhJgtqAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/Bacteria&usg=ALkJrhg_FYHlT9tc-_yce1J41NduhJgtqAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/Bacteria&usg=ALkJrhg_FYHlT9tc-_yce1J41NduhJgtqAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/CAMP_receptor_protein&usg=ALkJrhjnfc6m_rCro9Hs4dIcHt0XtS7hlAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/CAMP_receptor_protein&usg=ALkJrhjnfc6m_rCro9Hs4dIcHt0XtS7hlAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/CAMP_receptor_protein&usg=ALkJrhjnfc6m_rCro9Hs4dIcHt0XtS7hlAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/Enzyme&usg=ALkJrhjhDe94YW-uofeoQZgsYmgSFKET4Ahttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/Lac_operon&usg=ALkJrhjAEcERs1GZVzdy9g74ab-q5fLt9ghttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/CAMP_receptor_protein&usg=ALkJrhjnfc6m_rCro9Hs4dIcHt0XtS7hlAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/CAMP_receptor_protein&usg=ALkJrhjnfc6m_rCro9Hs4dIcHt0XtS7hlAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/CAMP_receptor_protein&usg=ALkJrhjnfc6m_rCro9Hs4dIcHt0XtS7hlAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/CAMP_receptor_protein&usg=ALkJrhjnfc6m_rCro9Hs4dIcHt0XtS7hlAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/Lac_operon&usg=ALkJrhjAEcERs1GZVzdy9g74ab-q5fLt9ghttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/Enzyme&usg=ALkJrhjhDe94YW-uofeoQZgsYmgSFKET4Ahttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/CAMP_receptor_protein&usg=ALkJrhjnfc6m_rCro9Hs4dIcHt0XtS7hlAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/Bacteria&usg=ALkJrhg_FYHlT9tc-_yce1J41NduhJgtqAhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/RAPGEF4&usg=ALkJrhjBlM7vAH6QhGQJ7Cs_4IQKCBdxoghttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/RAPGEF3&usg=ALkJrhjXNZ8mawE4mj9WscEgEEQKSIdPQwhttp://translate.googleusercontent.com/translate_c?hl=vi&prev=/search%3Fq%3DcAMP%26hl%3Dvi%26biw%3D1517%26bih%3D701%26prmd%3Dimvnsz&rurl=translate.google.com.vn&sl=en&u=http://en.wikipedia.org/wiki/Guanine_nucleotide_exchange_factor&usg=ALkJrhigsCNzkn0QYs42DeS598jyMAtu6Q


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(Ngun:http://thuviensinhhoc.com/chuyen-de-sinh-hoc/sinh-ly-hoc-dong-vat/3125-cac-hormon-tac-dung-thong-qua-cac-chat-truyen-tin-thu-2.html)

- Lipit nhl cc tn hiu thhai

Hthng cc tn hiu thhai c nghin cu y lin quan n cc sn phm caqu trnh thy phn thnh photpholipit b (photphattidyllinositol (PI)) ca mng tbo. PI c

thchuyn ha thnh poliphotphattidylinositol (PIP2) bi enzim kinaza. Mc du PIP2trongmng tbo t hn nhiu so vi PI, n ng vai tr trung tm trong struyn tn hiu.

Trong cc tbo ng vt, hoocmon, v dnhvasoprexin lin kt vi cht nhn, nc tc dng hot ha G- protein dtam phn. Sau di n vphn li ra tG v hot ha

photpholipaza photphoinositit c hiu, photpholipaza C (PLC). PLC c hot ha nhanhchng thy phn PIP2, ti sinh cc sn phm inositol triphotphat (IP3) v diaxylglixerol(DAG). Mi mt phn ttrong hai phn ty c vai tr quan trng trong tn hiu tbo.
http://thuviensinhhoc.com/chuyen-de-sinh-hoc/sinh-ly-hoc-dong-vat/3125-cac-hormon-tac-dung-thong-qua-cac-chat-truyen-tin-thu-2.htmlhttp://thuviensinhhoc.com/chuyen-de-sinh-hoc/sinh-ly-hoc-dong-vat/3125-cac-hormon-tac-dung-thong-qua-cac-chat-truyen-tin-thu-2.htmlhttp://thuviensinhhoc.com/chuyen-de-sinh-hoc/sinh-ly-hoc-dong-vat/3125-cac-hormon-tac-dung-thong-qua-cac-chat-truyen-tin-thu-2.htmlhttp://thuviensinhhoc.com/chuyen-de-sinh-hoc/sinh-ly-hoc-dong-vat/3125-cac-hormon-tac-dung-thong-qua-cac-chat-truyen-tin-thu-2.html


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- Ion canxi nhl tn hiu thhai

Nng ion canxi trong tbo kim sot nhng qu trnh khc nhau kcqu trnhtrao i cht, tng sinh v cc chc nng bit ha nhtnh thm ca mng, vn ng co rt.Con ng tn hiu canxi lin quan n cc protein lin kt canxi m khi n c canxi hotha th lin kt vi protein bt hot v sau kch thch chng, bng cch nhvy n nhhng n slng cc con ng xui dng. Nng Ca2+trong tbo cht c duy tr mc thp nhsbi tit chng, nhng cho n 10000 ln cao hn trong dng ngoi bo vtrong mt sbo quan xc nh. Cc tn hiu ngc dng xut pht tcc cht nhn trn b

mt tbo gy ra dng canxi xm nhp vo tbo mcc knh canxi, iu lm tng lngcanxi lin kt ca canxi c hot ha v tng cc con ng tn hiu xui dng. C haicon ng kch thch s xm nhp ca canxi l cc cht nhn lin kt G- protein v chtnhn tirosin kinaza (RTK).Chai cht nhn ny kch thch enzim photpholipaza vn gia tnghm lng inositol-1, 4, 5-triphotphat (Ins[1, 4, 5]P3) (IP3) c tc dng m cc knh Ca

2+cng IP3(inositol-1, 4, 5-triphotphat) trong mng li ni sinh cht.


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Trong tbo cht c nhiu protein lin kt canxi. Mc du sln trong chng tc ngnhl cchm (btr) gim nng Ca2+trong tbo cht, mt shot ng nhccmy kim tra canxi v protein tn hiu. Annexin l nhng protein lin kt mng ph thuc

canxi, cc protein ny c th ti t chc thnh phn khung ca tbo; chng cng c chphotpholipaza A2. Trong cc protein ln lin kt iu tit canxi c calmodulin (CaM) vtroponin C. CaM c mt khp ni v l protein a chc nng; troponin C l protein chacanxi c c hiu v iu tit s tng tc gia myosin v actin khi co c. CaM hot hanhiu protein kinaza (CaM kinaza II, kinaza tc nhn ko di), photphataza (calcineurin) vcc thnh phn khung t bo, n cng hot ha Ca2+ -ATPase, nh vy n khi ng schuyn dch Ca2+ra khi tbo cht, CaM cng tng tc vi cc thnh phn ca cc h.


4. Struyn tn hiu trong cthc nhn

Nhiu vi sinh vt sdng tn hiu ha hc trong struyn tin gia cc tbo. V d,nm nhyDictyostelumi cm ng mt stbo tit ra cAMP. Cht ny khuch tn qua ccht v cm ng cc tbo ttp li. Sttp ca nm men l mt v dkhc vstruyntin ha hc gia cc tbo vi sinh vt. Tuy nhin, hng tnm vtrc, tn hiu tbo nh


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vy l bc tin phc tp khi cc tbo c nhn bt u lin kt vi nhau nhl cthabo, xut hin chiu hng chuyn ha cc tbo cng nhpht trin ca m v cquan hnh thnh nhng chc nng c hiu.

Vic iu phi spht trin v cm ng trli i vi mi trng ca cc cthabo phc tp i hi phi c nhiu cchtn hiu. Hai hthng tn hiu chyu c cthng vt l h thng thn kinh v hni tit. Thc vt khng di chuyn, khng c h thnkinh nhng li c cc hoocmon nhl cc tn hiu ha hc. L c thquang hp, thc vtcng c nhng c ch thch nghi s sinh trng v pht trin ca chng i vi mt nhlng v nh tnh ca nh sng.

a) Hai lp tn hiu xc nh hai lp cht nhn

Hoocmon c chia thnh hai lp da trn kh nng vn chuyn qua t bo cht:hoocmon a m vn d dng khuch tn qua lp ght nc ca mng sinh cht v cchoocmon tan trong nc vn khng c khnng xm nhp vo tbo. Hoocmon a mlinkt chyu vo cc cht nhn trong tbo cht hoc nhn, cc hoocmon a nc lin kt vicc cht nhn nh ctrn mng tbo. Trong mi trng hp, phi tlin kt lm thay icu hnh cht nhn bng cch gy nn sbin dng.

b) Cc cht nhn steroid hot ng nhlcc tc nhn phin m

Cc hoocmon steroid, hoocmon tiroit, hoocmon retinoit v vitamin D ddng i qua

mng sinh cht v chng l nhng cht ght nc v chng lin kt vi cc protein cht nhnni bo. Khi c hot ha bng cch lin kt vi cc phi t, nhng protein ny hot ngnhl cc tc nhn phin m. Tt ccc cht nhn steroid nhvy c min lin kt ADN.Cc thnh phn phn ng steroid c c trng nh ctrong cc min tng cng ca cc genkch thch steroid. Hu ht cc cht nhn steroid nh ctrong nhn, chng c neo vocc protein nhn dng mt hot tnh. Khi cht nhn gn vo steroid, n c tch ra khi

protein neo v trnn hot ng nhl cc tc nhn phin m. Sau cc tc nhn phin mc hot ha lin kt vo gen tng cng (enhancer) v kch thch sphin m. Khng linkt vi hoocmon, cht nhn hoocmon tiroit khng c khnng kch thch phin m.

Khng phi tt ccc cht nhn steroid ni bo nh ctrong nhn. Cht nhn i vihoomon glucocorticoit (coctisol) khc bit vi cc cht nhn khc ch n nh c trongxitosol (ccht ca tbo) v c neo vo protein xitosol trng thi mt hot tnh. Linkt vi hoocmon l cho cht nhn ri khi protein neo xitosol v sau phc hhoocmon-cht nhn di cvo nhn, n gn vo gen tng cng v kch thch sphin m. Miy, chng mnh c rng brassinosteroit l hoocmon steroid trong thc vt v gen i


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vi cht nhn brassinosteroit mi c chn dng v xc nh trnh tcc nucleotit. N mha kiu cht nhn xuyn mng gi l cht nhn giu ip khc loxin.

c) Cht nh

n b

m

t t

bo c th

t

ng tc v

i G- protein

Cc cht nhn b mt t bo c nghin cu tt ng vt c v. Tt c cchoomon tan trong nc ng vt c v lin kt vi cc cht nhn c trn bmt tbo. Ccthnh phn ca cht nhn bmt tbo tng tc vi cht truyn tn hiu l cc protein iutit lin kt GTP gi l cc G- protein dtam phn (heterotrimerric G- proteins). G- protein c hot ha, n lt mnh, hot ha enzim effector(enzim tc ng, enzim thc hin).Enzim effector c hot ha ti sinh ra tn hiu ni bo thhai vn kch thch nhiu qutrnh tbo.

Cc cht nhn sdng G- protein dtam phn (G- protein tam phn khc nhau) gingnhau v mt cu trc v khc nhau v chc nng, cu trc y ca chng ging vibacteriorhopsin, sc t mu ta lin quan vi quang hp trong t bo ca chi (genus)Halibacteriumv ging vi rhodopsin, sc tthgic ca mt ng vt c xng sng. Gny ngi ta xc nh c cc cht nhn khu gic ca mi ng vt c xng sngcng thuc nhm ny. Cc protein cht nhn tn ti dng chui xon by cu xuyn mng(seven transmembrane helices)(hnh 1.13). i khi ngi ta gi cht nhn y l cht nhnby cu, cht nhn by ln vt hay cht nhn setpentin.

(A)-Cc min lin kt phi tngoi bo ln c trng ca cc cht nhn by cu linkt protein. Cha r min ni bo tng tc vi G- protein dtam phn (G- protein gm , v di n v).

(B)-Cc min ngoi bo nhc trng ca cc cht nhn by cu lin kt vi cc phitb nhepinephrine. Mt scc chui xon n trong chui kp ca mng thng to nnvtr lin kt phi t.


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d) Chu trnh G- protein dtam phn lcu dao phn tca cc dng hot tnh vbt hot

G- protein truyn tn hiu t cc cht nhn by cu c tn gi l d tam phn(heterotrimeric). C tn nh vy v G- protein c ba di n v hp thnh: , v .Chng khc bit vi n phn G- protein. Chu trnh G- protein dtam phn gia dng hottnh v dng bt hot, hot ng nh l cu dao phn t. Cc di n v v to thnh

phc hgn cht G- protein vo mng pha tbo cht. Khi lin kt vi cht nhn by cuhot ha phi t, G- protein trnn hot tnh. dng bt hot, G- protein tn ti nhl mt

cu trc tam phn i vi GDP lin kt vo di n v .Khi lin kt vo phc h chtnhn- phi tn cm ng di n v chuyn h GDP thnh GTP. Schuyn ha y lmcho di n v phn li t v cho php lin kt vi enzim tc ng. Di n v chot tnh GTPaza. N c hot ha khi n lin kt vi cc enzim tc ng. Trong trnghp ny l enzim adenylyl-xiclaza (cn c tn l adenylat xiclaza) GTP bphn hy thnhGDP, bng cch n lm mt hot tnh di n v m n lt mnh n li lm bt hot


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adenylat xiclaza. Di n v lin kt vi GDP ti kt hp vi v v sau li chot ha bng cch lin kt vi phc hcht nhn- hoocmon.

e) Hot ha adenylat x iclaza lm t

ng l

ng cAMP

cAMP l phn ttn hiu quan trng ctrong cthtin nhn, trong tbo thc vt vng vt. Shot ha adenylat xiclazabi G- protein dtam phn lm tng nng cAMPtrong t bo. cAMP thng c duy tr mc thp bi tc ng ca cAMP

photphodiesterasevn thy phn cAMP thnh 5-AMP.

Trong cc tbo ni cAMP iu tit sbiu hin gen, enzim protein kinaza (PKA)photphorin ha tc nhn phin m gi l CREB (protein lin kt thnh phn cAMP phn ngtrli- cAMP response element- binding protein). Vi shot ha bi PKA, CREB lin kt

vi thnh phn phn ng trli cAMP (CRE- cAMP response element) nh cti cc vngkhi u ca cc gen vn c cAMP iu tit. V vy lng cAMP trong dch chit m thcvt qu thp nn vai tr ca cAMP trong con ng truyn tn hiu thc vt b tranh cinhiu (Assmann, 1995). Tuy nhin, ngy cng tch ly nhiu bng chng vvai tr ca cAMPtrng tbo thc vt. V d, ngi ta nh r c cc gen m ha CREB trong thc vt(Kategiri v CS., 1989) chng minh c s sinh trng ca ng phn hoa loa kn(Lilium) c cAMP nng thp, khong 10nM kch thch (Tezuka v CS., 1993). Li vCS. (1994) chcho thy cAMP hot ha cc knh K+trong mng sinh cht ca cc tbonhu m ca l cy u tm (Vicia faba). Mi y, Ichikwa v CS (1997) nhn din cc gen

c thca adenylat xiclaza trong cy thuc l (Nicotiana tabacum) v trong cyArabidopsis.Nhvy, mc cho c nhng nm thng nghi ng, vai tr ca cAMP nh l tc nhn tnhiu vn nng trong cc cthsng, kctrong thc vt r rng.

g) Shot ha photpholi paza Ckhi u con ng IP3, canxi nhl tn hiu thhaii vi sbin ng ln ca nhng skin tbo. Vi tr ny ca canxi c xc lpchc chn trong cc tbo ng vt cng nhthc vt. Nng Ca2+ tdo trong phn botan (cytosol) bnh thng c duy tr mc rt thp (1.10-7M). Ca2+-ATPaza trn mng sinhcht v trn li ni sinh cht (LNSC) bm cc ion Ca2+i ra v i vo khoang ca LNSC.

Trong t bo thc vt, hu ht canxi tp trung trong khng bo. Gradient proton in haxuyn qua mng khng bo c ti sinh bi cc bm proton c tc dng khi u con nghp thCa2+- H+vn chuyn canxi.

h) IP3mcc knh canxi vpha LNSC vpha mng khng bo

Photpholipaza C c hot ha ti sinh li IP3. Photpholipaza C tan trong nc vkhuch tn qua phn bo tan cho n khi gp c cc vtr lin kt IP3trn LNSC v trn


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mng khng bo. Nhng v tr lin kt ny l cc knh IP3 cng Ca2+ bn trong mc

milimol, canxi khuch tn nhanh vo phn bo tan theo gradient nng . Phn ng kt thckhi IP3bphn gii bi photphataza c hiu hay khi canxi c bm ra khi tbo cht bi

Ca2+

- ATPaza (bm ion canxi).

Tn hiu canxi thng bt ngun tmin nh cca n trong tbo v lan truyn nhsng truyn qua phn bo tan. Nhng song lp li c gi l dao ng canxic ththeo sautn hiu gc, mi t dao ng c thko di vi giy n vi pht.

Mi y, ngi ta pht hin c nhng dao ng sng nhvy trong cc tbokh khng ca thc vt (Mc Ainsh v CS., 1995).

i) Cc hthng hai thnh phn trong cthc nhn

Mi y, ngi ta pht hin ra trong cthnm men v thc vt c cc protein iutit sphi hp cm th- trli lin quan vi cc hthng hai thnh phn vi khun. V d,gen SLN1 ca nm men Saccharomyces cerevisiaem ha protein 134 kDa. Gen ny c cctrnh tging vi hai min truyn v min nhn ca vi khun v thhin chc nng trong siu tit thm thu (Ota v Varhavsky, 1993).

c bng chng rng mt sthc vt c hthng tn hiu gm tcc hthng haithnh phn ca vi khun. V d, sc tphitocrom hp thnh sng /xa c minhha khun lam (Cyanobacteria) v n lin hn cc protein cm thvi khun.

Ngoi ra, cc gen m ha cc cht nhn c ginh i vi hai hoocmon thc vt lxitokinon v etylen, chai cha cc min kinaza tphotphorin ha histidin cng nhcc muhnh iu tit trli tip gip.

5. Spht tn hiu

- Tr li tn hiu: Nhng tn hiu khc nhau t n bmt tbo c truyn vothc trnh thot tnh kinaza hoc sn xut ra tn hiu thhai. Cc tn hiu thhai tsdngtc ng ca chng bng cch bin i hot tnh ca kinaza tbo. Thc kinaza c protein

photphataza iu tit. Slng v hot tnh ca protein photphataza l i tng cho siutit phc tp.

-nh ngha: Spht tn hiu l on kt thc ca con ng truyn tn hiu ni ccthnh phn ca tbo vn l

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