giao trinh sinh ly thuc vat

B GIO DC V DOTODI HC S PHM THAI NGUYN PGS. Nguyn B Lc, PGS. Trng Vn Lung, TS. V Th Mai Hng, ThS. L Th Ha, ThS. L Th Tr! GIO TRINHSINH L HC THC VAT Thi Nguyn, "#$$ LI NI DAU Snh hc thc vt khoa hc snh hc nghn cu v cc hot ng sng ca thc vt. y mn khoa hc thc nghm v khoa hc co so cho cc ngnh khoa hc k thut nng nghp.Do ngha quan trng ca nh vc khoa hc ny cho nn t khra vo cu the k XVIII en nay n uoc pht tren nhanh chng vc nhu ng gp to n cho khoa hc cng nhu cho sn xuat v sng con ngu.Snh hc thc vt khoa hc uoc gng dy o cc trung hc hng tram nam nay. Cng c nhu go trnh Snh hcthc vt uoc vet phuc vu cho vc gng dy, hc tp v nghn cunh vc khoa hc ny.O Vt Nam Snh hc thc vt cng uoc gng dy o nhutrung hc (HSP, HKHTN, HNL ...) v cng c nhu gotrnh Snh hc thc vtuoc pht hnh.Trn co so nhng go trnh hn c, e c tu u hc tp, nghn cu cho snh vn, truc het snh vn ca hc Hue, chng t bn son go trnh Snh hc thc vtny. Sch uoc dng m gotrnh cho snh vn cc khoa Snh HSP, HKH v HNL thuc hc Hue v m t u tham kho cho snh vn, cn b cc ngnh n quan.Go trnh do mt tp the cc nh Snh hc thc vt o H Huebn son do PGS.TS. Nguyn B Lc ch bn v bn son cc Chuong4, Chuong 5, Chuong 7. PGS.TS. Truong Van Lung bn son Chuong 2,ThS. L Th} Tr bn son Chuong 1, ThS. L Th} Hoa bn son Chuong6. ThS. L Th} Ma Huong bn son Chuong 3.Trongqutrnhbnson, tpthetcgcgangcpnhtnhng ken thc hn v thc tn vo. Tuy nhn, do th gan, trnh, nguon tu u c hn nn khng trnh kho nhng theu st. Chngt mong nhn uoc s gp ca c g e n t bn sau go trnhc chat uong tt hon. Tha nguyn, thng 5 nam 2011 Cc tc g M U I. i tng, ni dung v nhim v ca Sinh l hc thc vt .!.i tng ca Sinh l hc thc vt "S#$%&'.Snh hc thc vtnghn cu hot ng sng ca thc vt chonn tuong nghn cu ca Snh hc thc vt co the thc vt.Khc v ng vt, thc vt snh vt t dung nn hot ngsng c nhng ac trung rng do vy vc nghn cu hot ng sngca thc vt c nhng ac trung khc v o ng vt.(. )i dung ca Sinh l hc thc vt.Snh hc thc vt mt khoa hc nghn cu v cc qu trnh sng trong co the thc vt. qu trnh nhn vt chat v nang uong t m trung ngo vo co the e chuyen ho chng thnh vt chat, nang uong ca co the nhm ken to nn co the, gp cho co the snh truong v pht tren. Ou trnh hot ng uoc the hn qua cc chc nang snh ca thc vt trao o nuc, dnh dung khong, quang hop, h hap, snh truong v pht tren.*. )him v ca Sinh l hc thc vt.Nhm vu ca Snh hc thc vt pht hn ra nhng qu ut ca cc hot ng snh dn ra trong co the thc vt. Nghn cu bn chat hc, ho hc v snh hc ca cc hot ng sng . ong th Snh hc thc vt cng nghn cu nhng tc ng ca cc nhn tsnh th (nh sng, nuc, nht , chat khong, chat kh ...) en cc hot ng sng ca thc vt.Muc tu cu cng ca Snh hc thc vt phuc vu cho vc c to thc vt theo muc tu ca con ngu nhm to nhu sn phmthu nhn t thc vt phuc vu cho nhu cu cuc sng ca con ngu ngy cng cao. Snh hc thc vt co so khoa hc ca cc bn php k thut tc ng vo thc vt nhm nng cao nang suat v c thn phm chat ca chng theo muc ch ca con ngu.II. Mi li+n ,uan gi-a Sinh l hc thc vt v.i c/c 0h1a hc 0h/c.Snh hc thc vt mt khoa hc thc nghm. Truc het Snh hc thc vt n quan en cc khoa hc co bn nhu hc, ho hc.Snh hc thc vt su dung cc phuong php, cc ken thc ca hc, ho hc e nghn cu trn tuong thc vt, do vy ten vk thut, v phuong tn nghn cu hc, ho hc c va tr quantrng trong s pht tren ca Snh hc thc vt.Trong snh hc, Snh hc thc vt c m quan h chat ch vnhu nh vc chuyn mn khc nhu Ho snh hc, L snh hc, Thcvt hc, Te bo hc, Snh th hc ... Nhu ket qu nghn cu caSnh hc thc vtda vo nhng thnh tu ca cc ngnh khoa hctrn.Tr Snh hc thc vt cng gp phn pht tren cc ngnhkhoa hc .Snh hc thc vt mn khoa hc co so cho cc ngnh khoahckthutnngnghpnhu: trongtrt, msnh, boqunnngsn ... nn un ca Snh hc thc vt gp phn pht tren ccngnh khoa hc .III. #c 23 4h/t t5i6n ca Sinh l hc thc vt.Snh hc thc vt mt mn khoa hc ra mun so v nhu khoa hc snh hc khc nhu phn o hc, g phau hc ...Cu the k XVIII, Snh hc thc vt ra kh cc nh khoa hcpht hn ra qu trnh quang hop, h hap ca thc vt (Presey-1771,Ingenhous, Seneber-1782, De Sanssure-1801 ...).Tuy nhn, truc nhu van v hot ng sng ca thc vt cng uoc mt s nhkhoa hc nghn cu mt cch e te.Sang the k XIX, nh nhng ten b v phuong tn v phuongphp nghn cu ca vt , ho hc gp phn cho Snh hc thcvt hon thn dn. Cc hc thuyet v quang hop, h hap, dnh dungkhong, trao o nuc ngy cng su vo bn chat v co che. nhng ng gp to n ca cc nh khoa hc nhu Lebg v dnh dungkhong (1840), Krgov v enzme (1810), Mayer v quang hop, Pasterv n men (1880), Pfeher v tham thau (1877), Vnogratxk v c }nhm t do ...ac bt quan trng nhng cng trnh nghn cu mt cchton dn, c h thng ca Tmradep v quang hop, h hap ... mcho Snh hc thc vt tro thnh mt khoa hc c p. C the xemTmrazep ngu sng p ra khoa hc Snh hc thc vt.Sang the k th XX, v s pht tren mnh m ca khoa hc,Snh hc thc vt cng pht tren nhanh chng. Nh nhng thet b}nghncungycnghn, ccphuongphpnghncungycng hon thn nn Snh hc thc vt cng c u kn su vobn chat, co che cc hot ng sng ca thc vt m cho n dungSnh hc thc vt ngy cng phong ph.Song song v vc su nghn cu co che cc hot ng sngca thc vt, cc nh Snh hc thc vt cn tptrunggquyetnhng van n quan en thc tn sn xuat, gp phn quan trngthc y tang nang suat cy trong.Tm , }ch su pht tren Snh hc thc vt gan n v s tenb ca cc ngnh khoa hc khc ac bt hc v ho hc v ngycng pht tren mnh m gp phn vo vc pht tren chung ca ccngnh khoa hc v s sng v thc y thc tn sn xuat. Chuong I SINH L T BO THC VTI. Khi nim t bo. 1. Hc thuyt t bo.T% &' (' )n * c +, -' ./. c0 cc c .h1 +2ng )3u h4nh .h'nh nn .5 )6. N- $778, 9&:r. H; )< =h. hi>n ra )n * c/u .r?c c +, c@a c .h1 +2ng (' A.%&'B. Cng )< -D .0 c/u .r?c )6. EFng .hGi *' )c (H= *Ii 9&:r. H;, nh' &c hJc H'Lan Kn.ni: Van L::uL:nhc; *' ngGi M Ma(=ighi )< nghin cNu , )2i .Ong )ng *H.*' cPng =h. hi>n ra .% &'.E%n .h% ;Q RSR, *Ii +T )6ng g6= c@a nh' .hTc *H. hJc Ma.hia+ Sch(:iU:n *' nh')ng *H. hJc Th:Ur SchLann hJc .huy%. .% &' chVnh .hNc ra )Gi W$XYXZ.2. D}c trung chung cua t bo.2.1.D}c trung v cu to.Th: Ma.hia+ Sch(:iU:n *' Th:Ur SchLann .h4 -Ji c .h1 .hTc *H. *' )ng *H.)3u U nh[ng .% &' c/u .\ nn *' ch?ng )Oc +]= ^%= .h: nh[ng .rH. .T ring )_c.rng ch .5ng c .h1. T/. c0 cc & =hHn c@a n6 )3u )\. )%n -Nc chuyn h6a *3 h4nh.hi *' chNc nng. E6 (' ;%. ù0 c@a c0 -. ù .r4nh .i%n h6a h%. +Nc (au U'i c@a ccU\ng +2ng nguyn .h@y, .hVch nghi ca ) *Ii cc )i3u ;i>n -Di .rGng =hNc .\= *' )aU\ng.MJi .% &' )3u c6 c/u .\ c &0n nh +aubcMJi.%&')3uc6-'ng+inhch/. &aùanh.Trn-'ngc6nhi3u;nhUdn.ruy3n *H. ch/. *' .hDng .in .\ ceu n2i gi[a .% &' *' -Di .rGng &n ng'i.cMJi .% &' )3u c6 nhan h_c nguyn (i>u nhan chNa .hDng .in Ui .ruy3n .% &'. f6*gng nhan )nh hIng *' )i3u .i%. -Ji h\. )ng c@a .% &'.cMJi .% &' )3u chNa ch/. n3n gJi (' .% &' ch/.. T% &' ch/. chNa cc &' ùan.2.2. D}c trung v chc nng.MJi h\. )ng +2ng c@a c .h1 cPng )Oc .hTc hi>n .5 -Nc ) .% &'.c Tra )hi ch/. *' nng (Ongb Gi[a c .h1 +inh *H. *' -Di .rGng (uDn (uDn ^0y raù .r4nh .ra )hi ch/. *' nng (Ong. NhG .ra )hi ch/. *' nng (Ong -' c .h1 .Fn .\i,+inh .r,ng *' =h. .ri1n.c Sinh .r,ng *' =h. .ri1nb Sinh .r,ng (' h> ù0 c@a ù .r4nh .ra )hi ch/. *'nng (Ong. Sinh .r,ng (' +T .Vch (Py *3 (Ong ('- ch ;h2i (Ong *' ;Vch .hIc .ng(n. ihi +inh .r,ng )\. )%n ngjng nh/. )nh .h4 c .h1 chuy1n +ang .r\ng .hi =h..ri1n. Ph. .ri1n (' +T &i%n )hi *3 ch/. (Ong c@a c0 c/u .r?c (dn chNc nng +inh (k c@a c.h1 .h: .5ng giai )\n c@a c .h1.c Sinh +0nb Sinh +0n (' .huc .Vnh )_c .rng nh/. ch c .h1 +2ng. NhG +inh +0n -'c .h1 +2ng .Fn .\i, =h. .ri1n .5 .h% h> n'y ùa .h% h> ;hc, c .h1 .hTc hi>n )Oc c ch%.ruy3n )\. .hDng .in Ui .ruy3n .5 .h% h> n'y ùa .h% h> ;hc. Sinh +0n (' )_c .Vnh ùan.rJng nh/. c@a c .h1 +2ng -' *H. .h1 ;hDng +2ng ;hDng c6 )Oc. Sinh +0n .h: ;i1u .rTc=han hay U cc .% &' chuyn h6a )0- nhHn.Nh *Hy -Ji h\. )ng +2ng c@a c .h1 )Oc .hTc hi>n .5 -Nc ) .% &'. VHy .%&' *5a (' )n *c/u .r?c *5a (' )n * chNc nng c@a -Ji c .h1 +2ng.II.Thnh phn ha hc cua t bo.1. Cc cht v co.lua +T =han .Vch c@a cc nh' ;ha hJc, ch/. +2ng .rung &4nh c6 ;h0ng 8mc XmnnIc, $#c $"n =r.iU:, "c Yn (i=iU:, $n g(uciU: *' gen $n -u2i *' cc hO= ch/. ;hc.1.1. Nuc. NIc (' .h'nh =hen ch@ y%u c@a ch/.nguyn +inh, n6 c6 *ai .ro ùan .rJng ;hDngnh[ng .rng *i>c hoa .an cc ch/. Uinh Ujng -' con (' -Di .rGng )1 .i%n h'nh cc (\i=h0n Nng h6a +inh, n6 con )i3u hoa nhi>. ) c .h1, .ha- gia *' ù .r4nh *Hn chuy1ncc ch/. .rng c .h1p *4 *Hy n6 c6 k ngh!a (In. LOng nIc .rng .% &' .hGng (' -.chq .iu *3 -Nc ) h\. )ng +2ng c@a .% &'. fhrng h\n, , -D nn .Ong .h@y h6a. Hi>n .Ong .h@y h6ac6 0nhh,ng r/. (In )%n h\. )ng +2ng c@a .% &'.Trng ch/. nguyn +inh, nIc .Fn .\i , hai U\ngb nIc (in ;%. *' nIc .T U.NIc .T U chi%- heu h%. (Ong nIc .rng .% &' *' c6 *ai .ro ùan .rJng .rng .ra )hich/. WTEfZ. NIc (in ;%. chi%- uc mn .hng (Ong nIc. NIc (in ;%. .hGng ;%. hO=*Ii nh6- a nIc c@a =r.:in &vng ceu n2i hyUrg:n. H'- (Ong nIc (in ;%. (In .h4;h0 nng ch2ng chu c@a ch/. nguyn +inh )2i *Ii ng\i c0nh &/. (Oi ca.1.2. Cc cht khong. Ng'i nIc, .rng .% &' con chNa nhi3u ch/. *D c ;hc (' cc nguyn .2 ;hng,(Ong chNa c@a .5ng nguyn .2 ;hng .rng ch/. +2ng ;hc &i>. nhau r/. nhi3up ng'inh[ng nguyn .2 )\i (Ong con c6 nh[ng nguyn .2 *i (Ong, +iu *i (Ong. fh?ng ,U\ng cc -u2i *D c Wif(, Naf(, faf("...Z, cc aciU WHf(, HYPwu...Z, cc (\i ;i3- WNHY,NH"wH...Z. Trng .% &', cc ch/. ;hng .hGng .Fn .\i UIi U\ng cc in .T U nhHfwYc, fwYc, NwYc, Nw"c, H"Pwuc, HPwuc, Swuc, f(c, Hx, faxx, ix, Mgxx, Nax, y:xx, ... haych?ng )Oc h?. &- .rn cc g2c -ang )i>n c@a cc -i^:n ;: h_c c6 -_. .rng .h'nh=hen cc hO= ch/. h[u c ;hc W(in ;%. h6a hJcZ. fh/. ;hng , .r\ng .hi .T U ùy)nh = +u/. .hz- .h/u c@a .% &' .5 )6 g6= =hen *' c ch% h/= .h{ nIc, cc ch/.;hng c@a .% &'. ST =han &2 ;hDng )Fng )3u c@a -. +2 in ;hng , hai &n -'ng+inh ch/. (' c +, c@a +T û/. hi>n .h% hi>u -'ng *' Uong )i>n +inh hJc. fc ch/. ;hng, U\ng h?. &- .rn &3 -_. cc h\. ;: n6 gi[ .rng .r\ng .hi &3n *[ng, -Nc ) =han.n, ) ngH- nIc, ) nhI. nh/. )nh c@a h> .h2ng ;: WSn h6a .r $, nh i .hGng ('-.ng ) ngH- nIc, ) =han .n *' gi0- ) nhI., con in h6a .r " nh fa *' in h6a .rY nh K( c6 0nh h,ngngOc (\iZ. fc nguyn .2 ;hng c6 .c U{ng )i3u .i%. cc h\. )ng +2ng U 0nh h,ng +au+]c )%n cc h> :n|y-:. fc nguyn .2 *i (Ong .hGng (' .h'nh =hen c/u .r?c &]. &ucc@a cc h> :n|y-:. Ng'i ra cc ch/. ;hng con (' .h'nh =hen c@a h'ng (\. ch/. h[uc ch@ y%u c@a .% &' +2ng nh =r.iU:, nuc(:ic aciU, (i=iU...1.3. Cc cht kh.fc ch/. ;hV w", fw" (' cc y%u .2 +2ng con c@a c .h1, n%u .hi%u cc ch/. )6, nh/.(' w" .h4 ;hDng .h1 c6 +T +2ng.w^y (' ch/. ;hV c@a +T +2ng, w" cen ch hD h/= .% &', .\ nng (Ong cen ch c.h1 h\. )ng. fw"(' nguyn (i>u ch ù .r4nh ùang hO=, ;hDng c6 fw" .h4 ;hDng c6 +inh *H.+0n û/.,+inh *H. .T Ujng +} ;hDng .Fn .\i, Uen Uen -Ji +inh *H. ;hc cPng +} & Ui>.*ng *4 ;hDng c6 fw", cay ânh ;hDng chuy1n )Oc nng (Ong -_. .rGi .h'nh nng(Ong h6a hJc.2. Cc cht hu co. Trng .% &' c6 r/. nhi3u (\i ch/. h[u c ;hc nhau, -~i (\i c6 chNc nng chuynh6a )_c .rng. Trng )6, ùan .rJng nh/. (' cc ch/. =r.:in, nuc(:ic aciU, g(uciU:, (i=iU:.T5 &2n ch/. h[u c cn &0n n'y, .5 )6 h4nh .h'nh nn cc ch/. nh :n|y-:, hr-n:,*i.a-in, +]c .2, ch/. .h-... V' cPng chq .5 &2n (Jai ch/. )6 -Ii c6 +T .ha- gia *' ù.r4nh chuy1n h6a *' cung c/= nng (Ong ch c .h1. fc ch/. n'y con )Oc gJi (' cc=han .t +inh hJc.2.1. Protein.Trng +2 ccch/. h[u c, =r.:in (' .h'nh =hen ùan .rJng nh/.. 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ST ;hc nhau *3 .h'nh =hen, +2 (Ong *' .rH. .T +]= ^%= cc a-in aciU .\ nn +T)a U\ng c@a =r.:in, .5 )6 .\ nn .Vnh )a U\ng c@a +inh giIi.f/u.r?cc@aa-inaciU)Oc)_c.rng&,i hainh6-chVnhbNh6-far&^y(cfwwH *' nh6- a-inc NH", =hen con (\i (' g2c W9Z c6 c/u .r?c ;hc nhau ,cc a-inaciU ;hc nhau. f/u .\ .hng ù. c@a a-in aciU nh +aub fc a-in aciU (in ;%. *Ii nhau &vng (in ;%. =:=.iU:, .\ nn chuhi =(y=:=.iU: ('c/u .r?c &Hc S c@a =r.:in.TVnh ch/. )a U\ng c@a =r.:in con gia .ng (?c .\ .h'nh cc -Nc ) c/u .r?c =hNc.\= hn Wc/u .r?c &Hc SS, &Hc SSS *' &Hc SVZ nhG cc (in ;%. ngang ;hc nhau. ii1u ^%=cun c@a -\ch ^]n Wc/u h4nh ;hDng gianZ cPng c6 .Vnh )_c .hg )2i *Ii .5ng (\i =r.:in.Pr.:in c6 ;h0 nng U U'ng .\ nn cc h4nh .hNc (in ;%. ;hc nhau *Ii cc ch/.*D c *' h[u c U -\ch &n c@a ch?ng c6 nhi3u nh6- )nh chNc ;hc nhaunh nh6-anIcWcfwwH, cwH, cfHw, cfw, cNH", NH, cfwNH",cSHZp nh6-ghs.nIcWfHY , fH" , fYH8 , nhan .h-...Zp nh6- c6 .Vnh ch/. aciU h_c &a+:, nh6- -ang )i>n .VchUng WNHxZ hay a- Wfwwc Z. ;h0 nng =h0n Nng ca nn =r.:in .hGng , U\ng =hNc hO= *Ii cc ch/. h[uc ;hc W(i==r.:iU, nuc(:c=r.:iU, =h+=hr=r.:iU, g(uc=r.:iUZ, =r.:in )6ng *ai .ro(' c +,, (' & +Gn c/u .r?c .inh .% c@a .% &' nh/. (' c/u .r?c cc h> .h2ng -'ng *' c/u.r?c ni .\i c@a cc &' ùan.Pr.:in con c6 *ai .ro )i3u .i%. cc ù .r4nh .ra )hi ch/.. fc h> :n|y-: )3u c6&0n ch/. h6a hJc (' =r.:in. Nh= ) ù .r4nh +inh .r,ng, =h. .ri1n, cGng ) *' chi3uhIng cc ù .r4nh .ra )hi ch/. c@a .% &' n6i ring *' c .h1 n6i chung )3u c6 (inùan .rTc .i%= *Ii +T .hng hO= *' h\. .Vnh ^?c .c c@a :n|y-:.Pr.:in c6 k ngh!a (In )2i *Ii ù.r4nh h?. nIc *' -u2i ;hng W $ga- =r.iU:(in ;%. ^/= ^q #,Y ga- nIcZ. Pr.:in ;han nIc c6 .h1 AcI= nIcB *Ii nh[ng (Tc r/.(In. B,i *Hy ) a nIc c@a =r.iU:, ù .r4nh .rng =hFng c@a ;: =r.iU: c6 0nhh,ng ùan .rJng )%n ù .r4nh .ra )hi nIc. Pr.iU: c6 .h1 (in ;%. c0 anin (dn ca.inc@a -u2i ;hng U .Vnh ch/. (jng .Vnh *3 )i>n c@a n6 W=han .t =r.:in chNa nhi3u g2ca-in WNH"Z*' car&^y( WfwwHZ .T U , -\ch &n nn c6 .h1 =han (y .rng Uung Uch.h'nh cc g2c -ang )i>n.Ng'i cc chNc nng .rn, =r.:in cPng c6 *ai .ro (' nguFn cung c/= nng (Ongch .% &'. Nng (Ong )Oc gi0i =h6ng (?c ^y h6a cc a-in aciU .rng .rGng hO=.hi%u g(uciU: *' (i=iU:, n6 )Oc +t U{ng )1 Uuy .r4 cc h\. )ng +2ng c@a .% &'. T/. c0nh[ng )_c )i1- *' .Vnh ch/. )6 c@a =r.:in gi0i .hVch )Oc =r.:in (' c +, *H. ch/. c@acc ù .r4nh +2ng.2.2. Lipide.Trng .% &', (i=iU: hJ= .h'nh nh6- ;h (In nh -j,Ueu, +=, =h+=hr(i=iU:,g(uc(i=iU:, +.:riU. fh?ng (' nh[ng hO= ch/. h[u c ;hDng .an .rng nIc, chq .an .rngcc Uung -Dih[u c nh :.h:r, ch(rr-, &:n|:n:, .(u:n:...Li=iU: c6 *ai .ro ùan .rJng .rng c/u .r?c .% &', )_c &i>. (' -'ng nguyn +inh,=h+=hr(i=iU: (' (i=iU: =hNc .\= c6 chNa =h+=hr (' .h'nh =hen c@a -'ng nguyn +inh*' nhi3u c/u .r?c ùan .rJng ;hc c@a .% &'. Li=iU: con (' ch/. cung c/= nng (Ongùan .rJng c@a .% &'.2.3. Glucide.G(uciU: con gJi (' +acchariU: (' hO= ch/. h[u c r/. =hh &i%n .rng c .h1. Th'nh=hen nguyn .2 c@a g(uciU: chq chNa f, H, w. .rng )6 +2 nguyn .t H (uDn g/= )Di w. G(uciU: )6ng *ai .ro (' ch/. UT .r[, )Oc +t U{ng nh -. nguyn (i>u .\ h4nh *'nng (Ong. M. =hen g(uciU: .ha- gia ây UTng ch/. +2ng, (Ong (In )Oc +t U{ng )1.\ .h'nh -'ng .% &', .rng )6 cen (u k )%n c:((u(+:, h:-ic:((u(+:, =:c.in.2.4. Mt s cht khc. Ng'i cc nh6- h[u c cn &0n nu .rn, .rng .% &' con c6 r/. nhi3u ch/. h[u cùan .rJng ;hc, -~i ch/. c6 c/u .\ *' chNc nng )_c .rng. Nh +]c .2 c6 *ai .ro ùan.rJng .rng ùang hO= c@a cay ânhp hr-n:, *i.a-in c6 *ai .ro ùan .rJng .rng )i3uhoa .ra )hi ch/.cnng (Ong *' h\. )ng +2ng c@a c .h1p cc +0n =hz- .rung gian c@aù .r4nh .ra )hi ch/. c@a .% &'.VHy .% &' +2ng (' ;h chNa *D +2 cc nh6- hO= ch/. c6 c/u .r?c, .Vnh ch/. *' kngh!a +inh hJc ;hc nhau, nhng ch?ng c6 -2i ùan h> ch_. ch} c0 *3c/u .\ (dn chNcnng, )_c &i>. .rng chNc nng .ra )hi ch/.c nng (Ong .rng .% &'.III. Cu to v chc nng cua t bo.1. D}c trung cu to cua t bo thc vt.T% &' (' )n * c/u .r?c c@a -Ji c .h1 +2ng *' cPng con .h1 hi>n nguFn g2cchung c@a +inh giIi. T% &' )ng *H. *' .hTc *H. c6 nhi3u )i1- gi2ng nhau, nhng &nc\nh +T gi2ng nhau, +T ;hcnhau c@a hai (\i .% &' .h1 hi>n +T =han h6a *3 chNc nngUdn )%n =han h6a *3 c/u .r?c &0 )0- .Vnh .hVch nghi c@a +inh giIi.Gi[a .% &' )ng *H. *' .% &' .hTc *H. c6 -. +2 -_. ;hc nhau U chNc nng;hc nhau .\ ra. f6 u +ai ;hc ch@ y%ub c T% &' )ng *H. c6 .rung .t, .% &' .hTc *H. ;hDng c6.c T% &' .hTc *H. c6 ({c (\=, .% &' )ng *H. ;hDng c6.c T% &' .hTc *H. c6 *ch .% &', .% &' )ng *H. ;hDng c6.c T% &' .hTc *H. c6 ;hDng &', .% &' )ng *H. ;hDng c6.2. Mng t bo.2.1. Mng cellulose. M'ng c:((u(+: chq c6 , .% &' .hTc *H., (' -'ng &0 *>, con gJi (' *ch .% &'. TrIc)ay ngGi .a ch *ch .% &' ('-. c/u .r?c ;hDng +2ng. Nay, .h'nh =henh6a hJc c@a-'ng &0 *> )< )Oc =han .Vch, ;h =hNc .\=, nIc chi%- 7#n )Oc chNa .rng cc;h0ng .T U c@a -'ng, Y#n c:((u(+:, cc +Oi c:((u(+: (in ;%. *Ii nhau .\ .h'nh cc-i^:n W;h0ng $## +Oi c:((u(+: &>n (\i *Ii nhau .\ nn -. -i^:n *Ii ;Vch .hIc mn-,cN "# -i^:n ;%. *Ii nhau (\i .\ nn -. +Oi &s W-icri&rinZ VIi ;Vch .hIc ;h0ng $#c"# n-, *' cN "m# +Oi &s (\i .\ nn +Oi (In W-acri&rinZ. fc +Oi )an chs *Ii nhau .h:nhi3u hIng ('- ch -'ng c:((u(+: r/. &3n *[ng, nhng (\i c6 ;h0nng )'n hFi. gi[a cc +Oi (' ;h2i ;hDng gian chNa cc ch/. *D )nh h4nh gF- :-ic:((u(+:, =:c.in *'nIc.NhG c/u .r?c .rn, -'ng c:((u(+: *5a &3n *5a -3- U .hVch Nng *Ii chNc nng&0 *> c@a n6. M'ng n'y )< gi?= ch .% &' c6 h4nh U\ng hn )nh. fc .ia +inh ch/. c@a-'ng *' cc :n|y-: .rn -'ng .\ ra nh[ng =h0n Nng .ng h~ =hNc .\= .ha- gia *'*i>c =han gi0i cc ch/. ;h6 .an .h'nh ch/. U .an, h_c ch?ng (' ch/. ^?c .c c@a =h0nNng gi[a -Di .rGng*' .% &'.2.2. Mng nguyn sinh cht.M'ng nguyn +inh ch/. con gJi (' -'ng ng\i ch/., (' -'ng &a &Jc ;h2i +inh ch/.c@a .% &' , -Ji c .h1. Th'nh =hen *' c/u .r?c c@a -'ng nguyn +inh ;h =hNc .\=, UhO= ch/. (i==r.:in c/u .\ nn. f6 nhi3u + )F gi0i .hVch c/u .r?c -'ng nguyn +inhnhng )3u chung -. nguyn (k (' -'ng nguyn +inh c6 c/u .r?c Y (I=p " (I= =r.:in *'$ (I= (i=iU:. Trn -'ng c6 nhi3u (~ nh *Ii )Gng ;Vnh ;h0ng #,X n-. fc + )F ;hcnhau chq nu ra cch +]= ^%= ;hc nhau c@a cc (I= )6. NhG c/u .r?c .rn ;hi%n -'ng c6 .c U{ng (In .rng *i>c &0 )0- .Vnh &n .h/- *';h0 nng .h/- c6 chJn (Jc c@a .% &' +2ng )2i *Ii cc ch/. ;hc nhau. M'ng nguyn+inh('=hen+inhch/. c6;h0nng.ra)hi ch/. r/. -. *4 n6chNanhi3uh>:n|y-:, )_c &i>. (' :n|y-: .h@y =han. Ng'i ra -'ng nguyn +inh con ('- nhi>- *{.ruy3n )\. .hDng .in .5 .% &' n'y +ang .% &' ;hc.3. T bo cht v cc bo quan.3.1. T bo cht.T% &' ch/. (' ;h2i ch/. +2ng nv- .rng -'ng nguyn +inh ch/., &a ùanh cc&' ùan c@a .% &'. T% &' ch/. ;hDng =h0i (' -. ;h2i c/u .r?c )Fng nh/., -' c6 c/u.r?c U .h1, .rng )6 c6 chNa cc .h1 *gi Wcc giJ. Ueu, cc h\. .inh &.Z, cc )\i =han .t=r.:in , cc +Oi K9N... fh/. ;hD c@a .% &' ch/. c6 ;h0ng 8mn =r.:in )n gi0n *'=hNc .\= WNuc(:=r.:in, G(uc=r.:in, Li==r.:in...Z $mc "#n (i=iU:. Trng .% &' ch/.con chNa nhi3u h> :n|y-: .ha- gia ù .r4nh .ra )hi ch/..3.2. Cc bo quan.c Ty .h1 f6 h4nh U\ng ;Vch .hIc *' +2 (Ong .hay )hi .gy .h: .% &' *' .gy .huc *' .hGi; +inh .r,ng c@a c .h1. Ty .h1 c6 U\ng h4nh ù:, h4nh +Oi, h4nh h\., h4nh .hi. S2 (OngD}ch n bo.y .h1c@a cc .% &' r/. ;hc nhau, c6.h1 .5*'i )%n *'i .r-.y .h1 .rng -. .% &'. .% &' c6 ù .r4nh .ra )hi ch/. -\nh, +2 (Ong .y .h1 r/. ca. Ty .h1 c6 .h1 Ui chuy1n.rng .% &' )%n *gng c6 ù .r4nh .ra )hi ch/. -\nh )1 .hTc hi>n chNc nng c@a n6.Th'nh =hen =r.:in c@a .y .h1 chi%- 7mc 8mn, (i=iU: "#c Y#n, K9N $n, KN#,mn, G(uciU: $n, y:, fu... Trng .y .h1 chNa nhi3u h> :n|y-:, nh :n|y-: .rng chuhihD h/=, .rng chu .r4nh fr:&+, cc :n|y-: .rng ù .r4nh .ra )hi ch/., nuc(:ic aciU *'=r.:in.f/u .r?c c@a .y .h1r/. =hNc .\=. Ba ng'i (' -'ng c +, c6 " (I=, (I= ng'i .\.h'nh -_. nhn c@a .y .h1, (I= .rng cun gG (n .h'nh ./- rng (Oc. Trn ./- rng (OcchNa nhi3u h> :n|y-: .ha- gia *' .ra )hi ch/. *' nng (Ong.Gi[a hai (I= -'ng (' ;h2i c ch/. U'y Xc $# n-,.rn )6 cPng chNa nhi3u (\i:n|y-:.Trn ./- rng (Oc (\i -ang cc h\. nh gJi (' ^y^- c6 )Gng ;Vnh Xc $# n-.fc ^y^- , -'ng .rng c6 chan ng]n " n- g]n *' -'ng, cc h\. , -'ng ng'i g]n.rTc .i%= *' -'ng, ;hDng c6 chan.fhNc nng c@a .y .h1 ch@ y%u .ha- gia *' ù .r4nh hD h/=, (' ni Uin ra chu.r4nh fr:&+, chuhi hD h/=, =h+=hry( h6a. Ty .h1 (' .r\- nng (Ong ch@ y%u c@a .% &'.fhNc nng c@a n6 (' gi0i =h6ng .ri>. )1 nng (Ong chNa )Tng .rng nguyn (i>u h[u c*' chuy1n h6a.h'nhU\ngnng (Ong.i>n U{ngWKTPZ.fhNcnng c@a .y .h1 Uin ra.rng Y nh6- ù .r4nh (in ùan -H. .hi%. *Ii nhau.x fc =h0n Nng ^y h6a cc nguyn (i>u W.rng chu .r4nh fr:&+Z, .\ ra cc +0n=hz- cu2i cgng (' fw" , H"w, )Fng .hGi gi0i =h6ng nng (Ong chNa .rng ch/. )6.x fc =h0n Nng chuy3n nng (Ong gi0i =h6ng ch h> .h2ng KTP. ST ^y h6a ccch/. )i )Di *Ii +T gi0i =h6ng nng (Ong *' .\ cc ch/. c6 (in ;%. ca nng.x VHn chuy1n )i>n .t*' hyUrg:n .5 nguyn (i>u hD h/= )%n ^yg:n c@a ;hV .rGi.Ng'i chNc nng ch@ y%u .rn, .y .h1 con c6 ;h0 nng .hng hO= =r.:in,=h+=hr(i=iU:, aciU &s, -. +2 h> :n|y-: nh cy.chr-:. Gen )ay, ngGi .a =h. hi>n.h/y -. (Ong KN *' -. (Ong (In K9N , .y .h1, ;hi%n -. +2 .c gi0 ch rvng .y .h1c6 ;h0 nng .hng hO= =r.:in )_c .hg *' U )6 cPng .ha- gia .Vch cTc *' *i>c ùy )nh.Vnh Ui .ruy3n c@a .% &' +2ng.c L{c (\=L{c (\= (' &' ùan )_c .rng c@a c .h1 .T Ujng. L{c (\= (' & -y ùang hO=c@a cay ânh.Th'nh=henh6ahJcc@a({c(\=gF-ccch/. ('-nhi>-*{c/u.r?cb =r.:in,(i=iU:, g(uciU:... *' cc ch/. ('- nhi>- *{ chNc nng +inh (kb cc +]c .2, cc h> :n|y-:,cc y%u .2 ;Vch .hVch...Th'nh =hen ùan .rJng nh/. .hTc hi>n chNc nng c@a ({c (\= (' cc +]c .2 *' cc h>:n|y-:. Trng ({c (\= c6 Y nh6- +]c .2 ;hc nhau, -~i nh6- c6 nhi3u (\i +]c .2bcNh6- fh(r=hy((b fh(a. fh(&, fh(c...cNh6- far.:niUbfar.:n:, Ran.h=hy((.cNh6- Phyc&i(inb=hyccyanin, =hyc:ry.hrin.Trng ({c (\= c6 h> :n|y-: .ha- gia *Hn chuy1n )i>n .t .rng ùang hO=,cc:n|y-: .ha- gia .rng =h+=hry( h6a ùang h6a, cc :n|y-: .ha- gia .rng .ra )hich/., )_c &i>. (' .rng ù .r4nh .hng hO= g(uciU: *' cc ch/. ;hc.L{c (\= c6 h4nh )!a, &a ùanh ({c (\= (' (I= -'ng ;s=. Bn .rng -'ng (' ;h2i cch/. c@a ({c (\= chNa nhi3u h> :n|y-: .ra )hi ch/..Trng ;h2i c ch/. c6 nhi3u &0n -ng, cc &0n -ng nv- r0i rc .rng c ch/. gJi(' Thy(aciU c ch/.p cc &0n -ng ^%= chFng (n nhau .\ nn grana )6 (' .hy(aciU h\.,(a-:n c6 c/u .\ .5 )n * -'ng c +, ^%= ^:n ;} *Ii cc +]c .2 *' cc h> :n|y-: .\nn -'ng ùang hO=.Trn .hy(aciU c6 nh[ng h\. nh W$7c $X n-Z, )6 (' ùangc.^-. luangc .^- (' )n * c/u .r?c c +, c@a ùang hO=. M~i ùangc .^- chNa $7# =han .t ch(a, 8# =han .t ch(&, uX =han .t ch(c, uX =han .t ùinn, $$7 =han .t =h+=hr(i=iU:, u7 =han .t +u((i=iU:, $" =han .t y:, " nguyn .t Mn, 7 nguyn .t fu.fN $# ùangc .^- .ha- gia h?. $# =h.n nh +ng )1 .i%n h'nh ;ht $ =han.tfw" . TH= hO= $# ùangc .^- (' -. )n * chNc nng ùang hO=.fhNc nng ch@ y%u c@a ({c (\= (' .hTc hi>n ù .r4nh ùang hO=. E6 (' ù .r4nh +tU{ng nng (Ong nh +ng )1 .hng hO= nn cc ch/. h[u c .5 fw" *' H"w.L{c (\= con .ha- gia *' cc ù .r4nh .hng hO= =r.iU:, (i=iU:, =h+=hr(i=iU:,aciU &s *' nhi3u hO= ch/. ;hc.c B -y G(gif/u .r?c & -y G(gi (' -. h> .h2ng nh[ng ;nh, )6 (' cc .?i U. u2n cng*ong cung U cc -'ng (i==r.:in .\ .h'nh. gi[a *' &n +Gn .?i U. )6 c6 cc ;hDng&' nh W"#c 7# n-Z *' ;hDng &' (In W#,mc "Z.B -y G(gi ('- nhi>- *{ .hu nhHn ch/. .h0i c@a .% &' )1 &'i .i%.p n6 c6 ;h0nng .hu nhHn ch/. (\, ch/. )c .ha- nhH= *' .% &' rFi .i%. ra ng'i nhv- &0 *> ch .%&'.c Li|^-. fon gJi (' .h1 hoa .an, )6 (' nh[ng .?i .ron nh, c6 -'ng nguyn +inh &a &Jc, )ay(' .?i chNa .rn $# h> :n|y-: .h@y =han ;hc nhau nh nuc(:a+:, =h+=ha(a+:. Th1 hoa.an c6 chNc nng =han gi0i cc ch/. h[u c, .r5 (i=iU:.c P:rî^- Eay('&'ùanh4nhceu, )Oc=h. hi>nn-$7m. P:rî^-chNanhi3u:n|y-: nh ca.a(a+:, =:rr^yUa+:, (a*in, cc :n|y-: .rng chu .r4nh g(iî(ic.P:rî^- (' .rung .a- .ra )hi cc ch/. =:rîU:, )_c &i>. (' H"w" c@a .% &'.N6con (' &' ùan chuyn h6a =h{ .rch ;hau cu2i cgng chuy1n h6a aciU &s.c M\ng (Ii ni ch/.c 9i&^- NhG ;Vnh hi1n *i )i>n .t , -\ng (Ii ni ch/. )< )Oc =h. hi>n. M\ng (Ii nich/. (' h> .h2ng 2ng Udn r/. -0nh nv- r0i rc .rng .% &' *' ch?ng n2i (i3n *Ii -'ngnhan .\ nn h> .h2ng .h2ng nh/. .rng .% &' *' n2i (i3n *Ii -\ng (Ii .% &' &n c\nh.Th'nh =hen h6a hJc ch@ y%u c@a -\ng (Ii ni ch/. (' =r.:in *' =h+=hr(i=iU:,ng'i ra con c6 K9N *' cc :n|y-:. f/u .r?c +iuhi1n *i c@a -\ng (Ii ni ch/. .ng .T nh -'ng c +,. f6 " (\i-\ng (Ii ni ch/.b -\ng (Ii ni ch/. .rn chq c6 -'ng ;s= (i==r.:in .\ nn *' -\ng(Ii ni ch/. c6 h\., .rn cc -'ng ;s= (i==r.:in c6 cc h\. ri&^- )Vnh *'. N6 (' h>.h2ng h[u c .rng .% &', &0 )0- +T *Hn chuy1n nhanh ch6ng cc ch/. .5 -Di .rGngng'i *' .% &' ch/. *' +T .ra )hi gi[a cc =hen ;hc nhau .rng ni & .% &'. N6 con.hng hO= nhi3u h> :n|y-:, .hng hO=, =han gi0i -j *' g(ucg:n.9i&^- (' &' ùan +iu hi1n *i, .rJng (Ong ;hD *Ii .h'nh =hen ch@ y%u gF- umcmmn =r.:in, K9N umc mmn. 9i&^- c6 -_. nhi3u ni .rng .% &' nh , .rn -'ngnhan, nhan cn, .y .h1, (\= .h1, -\ng (Ii ni ch/. hay nv- r0i rc .rng .% &' ch/..9i&^- (' .rung .a- .hng hO= =r.iU: c@a .% &'. E6 (' ni )1 K9N-)%n )Vnh *',)Fng .hGi )1 ch =hNc h> K9N. aa )%n g]n aa *' chuhi =:=.iU: )Oc .hng hO= .\i )6.4. Nhn.Nhan (' c ùan ùan .rJng nh/. .rng ch/. nguyn +inh.Th'nh=henh6ahJcc@anhanchNanhi3uch/. ;hcnhau, ùan.rJngnh/. ('=r.:inWm#c X#nZ , KNWmc $#nZ, K9NW#,mc Y,YnZ, (i=iU:WXc $"nZ... Trngcc=r.:in, hi+.n ùan .rJng nh/., n6 (in ;%. *Ii KN .\ nn cc fhr-a.iU .rng c/u.r?c c@a nhi- +]c .h1. Trng nhan c6 nhi3u (\i :n|y-: .ha- gia .rng cc ù .r4nh.hng hO= KN, K9N, -. +2 ù .r4nh .ra )hi ch/. ;hc.Nhan c6 -'ng nhan &a &Jc ;h2i ch/. nhan &n .rng, .rng ch/. nhan c6 cc nhancn *' cc nhi- +]c .h1.M'ng nhan (' -'ng " (I=, -~i (I= c6 c/u .\ gi2ng -'ng nguyn +inh ch/. c@a .%&'. M'ng ng'i c@a nhan .i%= ^?c *Ii -\ng (Ii ni ch/., .rn )6 c6 (~ .hDng c6 U "#cY# n-, )i3u n'y &0 )0- +T .ra )hi ch/. .hGng ûyn gi[a nhan *Ii .% &' ch/. fh/. nhanb Nhan chNa )ey )@ Uch nhan, ch@ y%u (' ch/. nhi- +]c .h1. Nhi- +]c.h1 (' c +, *H. ch/. -Nc ) .% &' c@a ù .r4nh Ui .ruy3n.Nhan cnb f6 *'i nhan cn .rng -~i nhanp nhan cn (' cc .h1 ceu ;hDng c6 -'ng&a &Jc. Nhan cn chNa ;h0ng X#c Xmn =r.:in, $#c $mn K9N, -. V. KN. Nhan cn(' .rung .a- .hng hO= =r.:in c@a nhan.Nhan ('.rung .a- )i3u ;hi1n *' )i3u hoa -Ji h\. )ng c@a .% &'. Nhan c6 *ai.roùy%. )nh.rngù.r4nh.hnghO==r.:in, cc:n|y-:*'cPng('ni .ra)hinuc(:ic aciU, .hng hO= KN .i +inh *' K9N +a -n chNc nng c@a -4nh, -~i &' ùan )3u c6 .h'nh =hen*' c/u .r?c r/. =hg hO= *Ii chNc nng )6. EFng .hGi gi[a cc &' ùan cPng c6 +T =h2ihO= nh= nh'ng .rng h\. )ng +2ng c@a .% &' cPng nh c@a c .h1. ST =h2i hO= n'ych .h/y -~i -. chNc nng U -. &' ùan )0- nhHn chVnh *' c6 +T )6ng g6= *Iinh[ng -Nc ) ;hc nhau c@a cc &' ùan *' c ch/. c@a .% &'. VV U{b ù .r4nh chuy1nh6a nng (Ong .rng .% &' .hTc *H. c6 +T .ha- gia c@a ({c(\=, .y .h1, .% &' ch/. *' -.+2 &' ùan ;hc, )_c &i>. (' h> -\ng (Ii ni ch/. )0- nhHn +T (in (\c gi[a cc =henc@a .% &', gi[a cc &' ùan *Ii nhau .\ .h'nh .h1 .h2ng nh/. .rng h\. )ng c@a .%&'. H\. )ng .h2ng nh/. n'y (\i )Oc +T )i3u ;hi1n c@a nhan. ThDng ùa c ch% .ruy3n)\. .hDng .in nhan )< .r, .h'nh .rung .a- )i3u ;hi1n -Ji h\. )ng c@a .% &'. Ei3u n'y&0 )0- ch .% &' .r, .h'nh -. )n * .h2ng nh/. *3 chNc nng.IV.Tnh cht cua nguyn sinh cht.T% &' ch/. c6 -. +2 .Vnh ch/. nh .Vnh ;:, .Vnh nhI., .Vnh *Hn )ng *' .Vnh )'nhFi.1. Tnh keo. TVnh ;: c@a .% &' ch/. (' ;h0 nng chuy1n Uch .5 .r\ng .hi S( W(ngZ +ang .r\ng.hi G:( Wnta (ngZ. TVnh ;: U cc =han .t =r.:in, nuc(:ic aciU *' cc ch/. h[u c anIc .rng .% &' ch/. gay nn.2. Tnh nht. E nhI. (' -a +. ni, (' (Tc c0n û/. hi>n ;hi cc (I= *H. ch/. .rO. (n nhau. EnhI. =h{ .huc *' h'- (Ong nIc. E nhI. (' chq .iu ùan .rJng ch =hs= )nh gi.r\ng .hi +inh (k c@a .% &'. fc .% &' c@a c ùan nn .hGng c6 ) nhI. .h/= hn )nhI. c@a cc .% &' , cc c ùan .r,ng .h'nh *' c ùan gi'. E nhI. c@a .% &' ch/.(in ùan *Ii -Nc ) .ra )hi ch/.. ihi ) nhI. .ng (n .ra )hi ch/. gi0- û2ng .ngNng *Ii .Vnh ch2ng chu ca c@a c ùan .hTc *H. )2i *Ii -Di .rGng &/. (Oi. T% &' ch/..rng cc .% &' , .r\ng .hi nghq nh h\. ;hD c6 ) nhI. ca. E2i *Ii cay chu n6ng .2.c6 ) nhI. ca *' n6 U & ch%. rs.p )2i *Ii c ùan +inh +0n .hGng c6 ) nhI. ca hnc ùan Uinh Ujng. ST ;hc &i>. )6 (' -. )_c )i1- c6 (Oi nhv- &0 *> noi gi2ng.3. Tnh dn hi.ih0nngùay(\i.r\ng.hi &an)eu+au;hi)ch?. nIc c@a .% &'. Th: VanhD=, = +u/. .hz- .h/u =h{ .huc *' nFng ) =han .t,nhi>. ), +T )i>n (y c@a Uung Uch *' .Vnh .h: cDng .hNcbP 9TfiPb = +u/. .hz- .h/u Wa.-Z , 9b hvng +2 ;hV #,#X"$Tb nhi>. ) .uy>. )2i W"8Y x .Z, fb nFng ) Uung Uch .h: M. ib h> +2 VanhD= &i1u .h -Nc ) in h6a Uung Uch.i $ x W n c $Z b h> +2 =han (y, nb +2 in -' =han .t =han (y1.1. T bo thc vt l mt h thm thu. .% &' .hTc *H., cc (I= -'ng c@a ch/. nguyn +inh (' nh[ng (I= -'ng gay nnhi>n .Ong .hz- .h/u .rng .% &'. T2c ) c@a nIc â- nhH= h_c .h. ra ;hi .% &'=h{ .huc *' .Vnh .hz- .h/u ;hc nhau c@a -'ng .% &' *' -'ng ch/. nguyn +inh. STâ- nhH= c@a nIc *' .% &' c6 .h1 ^zy ra .gy .huc *' nFng ) c@a Uung Uch *IinFng ) c@aUch .% &'. f6 Y .rGng hO=bc Erng .rngb f-. fTBc NhOc .rngb f-. fTB c u .rngb f-. fTBN%u nga- .% &' *' nIc h_c Uung Uch nhOc .rng Wf-. fTB Z .h4 nIc .5-Di .rGng )i *' ;hDng &' *' ('- .ng .h1 .Vch c@a ;hDng &'. = +u/. ('- ch ;hDng&' . ra s= *' .h'nh .% &' gJi (' = +u/..rng nIc WPZ. = +u/. n'y ('- -'ng .%&' cng ra. M'ng .% &' +inh ra -. +Nc ch2ng (\i gJi (' +Nc cng .rng nIc WTZ. ihihai = +u/. n'y &vng nhau .h4 +T .hz- .h/u U5ng (\i. T% &' , .r\ng .hi &< hoa *' .h1.Vch .% &' (?c n'y cTc )\i. fhVnh nhG +Nc cng WTZ n'y -' nh[ng =hen nn c@acay *dn)Nng *[ng. , ;hDng & & gH= (\i.N%u ):- .% &' )6 nga- *' Uch u .rng, nIc .5 .rng .% &' ra ng'i *' .h1.Vch .% &' nh )i, -'ng .% &' .r,(\i .r\ng .hi &4nh .hGng, +Nc cng WTZ &vng #. N?Uung Uch nga- .% &' ù u .rng, nIc .5 ;hDng &' .i%= .{c )i ra ng'i ('- ch;hDng &' c, ;s .h: nguyn +inh ch/. .ch rGi ;hi -'ng .% &'.Hi>n .Ong ch/. nguyn +inh .ch ;hi -'ng .% &' gJi (' hi>n .Ong c nguyn+inh.N%u ):- .% &' )ang c nguyn +inh n'y )_. *' Uung Uch nhOc .rng .h4 .% &'Uen Uen .r, *3 .r\ng .hi &4nh .hGng *' ^0y ra hi>n.Ong =h0n c nguyn +inh.Hi>n.Ongcnguyn+inh*'=h0ncnguyn+inh.h1hi>n.Vnh)'nhFi c@anguyn +inh ch/. n6i (n +T +2ng c@a .% &'. ihi .% &' ch%., -'ng &n .h/- & =h [email protected] +, c@a hi>n .Ong c *' =h0n c nguyn +inh (' .Vnh ch/. .hz- .h/u c@a .% &'.1.2. S ht nuc cua t bo theo co ch thm thu. ihi nga- .% &' *' Uung Uch nhOc .rng, nIc )i *' .rng .% &' *'.% &'&< hoa hi nIc. Tuy nhin, .rng -. cay nguyn *n, (?c n' cPng c6 +T .h. hinIc .5 (. )6 V. ;hi c6 +T &< hoa nIc .rng .% &'. fay .hGng ,.r\ng.hi .hi%unIc. .rGng hO= .% &'&< hoa nIc.h4 = +u/. .rng nIc P &vng *Ii +Nc cng.rng nIc T WPTZfon , .r\ng .hi .hi%u nIc c@a .% &' .h4 PT. *' PcTS. Nh *Hy +T +ai (>ch gi[aP *' T gay ra +Nc h?. nIc S. NhG +Nc h?. nIc S -' nIc c6 .h1)i (in.{c *' .% &'.S =h{ .huc *' .r\ng .hi &< hoa nIc c@a .% &'. ihi .% &' hs .h4 S (In, ;hi .% &'&< hoa .h4 S #, *4 (?c /yP T Pc T #.VHy .r +2 KSTT WPZ c6 k ngh!a (In .rng *i>c ^c )nh +Nc h?. nIc .h: c ch%.hz- .h/u. lu .r4nh n'y ;hDng .iu .2n nng (Ong c@a .% &', ^0y ra -. cch nhnh'ng *' =h{ .huc *' KSTT c@a -Di .rGng *' .% &'. 1.3. S ht nuc cua t bo theo co ch khng phai thm thu.Ta )< &i%. S û/. hi>n U c6 Pa.-.rng ;hDng &'. Tuy nhin +Nc h?. nIc c@a .%&' ;hDng =h0i )n .huen (' -. ù .r4nh *H. (k W.hz- .h/uZ. N6 con (in ùan )%n .r\ng.hi c@a ch/. nguyn +inh, =h{ .huc *' ù .r4nh .ra )hi ch/. *' nng (Ong. fhrngh\n , .% &' cha h4nh .h'nh ;hDng &' -. cch r r>. *dn c6 S. S .rng .rGng hO= n'y(' U = (Tc =hFng c@a ;: gay nn ;hi cc -i^:n ;: h/= .h{ nIc. SNc h?. nIc ;hDng=h0i chq +inh ra U ù .r4nh .hz- .h/u .huen .?y -' con U .Vnh ch/. (k h6a c@a h> ;:nguyn +inh ch/.. Nh *Hy ;hDng .h1^:- .% &' nh .hz- .h/u ;% )n gi0n. ST h?. nIc c@a .% &'U nhi3u c ch% -' -Nc ) )6ng g6= c@a .5ng c ch% (> .huc *' .5ng )i3u ;i>n &n.rng *' &n ng'i.L?c .% &' ;han nIc, h> ;: nguyn +inh c6 *ai .ro h?. nIcp (?c .% &' gi', h\.)ng +2ng & y%u, +Nc h/= .h{ ch@ )ng c6 k ngh!a ;hDng )ng ;1.2. S ht cht tan.T% &' ch/. ;hDng chq ch Uung -Di )i ùa, n6 cPng con ch -. +2 ch/. .rngUung -Di )i ùa. T% &' ch/. ;hDng =h0i (' -. -'ng &n .h/- h'n .'n -' n6 (' -.-'ng &n .h/- chJn (Jc . N6 h?. cc ch/. Uinh Ujng .5-Di .rGng &n ng'i. T% &'+2ng c6 ;h0 nng .Vch (Py, chJn (Jc cc ch/. Uinh Ujng. M. +2 ch/. .h/- +n +'ng ùa*ch .% &' nhng h'n .'n ;hDng chui ùa )Oc -'ng ng\i ch/. )1 *' &n .rng .%&'. M. +2 ch/. ;hc +au ;hi chui ùa )Oc -'ng ng\i ch/. (\i & gi[ (\i , .% &' ch/.*' ;hDng chui ùa )Oc -'ng ni ch/. )1 *' ;hDng &'. f6 nh[ng ch/. (\i c6 ;h0 nngchui ùa )Oc cc h> -'ngc@a .% &' *' .H= .rung )Oc .rng ;hDng &'. T% &' c6 ;h0nng h?. *' nhi3u ch/. ;hc nhau -_c Ug -Nc ) ;hDng gi2ng nhau.c E2i *Ii cc ch/. ;hDng )i>n (y fh?ng (> .huc *' .Q (> .Vnh .an .rng -j *' .rng nIc. Nh[ng ch/. c6 .Vnh .an.rng -j gi2ng nhau, .gy .h: ;Vch .hIc, +T â- nhH= c@a ch?ng *' .% &' cPng h'n.'n ;hc nhau.c E2i *Ii ch/. )i>n (y fhVnh )i>n .Vch c@a ch?ng )< c6 c0n .r, .Ii *i>c ch?ng â- nhH= *' .% &'. fh/.c6 )i>n (y c'ng .h/= .h4 ch?ng chui *' c'ng nhanh. fc in h6a .r $ WNax, ix Z chui *'.% &' nhanh hn cc in c6 h6a .r " Wfaxx, Mgxx Z, f(c, Sc *' .% &' U hn Swucc . N%ucgng ) )i>n (y, ch/. n' c6 in -'ng hyUra.: (In ;h6 .hz- .h/u hn ch/. c6 ;Vch .hIcin (In. Nh[ng in cen ch )Gi +2ng c@a cay nh P, i c6 .h1 )i *' .% &' r/.nhanh *'.H= .rung , .rng )6 -_c Ug nFng ) )< ca hn r/. nhi3u (en + *Ii nFng ) c@a n6 ,-Di .rGng. VI. Khi nim v nui cy m - t boT5 -. =hen nh c@a c ùan +inh Ujng nh )qnh +inh .r,ng, .han, r... &vngcch nuDi c/y -D .% &' +} .\ )Oc -. cay h'n chqnh. *Ii ; .huH. nuDi c/y -D c .%&' nh *Hy c6 .h1 .\ ra h'ng (\. cay cn .5 =hen nh c@a cayp nhan nhanh cc gi2ngcay nh/. (' cc gi2ng ùk hi%-.TI LIJU THAM KHAOI. TI LIJU TIENG VIJT$. Tren Eng i%, Nguyn Nh ihanh. "###. Sinh (k .hTc *H.. TH= -.. NRBG.". Ph\- E4nh Thi, Nguyn uy Minh, Nguyn Lng Hgng. $X8. Sinh (k hJc .hTc*H.. NRBG.Y. VP VnV{, H'ng Minh T/n, VP Thanh Ta- $. Sinh (k hJc .hTc *H.. NRBG. II. TI LIJU TIENG ANH$.Mar+chn:r, H. $X7. Min:ra( nu.ri.ininhigh:r =(an.+. KcaU:rnic=r:++. LnUnr(anU +an i:g N:L r;. Ku+.in B+.n SyUn:y T;y Trn..Chuong 2 S TRAO I NC THC VT2.1. Cc dng nuc trong dt, trong cy v vai tr cua n.2.1.1. Cc dng nuc trong dt, trong cy v vai tr cua n.Trng ù .r4nh .i%n h6a, .hTc *H. .5 )\i Ung .i%n Uen (n c\n *' â- n#hH= +au*' cc ({c )a. fh?ng g_= -au .hudn (In (' )i3u ;i>n cung c/= nIc .r, nn ;h6 ;hn*' c .h1 .hGng ûyn & .h0i -/. nIc r/. nhi3u *' ;hV ùy1n. Vi>c .ha -c nghin cNu ù .r4nh .ra )hi nIc c@a cay. Cng )< nu kngh!a +inh hJc c@a ù .r4nh .h. hi nIc *' )3 ra ùan ni>- -Ii *3 &0n ch/. .Vnh chuh\n c@a cay. V..ran W$X8Z )< =h. hi>n +T *Hn chuy1n nIc .rng h> -\ch .uan .h: ccùy (uH. .h@y )ng hJc.Nh[ngcDng.r4nhc@a*i>n+! Macî-*W$$7c$m"Z)n +T .iu ha nIc U U_. *' ;hDng nh[ng chq (in ùan *Ii cc )_c)i1- .hVch nghi *3 gi0i =hdu c@a cay -' ch@ y%u *Ii cc .Vnh ch/. h6a ;: *' +inh h6ac@a ch/. nguyn +inh *Ii .'n & ù .r4nh .ra )2i ch/. Uin ra , .rng cay.Nh[ng cDng .r4nh nghin cNu gen )ay chNng . rvng ù .r4nh .ra )hi nIc c@acay ;hDng )n .huen .uan .h: cc ùy (uH. *H. (k gi0n )n nh .rIc )ay ngGi .a .,ng.2.1.1.1. Cc dng nuc trong dtTr\ng .hi nIc .rng )/..Trng )/. ;hDng c6 nIc nguyn ch/. -' (' Uung Uch )H- )_c V. nhi3u .rng )6cc ch/. hoa .an c6 nFng ) nh/. )nh gay ra =h0n (Tc .hz- .h/u W+Nc (in ;%. .hz- .h/uZch2ng (\i +T *Hn chuy1n nIc *' cay. Trng )/. c6 ^c )ng *H., .hTc *H., c6 cc ch/.*D c nh hyUr^yU +]., hyUr^yU nhD-, )3u (' nh[ng U\ng ;: a nIc, nn c6 .h1.ranh ch/= -. =hen nIc c@a .hTc *H.. B3 -_. h\. ;: )/. c6 ;h0 nng h/= =h{ -.=hen nIc gay nn cc .r, (Tc ch *i>c h?. nIc c@a r *' cay. fay chq h?. nIc )Oc&vng c ch% .hz- .h/u .rng .rGng hO= nFng ) c@a Uch )/. &s hn nFng ) c@a ccch/. c6 h\. .Vnh .hz- .h/u , .rng &0n .han r. SNc (in ;%. .hz- .h/u c'ng .ng (?c )/.c'ng ;hD h_c (?c &6n .h- =han *' )/..Ng'i ra, nIc & (in ;%. ch_. .rn )/. &vng nh[ng (in ;%. h6a hJc &3n *[ng *Iinh[ng .h'nh =hen *D c, h[u c c@a )/. *' &a nIc -ng & h/= =h{ , .rn &3 -_. h\.;:. \ng nIc n'y c6 .h1 & gi[ )%n $###a.-. N6 c6 nhi3u .Vnh ch/. c@a .h1 r]n *' cayh'n .'n ;hDng +t U{ng )Oc Wc6 ngGi gJi (' nIc ngH-Z. TQ (> U\ng nIc (in ;%.=h{ .huc *' .h'nh =hen c giIi )/.. Th'nh =hen c giIi c'ng n_ng .h4 .Q (> nIc (in;%. ch_. c'ng ca Wc. .hD #,mn, )/. +s. n_ng $Y,"nZ. Ng'i U\ng (in ;%. ch_. *' .ng)2i y%u .rng )/. con c6 U\ng nIc .T U, (Tc h/= Udn c@a )/. heu nh ;hDng )ng ;1.NIc /y chNa )ey cc ;h: h, c@a cc h\. )/. *' , .r\ng .hi ;h (inh )ng, ch?ng )OcgJi (' nIc h/= Udn hay nIc .rJng (Tc. NIc n'y UIi .c U{ng c@a .rJng (Tc nn ch0y.5 ch~ ca )%n ch~ .h/=. ihi ch0y ùa r cay .h4 )Oc cay +t U{ng, nhng n%u n6 ch0yù nhanh .h4 cay chq +t U{ng )Oc V., n%u ch0y ùa chH- *' )Jng (\i , ch~ .h/= .h4 .\ra )i3u ;i>n y%- ;hV c6 h\i ch cay. Trng cc -a ù0n )/. h= nIc )Oc gi[ ch_. hn&,i +Nc cng &3 -_. c@a -_. (- *' ;hDng & ch0y û2ng .h: .rJng (Tc, =hen nIc n'y)Oc gJi (' nIc -a Udn. NIc Uang (n c'ng ca n%u -a ù0n c'ng &s. Eay (' U\ngnIc c6 k ngh!a ch@ y%u .rng canh .c.NgGi .a c6 .h1 &i1u .h (Ong nIc ;hDng h?. )Oc &vng h> +2 hs. E6 (' U\ngnIc UT .r[ ch%. W&i1u .h &vng n c@a )/. ;hDZ con (\i .rng )/.. ihi ( -Jc .rn )/. )6&]. )eu c6 .ri>u chNng hs. Th: cDng .hNc .hTc nghi>- c@a Brigg+ *' fhan.| W$$YZb Nh[ng nghin cNu gen )ay ch .h/y h> +2 hs ;hDng nh[ng (> .huc *Ii )_c .Vnhc@a )/. -' con =h{ .huc .Vnh ch/. +inh (V c@a cay.Trn cgng -. .hN )/. cc cay c6 ;h0 nng h?. nIc ;hDng gi2ng nhau. Ng'i ra h>+2 hs con &i%n .hin .rng ù .r4nh =h. .ri1n c .h1 c@a cay. Nghin cNu c@a Macî-*cPng ch .h/y cay &]. )eu .hi%u nIc ;hDng =h0i )\. .Ii .r +2 h> +2 hs -' con +I- hnnhi3u. E2i *Ii )a +2 cay ) z- .2i .hVch c@a )/. (a 7#cX#n c@a z- Uung cTc )\i.2.1.1.2. Cc dng nuc trong cy Nng (Ong .T U c@a nIcM~i =han .t *H. ch/. )3u c6 nng (Ong &n .rng chung W.hng ni nngZ gF- )ngnng *' .h% nng. Nng (Ong .T U (' nng (Ong .rng )i3u ;i>n .hVch hO= c6 ;h0 nng+inh ra cDng. NIc (' -. U\ng *H. ch/. cPng c6 nng (Ong .T U.NgGi .a )< )a ra -. nguyn (V c &0n (' nIc +} chuy1n Uch .5 ni c6 nng(Ong .T U ca )%n ni c6 nng (Ong .T U .h/=. 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Ung +} gi[ ) .rng ch .% &'. E .rng n'y +} )zy ch/. nguyn +inh ch2ng (\i -'ng *' .h'nh .% &' ('- ch .% &' c6 h4nh U\ng nh/. )nh. ihi .h% nng = +u/. gi0- .Ii ;hDng U .% &' -/. nIc, .% &' )6 +} c nguyn +inh, ch/. nguyn +inh .ch ;hi .h'nh .% &'. H4nhSS.". -iu .0 -2i ùan h> gi[a gi .r .h% nng nIc WZ, .h% nng .hz- .h/u WZ*' .h% nng = +u/. WZ ;hi nIc â- nhH= *' .% &'.ihi nIc *' .% &', cc gi .r , , )3u .ng. .ng ;hi nIc *'.% &' *' +T .ng *O. c@a nIc .rng .% &' ('- ch .ng. .ng *4 ;hi nIc *' +} ('- gi0- nFng ) ch/. .an. T\i )i1- .h1 .Vch .ng )2i c@a .% &' (' $,# .h4 .% &' , .r\ng .hi hs, -3-. ihi .h% nng c@a Uch .% &' .i%n .Ii ;hDng .h4 +T â- nhH= nIc *' .% &' +} ng5ng.V3 .h% nng c ch/. -. Trng .% &' c6 nhi3u c ch/. c6 ;h0 nng h/= .h{ nIc.\ ra .h% nng c ch/.. ST h/= .h{ nIc .rn &3 -_. c ch/. ('- gi0- nng (Ong .T Uc@a nIc *' *4 *Hy, .h% nng c ch/. c6 gi .r a-, n6 ('- gi0- .h% nng nIc c@a .% &'.Gi .r c@a .h% nng c ch/. r/. nh, .rng )\i & =hHn cc .rGng hO=-#,$ &a. 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Sau )ay (' h'-(Ong nIc c@a -. +2 c ùan *' cay ;hc nhaub Tn thc vt v co quanHm lung nuc ()T0 #cXL cay R' (ch, H'nh, ù0 f' chua,a chu.ucmL f0i &]=, c@ Su h' "cYf@ f' r2. X8c$lu0 T, L XYcX7f@ ihai .ay 8ucX#L cay g~, cay &{i 8cX"Than cay g~ Wg~ .i *5a ^Z u#cm#H\. ;hD ;hDng ;hVWL?a -4, L?a nIc, NgDZ $"c$u Trng cay, nIc , .rng cc .% &' +2ng, cc y%u .2 ^y(:- ch%. Wcc -\ch *' ù0n&'Z *' .rng cc ;h0ng gian &'.Trng .h'nh .% &', ch/. nguyn +inh *' Uch &' nIc, .r\ng .hi (ng. Trng cc ;h0ng gian &' nIc , .r\ng .hi hi.H'- (Ong nIc con ;hc nhau , cc .eng (b .eng ( c'ng , =hVa UIi c'ng c6 h'-(Ong nIc ca. VV U{b .rng ;hDng &', cc .% &' ( BDng h'- (Ong nIc , cc .eng( nh +aub , .eng UIib YYcY8np , .eng gi[ab "7cY#np , .eng .rnb "mc"8n.H'- (Ong nIc con =h{ .huc *' )i3u ;i>n -Di .rGng *' +T =h. .ri1n c .h1c@a .hTc *H.. fc cay .h@y +inh c6 h'- (Ong nIc (In hn cay .rung +inh, h\n +inh. fccay nn chNa nhi3u nIc hn cay gi'.H'- (Ong nIc c@a cay con .hay )hi .h: nh= )i>u ng'y. H'- (Ong nIc (?c&uhi .ra n]ng .h/= hn &uhi +ng. Tuy h'- (Ong nIc .hay )hi .gy .huc *' nhi3u y%u.2 nh *Hy nhng , .rng cay nIc *dn gi[ )Oc .r\ng .hi can &vng )ng gi[a +T h?.nIc ùa r *' +T .h. hi nIc ùa ( .rng )i3u ;i>n &4nh .hGng.ihi h'- (Ong nIc .rng .% &' )\. .Ii -Nc 8#c#n .h4 cc ù .r4nh +2ng .rngch/. nguyn +inh ^0y ra -\nh nh/.. Thi%u nIc cc ù .r4nh +inh (V & *i =h\-, cc ù.r4nh =han gi0i .ng (n. fay =h0i .ng cGng h?. nIc )1 &g (\i (Ong nIc )< -/. ùaù .r4nh .h. hi nIc. NgGi .a )< .Vnh .n )Oc +2 (i>u nh +aub cN $### g nIc)Oc h?. *' cay .h4 $,mc",# g )Oc +t U{ng )1 .hng hO= cc ch/. h[u c, con (\i (' )1 &g*'(OngnIc) -\ch Udn =h. .ri1n .2. )1 Udn nIc )< h?. (n cc c ùan .h. hinIc.c Ph0i c6 h> -D &4 =h. .ri1n )1 h\n ch% +T .h. hi nIc c@a cay.Nhu ceu nIc c@a cay r/. (In, *V U{ -. cay NgD cen )%n "## ;g nIc h_c hn.rng )Gi +2ng c@a n6. Nhu ceu nIc =h{ .huc *' cc )_c )i1- +inh .hi Wcay , *gngn6ng c6 nhu ceu nIc ca hn , *gng (\nhZ. Nhu ceu nIc con =h{ .huc *' cc (Na.uhi ;hc nhau .rng cgng -. cay, cc ('i cay ;hc nhau, cc nh6- cay ;hc nhau. f6 .h1 cn cN *' h> +2 .h. hi nIc W+2 g nIc .h. ra )1 h4nh .h'nh nn $ gch/. ;hDZ )1 ^c )nh nhu ceu nIc c@a cc (\i cay. Tr\ng .hi nIc *' cc U\ng nIc .rng cay.- Trng thi ca nc trong cy.Phan .t nIc c6 .Vnh =han cTc W--:n (jng cTcZ. H> .h2ng gF- hai )i>n .Vch&vng nhau *3 .r +2 nhng ;hc nhau *3 U/u Wx*' cZ, nv- cch nhau .rng ;h0ng rn' )6, )Oc gJi (' (jng cTc )i>n. Phan .t nIc )Oc )nh hIng -. cch ^c )nh.rng )i>n .rGng c@a in .Nc (' ('- .h@y h6a in )6.- Trng thi thy ha ha hc.fc =han .t h[u c cPng c6 --:n (jng cTc, n6 (' .hng cc --:n (jng cTcc@a cc nh6- =han cTc c6 .rng .h'nh =hen c@a =han .t. V4 *Hy, ûng ùanh cc =han .t)6 .\ ra -. )i>n .rGng .hu h?. +T )nh hIng ^c )nh c@a cc =han .t nIc, .Nc ('gay nn +T .h@y h6a. Eay (' +T .h@y h6a .rung hoa )i>n. fhq -. +2 nh6- nh/. )nh nhcar&^y( WcfwwHZ, hyUr^y( WcwHZ,a(U:hyU WcfHwZ,car&ny(W fwZ,i-in W NHZ,a-in WcNH"Z, a-iU W fwNH"Z -Ii gay ra +T )nh hIng W+T .h@y h6aZ c@a cc =han .tnIc (jng cTc ;hi , gen cc nh6- )6.Ng'i ra, cc =han .t nIc (jng cTc con )nh hIng gen cc nh6- in h6a, *V U{bcc =hen in h6a c@a aciU a-in .rng =han .t =r.:in WNHYxp fwwcZ. E6 (' +T .h@y h6ain h6a. gen -. nh6- =han cTc hay in h6a c6 .h1 c6 -. *'i (I= =han .t nIc (jngcTc )nh hIng .\ nn (I= * .h@y h6a, .rng )6 cc (I= .rng cgng )Oc )nh hIng.rH. .T nh/. *' (in ;%. ch_., cc (I= .i%= .h: (ng ( hn *' cc (I= c'ng â +T .ng .cc'ng ;s- *' ;hDng con +T .h@y h6a n[a. ST .h@y h6a nu .rn (' -. ù .r4nh h6a hJcgay nn &,i cc (Tc h6a .r nn gJi (' +T .h@y h6a h6a hJc. E6 (' .r\ng .hi chVnh c@anIc .rng .% &'. Ng'i .r\ng .hi .h@y h6a, .rng .% &' nIc con , .r\ng .hi (in ;%.c/u .r?c Wcon gJi (' +T &/. )ng h6aZ *' .r\ng .hi h?. .hz- .h/u.ST &/. )ng h6a nIc c6 .h1 ^0y ra , &n .rng )\i =han .t *' .rng cc ;h0ngh= nv- gi[a cc )\i =han .t ch nn h\n ch% +T chuy3n c@a cc =han .t nIc -. cchc hJc.Tr\ng .hi h?. .hz- .h/u cPng c6 .h1 ^0y ra &n .rng cc )\i =han .t cPng nh.rng cc ;h0ng h= gi[a ch?ng. )ay, nIc & h?. &,i cc =hen =han .t .h/= U cchO= ch/. ca =han .t =han gi0i ra.Trng cc ùan ni>- +au n'y, cc =han .t nIc .rng c .h1 +2ng .Fn .\i , hai .r\ng.hibM. =hen nIc ('- ;hung W-\ngZ .\ nn c/u .r?c nIc W.Vnh +]= ^%= .hN .T .h:-\ng c@a nIcZ )Oc h4nh .h'nh nhG cc (in ;%. hyUrg:n gi[a cc =han .t.Phen .hN hai (/= )ey cc (~ .r2ng c@a ;hung )6.- Cc dng nc trong cy. Trng cay nIc .Fn .\i , hai U\ng (' nIc .T U *' nIc (in ;%.. Tuy nhin, nIcn' )Oc ci (' nIc .T U *' nIc n' )Oc ci (' nIc (in ;%. .h4 c6 nhi3u ùan )i1-;hc nhau. M. +2 ch rvng nIc (in ;%. (' nIc ;hDng & )Dng (\i , nhi>. ) .h/= hnc$#

f *' ;hDng c6 .h1 Ugng ('- Uung -Di ngay ch nh[ng ch/. U & hoa .an nh )GngWMacî-*Z.luan ni>- .hN hai ch rvng, =hen (In nIc & (in ;%. &vng cch .ha- gia *' +T.h@y h6a h6a hJc cPng nh *' +T (in ;%. c/u .r?c. Phen nIc con (\i )Oc ci (' nIc.T U. E6 (' nIc h?. .rng cc -a ù0n W.rng .h'nh .% &'Z, nIc & h?. .hz- .h/u,;hDng .ha- gia *' .h'nh =hen c@a * .h@y h6a ûng ùanh cc =han .t *' in, .r5 nIc.huc (I= ;hu%ch .n c@a * .h@y h6a *' con gi[ )Oc .Vnh (inh )ng WK(:c^:i:*Z.M. +2 nh' nghin cNu ;hc (\i .h/y rvng, .rng -Ji .rGng hO= nIc .rng .hTc*H. )3u (' nIc (in ;%. *' .gy .h: .c U{ng c@a cc (Tc gi[a cc =han .t *' ni =han .t-' h\. .Vnh c@a nIc & &i%n )hi )i. V4 *Hy, cc .c gi0 n'y )< nghi ngG +T .Fn .\i c@aU\ng nIc .T U. HJ )< )a ra +T =han chia .h'nh hai U\ng nIc (in ;%.b nIc (in ;%.ch_. *' nIc (in ;%. y%u .rng )6 nIc (in ;%. y%u (' nIc *dn gi[ )Oc .Vnh ch/. c@anIc .hDng .hGng.SI U! c6 nh[ng ùan )i1- ;hc nhau *3 cc U\ng nIc .rng cay (' *4 ;hDng c6ranh giIi r r>. *3 hai U\ng nIc .T U *' nIc (in ;%. WU\ng nIc n'y c6 .h1 chuy1n.h'nh U\ng nIc ;iaZ. Sau n'y ngGi .a )< )a ra ùan ni>- nh +au )1 =han &i>. haiU\ng nIc .rnbNIc .T U hay nIc (in ;%. y%u (' nIc & r?. ra ;hi .hTc *H. nhG nh[ng (Tc h?.nIc ^c )nh *' nIc )6 c6 nh[ng .Vnh ch/. gen gi2ng *Ii .Vnh ch/. c@a nIc .hGngWngh!a (' c6 .h1 Ugng ('- Uung -Di *' )Dng )_c , nhi>. ) gen #

fZ.NIc (in ;%. hay nIc (in ;%. ch_. (' =hen nIc con (\i -' .Vnh ch/. c@a ch?ng ). ) .h/= hn#

fZ.Ti%= .h: ngGi .a (\i =han chia c{ .h1 hn .h'nh Y U\ng nIc nh +aubx NIc (in ;%. ch_. (' nIc & gi[ (\i U ù .r4nh .h@y h6a h6a hJc cc in *' cc=han .t, cc ch/. .rgng hO= .h/= *' .rgng hO= ca.x NIc (in ;%. y%u (' nIc .huc cc (I= ;hu%ch .n c@a * .h@y h6a, nIc (in;%. c/u .r?c *' nIc h?. .hz- .h/u.x NIc .T U (' nIc & h?. .rng cc -a ù0n c@a .h'nh .% &' *' =hen nIc &h?. .hz- .h/u c@a Uch .% &', ;hDng .ha- gia *' .h'nh =hen * .h@y h6a ûng ùanhcc in *' =han .t. nghia cua cc dng nucST ;hc nhau *3 .Vnh ch/. c@a nIc .T U *' nIc (in ;%. )< )a )%n +T ;hc nhau*3 k ngh!a c@a ch?ng )2i *Ii )Gi +2ng c@a .hTc *H..x NIc .T U chi%- -. (Ong (In .rng .hTc *H. W8#nZ (\i (' U\ng nIc con Ui)ng )Oc.*' con gi[ nguyn nh[ng )_c .Vnh c@a nIc ch nn )6ng *ai .ro ùan .rJng.rng ù .r4nh .ra )hi ch/. c@a .hTc *H.. )6, ngGi .a )< ^c )nh rvng, (Ong nIc.T U ùi )nh cGng ) cc ù .r4nh +inh (V.x NIc (in ;%. ch_. *' ;hDng ch_. (' U\ng nIc chi%- Y#n (Ong nIc .rng cay.Tgy .h: -Nc ) ;hc nhau U\ng n'y -/. .Vnh ch/. &an )eu c@a nIc nh ;h0 nng ('-Uung -Di ;s-, nhi>. Uung gi0- û2ng, ) )'n hFi .ng (n, nhi>. ) )Dng )_c .h/=.Vai .ro c@a U\ng nIc n'y (' )0- &0 ) &3n *[ng c@a h> .h2ng ;: .rng ch/.nguyn +inh *4 n6 ('- ch cc =han .t =han .n ;h6 (]ng û2ng, hi>n .Ong ngng ;%. V.^0y ra.Trng cc c .h1 nn, h'- (Ong nIc (in ;%. .h/y nh hn .rng cc c .h1 gi'.ihi .hTc *H. g_= )i3u ;i>n ;hD h\n, h'- (Ong nIc (in ;%. .ng (n. fh nn, c6 .h1 ('h'- (Ong nIc (in ;%. (in ùan *Ii .Vnh ch2ng chu c@a .hTc *H. nh chu h\n, churs., chu -_n. NgGi .a )< Ugng .Q +2 h'- (Ong nIc (in ;%. *' nIc .T U )1 )nh gi;h0 nng ch2ng chu c@a .hTc *H. *' Nng U{ng .rng *i>c chJn cc gi2ng cay c6 ;h0 nngch2ng chu .2. nh/..S phn b cua cc dng nuc trong cyTrng .h'nh .% &' ch@ y%u gF- nIc (in ;%. &,i cc ch/. ca =han .t c:((u(+:,h:-ic:((u(+:, cc ch/. =:c.in, nIc (in ;%. c/u .r?c *' nIc & h?. .rng cc -a ù0n.Trng ch/. nguyn +inh, *i>c ^c )nh cc U\ng nIc .rng ch/. nguyn +inh g_=nhi3u ;h6 ;hn U .Vnh ch/. =hNc .\= *' .Vnh ;hDng hn )nh *3 .h'nh =hen *' c/u .r?cc@a n6. Tuy nhin, ngGi .a cPng ^c )nh rvng, .rng ch/. nguyn +inh nIc & (in ;%.&,i +T .h@y h6a h6a hJc c@a =r.:in *' cc ch/. ca =han .t ;hc, nIc (in ;%. c/u .r?c,nIc (in ;%. &,i cc ch/. .rgng hO= .h/= *' nIc & h?. &vng .hz- .h/u W&n .rng cc)\i =han .tZ.:nh h;ng ca .t $( icc ch/. .Vch nIc V. hay nhi3u .rng .% &'.nh h,ng .rTc .i%= c@a cc in )%n +T .h@y h6a h6a hJc (' U ch?ng & h?. &-.rn &3 -_. cc .i1u =hen & & .h@y h6a Wcc )\i =han .tZ ch nn ch?ng c6 .h1 .c )ng)%n +T .h@y h6a h6a hJc cc in cPng nh +T .h@y h6a h6a hJc .rung hoa )i>n.NgGi .a )< chNng -inh rvng, cc ch/. )i>n (i .ng (n +} ('- ch (Ong nIc (in;%. .ng (n. Tuy nhin, )2i *Ii cc in ix, f+x, Sc .h4 ngOc (\i, .Nc (' (Ong nIc (in ;%.gi0- )i U c/u .r?c c@a nIc .rng cc * .h@y h6a ;s- &3n *[ng )i.ST h?. &- -. (\i in n'y c6 .h1 ;- .h: +T .h0i ra cc in ;hc W+T .ra )hi h?.&-Z. N%u -Nc ) .h@y h6a c@a in & h?. &- (In hn -Nc ) .h@y h6a c@a in & .h0ira .h4 (Ong nIc (in ;%. *Ii )\i =han .t +} .ng (n. Trng .rGng hO= ngOc (\i, (OngnIc (in ;%. +} gi0- )i.Bvng .hTc nghi>- ngGi .a )< chq ra rvng, *i>c .i- cc Uung Uch c@a ./. c0 cc-u2i gay ra +T gi'- (Ong nIc .T U *' ('- .ng (Ong nIc (in ;%. *4 cc in )< 0nhh,ng )%n +T .h@y h6a h6a hJc cc )\i =han .t.nh h,ng c@a Uinh Ujng ;hng con .h1 hi>n , ch~ cc in ;hng 0nh h,ng)%n .h'nh =hen cc ch/. h[u c U cay .hng hO=. E6 (' cc )\i =han .t c6 chNa cc nh6-a nIc Wnh6- =han cTc hay inZ -' +2 (Ong *' +T =han &2 c@a cc nh6- n'y chi =h2i-Nc ) .h@y h6a c@a cc )\i =han .t nn c6 .h1 ('- &i%n )hi cc (in ;%. c/u .r?c c@anIc. cc ;%. ù0 nghin cNu )< ch .h/y ;hi .h@y =han =r.:in c6 .h1 ^0y ra +T =han gi0i=:=.iU )1 .\ .h'nh cc nh6- =han cTc -Ii WNH"*' fwwHZ (in ;%. *Ii cc =han .tnIc -Ii.x nh h,ng c@a nhi>. ). ST .h@y h6a h6a hJc (' -. ù .r4nh ng\i nhi>.. V4*Hy, ;hi h> h?. nhi>. .h4 =h0i ^0y ra ù .r4nh ngOc (\i .Nc (' +T .h@y h6a U chuy1n )ngnhi>. c@a cc =han .t nIc .ng (n, gay .c )ng ngOc (\i +T )nh hIng c@a cc =han.t nIc. *Hy, ;hi nhi>. ) .ng .h4 h'- (Ong nIc (in ;%. gi0-, nhng h'- (Ong nIc(in ;%. c/u .r?c *' nIc h?. .hz- .h/u .h4 (\i .ng (n, U +T gi0- (Tc (in ;%. c@a ccnh6- =han cTc ('- ch c/u .r?c cc )\i =han .t .r, nn ^2=, cgng *Ii *i>c .ng )ngnng c@a cc in *' cc =han .t ('- ch (Ong nIc (in ;%. c/u .r?c *' +T h?. nIc.hz- .h/u .ng (n.Ng'i 0nh h,ng )%n +T .h@y h6a, +T .ng nhi>. ) con 0nh h,ng )%n ù .r4nh.ra )hi ch/. ùa *i>c .h?c )zy ù .r4nh .h@y =han ('- gi0- (Ong cc ch/. ca =han .t,Udn )%n *i>c gi0- (Ong nIc (in ;%. *' .ng (Ong nIc .T U. 2.1.2. Vai tr cua nuc di vi thc vt. NIc (' .h'nh =hen &]. &uc c@a .% &' +2ng. f6 nhi3u nIc .hTc *H. -Ii h\.)ng &4nh .hGng )Oc. Nhng h'- (Ong nIc .rng .hTc *H. ;hDng gi2ng nhau, .hay)hi .gy .huc ('i hay cc .h chNc ;hc nhau c@a cgng -. ('i .hTc *H.. H'- (OngnIc con =h{ .huc *' .hGi ; +inh .r,ng c@a cay *' )i3u ;i>n ng\i c0nh -' cay+2ng. V4 *Hybc NIc (' .h'nh =hen c/u .r?c .\ nn ch/. nguyn +inh W#nZ.c N%u nh h'- (Ong nIc gi0- .h4 ch/. nguyn +inh .5 .r\ng .hi +( chuy1n .h'nhg:( *' h\. )ng +2ng c@a n6 +} gi0- +?..c fc ù .r4nh .ra )2i ch/. )3u cen nIc .ha-gia.NIcnhi3uhayV.+} 0nhh,ng )%n chi3u hIng *' cGng ) c@a ù .r4nh .ra )2i ch/..c NIc (' nguyn (i>u .ha- gia *' -. +2 ù .r4nh .ra )2i ch/..c ST *Hn chuy1n cc ch/. *D c *' h[u c )3u , .rng -Di .rGng nIc.c NIc &0 )0- ch .hTc *H. c6 -. h4nh U\ng *' c/u .r?c nh/. )nh. nIcchi%- -. (Ong (In .rng .% &' .hTc *H., Uuy .r4 ) .rng c@a .% &' ch nn ('- ch.hTc *H. c6 -. h4nh Ung nh/. )nh.c NIc n2i (i3n cay *Ii )/. *' ;hV ùy1n g6= =hen .Vch cTc .rng *i>c &0 )0- -2i(in h> ;hng ;hV. +T .h2ng nh/. gi[a c .h1 *' -Di .rGng. Trng ù .r4nh .ra )hi gi[acay *' -Di .rGng )/. c6 +T .ha- gia .Vch cTc c@a in Hx *' wHc U nIc =han (y ra.c NIc g6= =hen *' +T Udn .ruy3n ûng )ng cc Uong )i>n +inh hJc , .rng cay;hi%n ch?ng =h0n Nng -au ( ;hDng ;s- -. +2 .hTc *H. &Hc .h/= UIi 0nh h,ng [email protected] nhan ;Vch .hVch c@a ng\i c0nh.c NIc c6 -. +2 .Vnh ch/. h6a (k )_c &i>. nh .Vnh Udn nhi>. ca, c6 (Oi ch .hTc*H. =h. .n *' Uuy .r4 nhi>. (Ong .rng cay. NIc c6 +Nc cng &3 -_. (In nn c6 (Oi ch*i>c h/= .h{ *' *Hn chuy1n *H. ch/.. NIc c6 .h1 ch .ia .t ng\i *' nh +ng .rDng .h/y)i ùa nn c6 (Oi ch ùang hO=. NIc (' ch/. (jng cTc r r'ng nn gay h4>n .Ong .h@yh6a *' ('- ch ;: a nIc )Oc hn )nh.M. +2 .hTc *H. h\ )rng Wru, )a yZ c6 h'- (Ong nIc V. Wmc8nZ, chu )Tng .hi%unIc (au U'i, )Fng .hGi c6 .h1 chu )Tng )Oc +T ;hD h\n h'n .'n. ThTc *H. .hOng)rng -Jc , n?i ) hay +a -\c cPng chu )Oc h\n con )\i )a +2 .hTc *H. n%u .hi%u nIc(au U'i .h4 ch%.. fung c/= nIc ch cay (' )i3u ;hDng .h1 .hi%u )Oc )1 &0 )0- .huh\ch .2..Vi>c .ha -=. 2.2. S ht nuc cua cy.2.2.1. S hp th nuc cua r.2.2.1.1. D}c dim sinh hc cua h r thch nghi vi chc nng ht nuc.lua nghin cNu c@a nhi3u nh' ;ha hJc ch .h/y r c6 ;h0 nng )a- +au, (anrng, ;h0 nng =han nhnh c6 &3 -_. *' ) U'i (In hn .han *' ( g/= &i. NhG c6 nh[ng)_c .Vnh .rn, r c6 ;h0 nng +t U{ng (Ong z- =han .n, V. i *' ch/. ;hng ;h nghn'n .rng -Di .rGng )/.. Vi>n +! Macî-* W$uuZ )< n6ib Tri *Ii ùan ni>- .hDng.hGng h> r ;hDng =h0i )Vnh ch_., &/. )ng .rng -. -i3n nh/. )nh n' )6 c@a )/. -'(uDn (uDn Ui chuy1n *3 )vng .rIc .Ta nh -. )'n ;hhng (F ('i *H. nh &s )' (i%-ùanh -~i h\. c. g_= =h0i *' .ch nh[ng -'ng nIc -ng UVnh .5 )6. )6 ;hDng chqnIc ch0y .h: -a ù0n .Ii )eu r, -' )eu r )Oc .rO. .h: nIc *' *4 -{c )Vch )6ch?ng @i )' )/. -. cch -0nh (i>. ;hDng & +6. -. (y ;h2i nIc n' ;hDng +t U{ng.L?c cc (I= )/. -_. c'ng ;hD, h> r c'ng ng'y c'ng )a- +au *' (ong )/..2.2.1.2. Hot dng ht nuc v bom nuc vo r.ih0 nng h/= .h{ .Vch cTc c@a nIc .5 )/. *' )zy n6 *' (ong -\ch (n .han c@ar cay &i1u hi>n r r'ng .rng hi>n .Ong rq nhTa *' N giJ.. NhTa rq ra c6 .h'nh =hen r/.=hNc .\=. Ng'i cc -u2i ;hng .rng nhTa con c6 cc aciU h[u c, aciU a-in, )Gng,=r.:in *' cc ch/. h[u c ;hc. Nguyn nhan gay ra hi>n .Ong rq nhTa (' U r +0n +inhra = (Tc. n%u .a ):- ch~ c]. g]n (i3n *Ii = (Tc ;% .h4 .a +} ) )Oc = (Tc r (In hay &s.fc ('i c .hGng ;hDng ù $ a.-, cay g~ ca hn V. nhi3u. Th: hi.: W$uZ, ngay rf' chua c6 .rGng hO= cay .\ nn -. (Tc )zy .Ii Yc$# a.-. Trng cgng -. cay c6 rqnhTa nhi3u hay V. =h{ .huc *' .r\ng .hi .uhi, .r\ng .hi +inh (V, +T +inh .r,ng -\nhhay y%u. E2i *Ii (\i cay -. n- .h4 +au ;hi ra ha hi>n .Ong rq nhTa gi0- û2ng rr>.. fhVnh = (Tc r )< gay ra ù .r4nh h?. nIc ch@ )ng ch cay. Gi0i .hVch c ch% =(Tc r, ch )%n nay cha h'n .'n nh/. .rV. Th: -. +2 .c gi0, r c6 .h1 h?. nIc ch@)ng (' nhG c ch% .hz- .h/u W)ng c UIiZ.

Hi>n .Ong N giJ. c6 .h1 .h/y )Oc (?c &an -ai. V' .hGi gian &an )- ;hV hHu(\nh, chung ùanh ;hDng ;hV )Oc &< hoa hi nIc, ;hi%n ù .r4nh h. nIc .5 ( &h\n ch%. Hi>n .Ong N giJ. c6 .c U{ng Uuy .r4 +T can &vng gi[a h/= .hu*' .h. nIc *'(' U/u *%. con (\i c@a h4nh .hNc .ra )2i n6c c@a .h .in .h@y +inh â â. S2 (Ong NgiJ. &i%n )hi r/. (In, c6 (?c chq c6 -/y giJ., c6 (?c .rn -. ( .rng -. &uhi .2i c6 )%n$#-( nIc. Th'nh =hen cc ch/. .rng nIc N giJ. cPng&a gF- c0 ch/. *D c *' h[uc.Hai hi>n .Ong rq nhTa *' N giJ. (' U ;h0 nng h?. nIc *' )zy nIc -. cchch@ )ng c@a r (n .han. fh?ng c6 (n ùan ;hng ;hV. *Ii h\. )ng +2ng c@a cay )_c&i>. (' ù .r4nh hD h/=.L?c ^t (k h6a ch/. gay - W:.h:r, ch(rr-...Z h_c cc )c .2 hD h/= WifN,fw ...Z .a .h/y hi>n .Ong rq nhTa cPng nh N giJ. & )4nh chq. fc Udn (i>u chNng . ù.r4nh h?. nIc ch@ )ng c@a r )oi hi .iu ha nng (Ong (' -. ;hau .rng =hNc h> ccù .r4nh .ra )2i ch/..2.2.2. Anh huong cua cc diu kin bn ngoi dn qu trnh ht nuc.ST h?. nIc c@a r =h{ .huc *' )i3u ;i>n ng\i c0nh nh nhi>. ), .h'nh =hen;hng, ) z- c@a -Di .rGng. . Nhi>. ) .h/= ) h?. nIc c@a r gi0- û2ng Wnh/. (' cay a +ngZ. fc cay c6nhi>. ) .hVch hO= )2i *Ii h\. )ng h?. nIc Wc' chua "m

f, chanh Y#

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Trng ;hi +T .h@y h6a .rung hoa )i>n, .c U{ng (i.r= c@a cc in (in ùan*Ii * .rV c@a ch?ng .rng Uc ('- c/u.r?c nIc ;s- &3n *[ng.ST h?. &- -. (\i in n'y c6 .h1 ;- .h: +T .h0i ra cc in ;hc. N%u -Nc) .h@y h6a c@a in & h?. &- (In hn -Nc ) .h@y h6a c@a in .h0i ra .h4 (Ong nIc(in ;%. *Ii )\i =han .t +} .ng (np ngOc (\i, +} gi0- )i. V/n )3 *3 0nh h,ng c@a ccin )%n +T (in ;%. nIc &,i cc ;:Whay *Ii ùan )i1- hi>n )\i, &,i cc )\i =han .t cchO= ch/. .rgng hO= caZ c@a ch/. nguyn +inh )< )Ic n6i )%n .rng nhi3u cDng .r4nhnghin cNu. M_. ;hc, c6 .h1 .hTc hi>n +T .c )ng c@a cc )i3u ;i>n Uinh Ujng )%n .r\ng.hi nIc .rng cay, )6 (' 0nh h,ng c@a ch?ng )%n .h'nh =hen cc ch/. h[u c )Oc cay.hng hO=. Nh , .rn )< ch .h/y rvng, +T .h@y h6a h6a hJc c@a cc )\i =han .t chq ^0y ra, nh[ng ch~ c6 nh6- .hVch nIcW=han cTc hay in h6aZ. )6, -Nc ) .h@y h6a cc )\i=han .t =h0i =h{ .huc *' +2 (Ong *' +T =han &h c@a cc nh6- )6. ihi cc (Tc gay ra+T n, =hFng cc ;: cen =h0i ;1 )%n )i>n .Vch c@a cc nh6- =han cTc *' ;hDng =hancTc. 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H'n .'n U hi1u(' -Nc ) .h@y h6a cc hO= ch/. .rgng hO= ca c@a ch/. nguyn +inh .rIc h%. =h0i =h{.huc *' nh[ng &i%n )hi (Ong hO= ch/. .hVch nIc nh/. nh cc =r.:in .an .rng nIc*' =r.:in ;hDng chi%. r?. ra )Oc, cPng nh cc nuc(:=r.:iU, con 0nh h,ng cc &i%n)hi c@a cc hO= ch/. ;hc (' .hN y%u. nh h,ng ) z- )%n h\. )ng h?. nIc chNng . ;hDng nh[ng c6 +T .ha- giac@a c ch% .hz- .h/u -' con c0 c ch% ;hDng .hz- .h/u .rng ù .r4nh )6.E1 )0- &0 h\. )ng nIc )Oc &4nh .hGng, ch?ng .a .\ nh[ng )i3u ;i>nng\i c0nh .2i .hVch ch +T +inh .r,ng =h. .ri1n c@a h> r n6i ring *' .'n & c .h1n6i chung.2.3. Dng lc vn chuyn nuc trong cy.2.3.1. Con dung vn chuyn nuc trong cy.NIc .5 r (n ( =h0i .r0i ùa " )\n )Gng c6 .Vnh ch/. *' ) U'i r/. ;hc nhau.E6 (' cc ù0n &' *' -\ch g~ ch%. )6ng *ai .ro nh 2ng Udn nIc, c6 ) U'i r/. ;hcnhau W*'i c- h_c *'i ch{c c- , cay c, h'ng ch{c h'ng .r- -s. )2i *Ii cay g~Z. E\n)Gng .hN " (' ùa cc .% &' +2ng .5 (Dng h?. )%n -\ch g~ c@a .rung .r{ r *' .5 -\chg~ )%n gan ( ùa .% &' nhu -D ( )%n cc gian &'.2.3.1.1. Con dung vn chuyn qua t bo sng.Nghin cNu ch .a .h/y ch/. nguyn +inh c6 +Nc c0n r/. (In )2i *Ii +T chuy1n*Hn nIc. B,i *4, -_c Ueu ;: nguyn +inh ch/. ngH- nIc r/. -\nh nhng &a nIcùanh cc =han .t (In r/. V. (inh )ng U (Tc h/= Udn .ng h~ -\nh -} c@a nIc *Ii ccg2c .ng Nng .rn cc =han .t )6. fh/. nguyn +inh ngay c0 (?c chNa nhi3u nIc cPng;hDng h3 c6 U\ng nIc h'n .'n .T U. lu .r4nh *Hn chuy1n nIc .rng .% &' +2ngnh (' -. ù .r4nh )hi -Ii .h'nh =hen c@a &a nIc .rng cc -i^:nWSa&inhin, $mmZ.ThTc nghi>- ch .h/y rvng, .rung &4nh (Tc c0n )2i *Ii +T Ui chuy1n nIc ùa .%&' +2ng (' $ a.- $-- )Gng )i.M_. ;hc, nIcc6.h1Uichuy1n.rngcc-aù0nc@a*ch.%&', +ngch?ng chq .hTc hi>n )Oc .5 (Dng h?. )%n ni &4 W, ni &4 c6 ;hung ca+=ri: ;hDng chnIc )i ùaZ. Nh *Hy, , ni &4 +T *Hn chuy1n nIc ùa ch/. nguyn +inh *' ùa *ch .%&' )Oc n2i (i3n *Ii nhau. )ay +T *Hn chuy1n n7h8ng IIIDINH DNG KHONG V NIT (NITROGN! THC VTDnh dung khong v nto ng va tr ac bt quan trng trong sngcathcvt. ukndnhdungkhongvntomttrong nhng nhn t ch ph c hu qu nhat qu trnh snh truong vpht tren ca thc vt. Kh phn tch thnh phn ha hc ca thc vt, ngu ta pht hnra c en hon 60 nguyn t c trong thnh phn ca cy. Tuy nhn chcmt snguyntnhat }nht cnthet chocyg ccnguyn t thet yeu.Mtnguyn t thit yu l nguyn t c vai trsinh l rt quan trng v rt cn cho sinh trng, h!t tri"n # nuthiu, c$y %h&ng th" hon thnh chu tr'nh sng c(a #'nh)Bng phuong php trong cy trong dung d}ch v cc phuong phpnghncu dnhdung chnh xckhc,nguta pht hn rackhong 19 nguyn t dnh dung thet yeu v cy. : C, H, O,N, O, P, K, Mg, Ca, Fe, Cu, Mn, Zn, B.Mo, C, Na, S, N. Kh c cc nguyn t thet yeu v nang uongnh sng, cy c the tong hop cc chat hu co cn thet cho cc hotng snh , qu trnh snh truong pht tren ca cy v hon thnhchu ky sng ca mnh.. .Ngo 19 nguyn t thet yeu ra cy cng cn rat nhu nguynt khc m neu theu cng c nh huong en snh truong pht trenca cy nhung cy van hon thnh chu ky sng ca mnh, van ra hoaket qu.C ha quan nm v nguyn t khong trong cy:Theo quan nm th nhat nguyn t khong cc nguyn t chatrong phn tro ca thc vt. e pht hn cc nguyn t khong cacy, ngu taphntchtrothcvt. t thcvt onht cao(khong 550-600 0C) Cc nguyn t C, O, H, N s mat du dng khCO2,ho H2O, NO2, O2 hoac N2. Phn cn tro thc vt Nguyn t Cchem khong 45%. O chem khong 42%, H khong trn 6,5% v Nkhong 1,5% hm uong chat kh. Cc nguyn t C, H, O, N thnhphn ch yeu cau to nn cc chat hu co trong cy. Chng xm nhpvo cy du dng H2O, kh CO2, O2, NH3, NO3-, s cn , xap x 5%kh uong chat kh ca cy, cc nguyn t khong. V quan emny N khng ph nguyn t khong.Theo quan nm th ha, tr cc nguyn t c nguon gc t CO2 vH2O (C, H v O), cc nguyn t cn uoc cy hap thu t at g cc nguyn t khong.Theo quan nm ny th N nguyn t khong v n uoc r hapthu t at. Do cc phn bn c N (phn m) u uoc g phnkhong. Ouan nm ny hn nay uoc nhu ngu tha nhn.Hmuongccnguyntkhongtrongcykhcnhauratn.Chng phu thuc vo o cy, vo cc b phn khc nhau, vo gaon snh truong...Nhu th nghm chng mnh .rng 95% vt chat trong cy do cy ay t khng kh v nuc, ch 5% ay trong at.Trong thnh phn ha hc ca thc vt, ngu ta thay: - Hm uong cc nguyn t c nguon gc t CO2 v H2OH 6% C 45% O 42%- Hm uong cc nguyn t c nguon gc t atN 1,5% K 1,0% Ca 0,5%Mg 0,2% P 0,1% S 0,1%- Hm uong mt s nguyn t v uongC 100ppm Fe 100ppm B 20ppmMn 50ppm Na 10ppm Zn 20ppmCu 6ppm N 0,1ppm Mo 0,1ppmCan c vo hm uong cha trong cy, ngu ta cha cc nguyn tkhong trong cy thnh ba nhm:- Nhm cc nguyn t uong, c hm uong ben ng t 10-1en 10-4 % chat kh, gom: N, P, K, Ca, S, Mg, S, ...- Nhm cc nguyn t v uong, c hm uong nho t 10-5 en 10-7% chat kh, gom cc nguyn t Fe, Cu, Mn, Zn, Mo, B, Co, T, Sr, Ba,...- Nhm cc nguyn t su v uong c hm uong rat nho, t 10-7en 10-14% chat kh chng gom cc nguyn t Hg, Cd, Cs, I, Pb, Ag,Au, Ra...Cc nguyn t uoc cy hap thu vo c the cva tr khc nhau.Oua phn tch ta thay mc cn thet ca cc nguyn t khong,songcng c mt s nhuA,S,Na cha vuong n trong cy,ngha snh ca chng khng ng ke trong kh mt s nguyn tv uong cn thet cho cy.!. 78 ch9 h:4 th ch:t 0h1/ng.!.!. S th;ch nghi ca ng h?t 0h1/ng.Chcnangquantrngnhatcarhapthunucvcconkhong. R cy c ac em v cau trc hnh th, kh nang snh tr-uongvhotngsnhphhopv chcnanghtnucvhtkhong ca chng.Truc het r c nhng ben o e thch ngh v chc nang hapthu: vch te bo beu b mong, khng tham cutn; t beu b hnh thnhv s ng ht m tang dn tch b mat tep xc ca r n rat n; tebo vo r c nhu khong gan bo e d tr nuc v on khong; tbo n b c a Caspary m cho r c kh nang u chnh dng vtchat vo tru mch dan.R cy c kh nang m su, an rng trong ng at e ch ngtm nguon nuc v chat dnh dung nu cy. Kh nang ny the hn otnh hung nuc v hung ha ca r. R cy c the m su 1,5-2 m,c o r m su t 5- 10 m. R cy thung an rng gap 2-3 n tn ca cy. Nh kh nang phn nhnh mnh, nhat s pht tren cah thng ng ht nn h r c b d tong cng v b mat tep xc vat rat n. S uong ng ht ca r cc o cy rat khc nhau. dchung ca r cc cy trong t t hng chuc tru mha, to nn bmat ht thu n. B mat tep xc ca r thung t cc o ga onra hoa. S xuat hn cc ng ht c d 2-3 m m cho b mat htthu ca r chon t 10-13 n tong the tch ca at. B mat tong cngca r v ng ht t 130 n n hon b mat ca b phn k snh. Hr ca mch en c 13 815 678 r, tong chu d 623 km, bmat tong cng 673.28 m.S phn b ca r trong at ch}u nh huong ca nhu nhn tbn trong v bn ngo.!.(. 78 ch9 h?t 0h1/ng ca h [email protected] vocyth truchetph tantrongdung d}ch at v uoc hap phu trn b mat r. Cc on khong uochap phu trn b mat r theo phuong thc trao o on ga at v nght. C ha phuong thc trao o on: trao o tep xc (trao o trctep) hoac trao o gn tep thng qua H2CO3 trong dung d}ch.Trong qu trnh h hap ca r, CO2uoc to thnh. Trn b matca r s xy ra phn ng: R trao o on H v cc caton, trao o on HCO3- v cc anontrong at. S trao o on ga r v at theo ng ha tr} v uong -uong ca cc on.Chat khong sau kh ht bm n b mat r s uoc vo te bo e vn chuyen vo bn trong r v n cc b phn trn mat at hoac tham ga mt s qu trnh chuyen ha ngay t r. Theo quannm hn nay, qu trnh ht cc chat khong ca cy mt qu trnhsnh rat phc tp, ten hnh theo nhu co che khc nhau va c tnh chat thu ng khng n quan en cc qu trnh trao o chat, va c tnh chat ch ng n quan mt thet en cc qu trnh trao o chat trong thc vt. Sau y ha co che hap thu chat khong co bn: co che thu ng v co che ch ng.!.(.!. 78 ch9 h?t 0h1/ng .h2ng î h c ;ht .ch H" .5 nguyn (i>u hDh/= chuy1n )%n ch w" .\ nIc. Nng (Ong gi0i =h6ng .5 cc =h0n Nng î h c ;ht)6 )Oc c2 )nh (\i .rng (in ;%. gi'u nng (Ong c@a KTP.f6 .h1 n6i chNc nng c &0n c@a hD h/= (' gi0i =h6ng nng (Ong c@a nguyn (i>uhD h/=, chuy1n nng (Ong ;h6 +t U{ng )6 +ang U\ng nng (Ong U +t U{ng ch c .h1(' KTP.5.1.2. Vai tr h hp.HD h/= (' )_c .rng c@a -Ji c .h1 +2ng, (' &i1u hi>n c@a +T +2ng. f .h1 chq .Fn .\i;hi con hD h/=. Tuy nhin , .hTc *H. &n c\nh -_. c6 (Oi c@a hD h/= cPng .Fn .\i nh[ng.c h\i nh/. )nh c@a hD h/=.TrIc h%. (' hD h/= cung c/= nng (OngU\ng KTP ch -Ji h\. )ng +2ng .rngc .h1. MJi h\. )ng +2ng c@a c .h1 )3u cen nng (Ong nhng ;hDng .h1 +t U{ng .rTc.i%= nng (Ong h hJc c@a cc HfHf -' chq +t U{ng nng (Ong U\ng (in ;%. canng c@a KTP U hD h/= .\ ra.Tuy nhin, k ngh!a hD h/= ;hDng chq *3 -_. nng (Ong. Trng hD h/= con .\ ranhi3u +0n =hz- .rung gian c6 *ai .ro ùan .rJng .rng h\. )ng +2ng c@a c .h1. lua hDh/= cc cn )Gng .ra )hi ch/.n2i (i3n *Ii nhau .\ nn .h1 .h2ng nh/. .rng c .h1.Bn c\nh -_. .Vch cTc (' ch@ y%u, hD h/= cPng .h1 hi>n nh[ng -_. .iu cTc, c6 h\inh/. )nh. TrIc h%. hD h/= ('- gi0- cGng ) ùang hO=. HD h/= c'ng ca .h4 ùanghO= &i1u ;i%n c'ng .h/=. E_c &i>. hD h/= +ng ('- gi0- -\nh ùang hO= U =han huQnguyn (i>u ùang hO=, c\nh .ranh nh +ng *Ii ùang hO= ....W^:- =hen ùang hO=Z. 5.2. Cc con dung bin di co cht h hp.Trng ù .r4nh hD h/= nhi3u c ch/. nh g(uî., =r.:in, (i=iU .... )Oc Ugng ('-nguyn (i>u ;h,i )eu. fc c ch/. &vng cc cn )Gng ring &i%n )hi .h'nh cc +0n =hz-.rung gian, .5 )6 .ha- gia *' cn )Gng c@a hD h/= .% &'. f ch/. ch@ y%u c@a hD h/=.% &' (' guc+:. ST &i%n )hi g(uc+: ^0y ra &vng nhi3u cn )Gng ;hc nhau. Tu )S3u;i>n -' hD h/= .i%n h'nh .h: " h4nh .hNcb hD h/= hi%u ;hV WgJi .]. (' hD h/= Z *' hD h/=; ;hV (n -:n W.hGng gJi (' (n -:nZ.5.2.1. H hp hiu kh.HD h/=hi%u ;hV (' ù .r4nh hD h/= c6 +T .ha- gia c@a w", (' ù .r4nh hD h/= ^0yra .rng -Di .rGng hi%u ;hV -Di .rGng c6 w".HD h/= hisu ;hV ^0y ra .rng .hTc *H. *Ii nhi3u cn )Gng ;hc nhaubEGng =han fhu .r4nh fr:&+fhu .r4nh =:n.| =h.=ha..fhu .r4nh g(yî(ic.5.2.1.1. H hp hiu kh theo dung phn - chu trnh Crebs.HD h/= hi%u ;hV ùa )Gng =han *' chu .r4nh fr:&+ (' cn )Gng chVnh c@a hD h/=.% &', ^0y ra =hh &i%n , -Ji +inh *H. *' -Ji .% &'.HD h/= .h: cn )Gng n'y ^0y ra ùa Y giai )\nbc EGng =han .i%n h'nh .rng .% &' ch/..c fhu .r4nh fr:&+ .i%n h'nh .rng c ch/. .y .h1.c ST *Hn chuy1n )i>n .t ^0y ra .rng -'ng .y .h1.`Dungphnb ('giai )\n=hanhuQ=han.tg(uc+:.\raaî. =yru*ic*'NKH". Ei1- )_c &i>. c@a ù .r4nh )Gng =han (' ;hDng =h0S =han .t )Gng .T U=han gi0i -' =han .t )Gng )< )Oc h\. h nhG ù .r4nh =h.=hry( h .\ U\ng)Gng =h.=ha.. , U\ng )Gng =h.=ha. =han .t .r, nn h\. )ng hn U & &i%n )hihn.EGng =han )Oc chia ('- " giai )\n, -~i giai )\n ^0y ra nhi3u =h0n Nng =hNc.\=bc Giai )\n )eu .in (' =han c]. )Gng g(uc+: .h'nh " =han .t )Gng Yfb K(PG *' PK. c Giai )\n hai (' &i%n )hi cc )Gng Yf .h'nh Kî. =yru*ic.i%. ù0 c@a )Gng =han c6 .h1 .6- .]. nh +aubf7H$"w7 x " NK x "KP x "HYPwu=>"fHYfwfwwH x "NKH" x "KTPfc =h0n Nng c@a )Gng =han )Oc .r4nh &'y .h: + )F +au.Trng hD h/= hi%u ;hV Kî. =yru*ic =han huQ .i%= ùa chu .r4nh fr:&+ con "NKH".hTc hi>n chu~i hD h/= )1 .\"H"w."NKH" x w"=> "NK x "H"w.VHy ;%. ù0 c@a chu .r4nh )Gng =han .rng hD h/= hi%u ;hV +} ('bf7H$"w7 x w"=> "fHYfwfwwH x "H"w` Chu trnh Crebsb Sau ;hi )Gng =han =han huQ g(uc+: .\ ra Kî. =yru*ic,.rng )i3u ;i>n hi%u ;hV Kî. =yru*ic .i%= .{c & =han huQ h'n .'n. ST =han huQ n'y^0y ra .h: chu .r4nh )Oc H.fr:&+ *' S.gygy ;h- =h .5 n- $Y8. E6 (' chu .r4nhfr:&+.lu .r4nh =han huQ aî. =yru*ic ùa chu .r4nh fr:&+ ^0y ra .rng c ch/. .y .h1 nhG+T ^?c .c nhi3u h> :n|i-:. B0n ch/. c@a cc =h0n Nng ^0y ra .rng chu .r4nh fr:&+ ch@y%u (' U:cac&^y( h *' U:hyUr h aî. =yru*ic.fhu .r4nh gF- " =henbc Phan huQ aî. =yru*ic .\ fw" *' cc c:n|i-: ;ht.c fc c:n|i-: ;ht .hTc hi>n chu~i hD h/= )1 .\ H"w *' .hng hO= KTP.f ch% chu .r4nh )Oc .r4nh &'y .h: + )F +aubi%. ù0 chu .r4nh ('i%. ù0 chung (' "fHYfwfwwH x mw" => 7fw" x uH"wN%u ;%. hO= *Ii giai )\n )Gng =han f7H$"w7 x w"=> "fHYfwfwwH x H"w)Oc =hng .r4nh .hng ù. c@a hD h/= hi%u ;hVf7H$"w7 x w"=> 7fw" x 7H"wfhu .r4nh fr:&+ .\ uNKH", $yKH"*' $ KTP. fc c:n|i-: ;ht NKH"*'yKH" .hTc hi>n chu~i hD h/= +} .hng hO= KTPbuKNH" Y $" KTP$ yKH" " " KTP$KTP $KTPccccccccccccccccccccccccccccccccc$mKTPNh *Hy cN $ Kî. =yru*ic =han huQ ùa chu .r4nh .\ ra )Oc $m KTP, nn .5 "K.=yru*ic +} .\ )Oc Y# KTP. Trng ch_ng )Gng =han .\ ra )Oc "KTP x "NKH"XKTP.VHyhDh/=hi%u;hV cungc/=ch.%&'YX KTP ;hi =hanhuQ-. =han.tg(uc+:.5.2.1.2. H hp hiu kh qua chu trnh glyoxilic.T5 c ch/. (' ch/. &s & ^y h .\ Kc:.y(cfK. Kc:.y(cfK n'y )Oc &i%n )hi.h: chu .r4nh g(yî(ic ^0y ra .rng g(y^y^-. fhu .r4nh n'y )Oc fn&:c *' ir:&+=h. hi>n *' n- $m8 , nhi3u )2i .Ong nh *i ;huzn, n/- -2c *' nh/. (' , .hTc *H.c6 Ueu. .hTc *H. c6 Ueu ;hi cc aî. &s & ^y h +} .\ ra Kc:.y(cfK. NhG cc:n|i-: c6 .rng g(yî^- nh i|î.ra.a|a .. .. -' Kc:.y(cfK ;hDng &i%n )hi .h: chu.r4nh fr:&+ -' &i%n )hi .h: chu .r4nh g(yî(ic.ihc *Iichu .r4nhfr:&+, ,chu.r4nhg(yî(ic, aî. i|î.ricUIi.c U{ngc@a:n|i-: i|î.ra..a+: =han gi0i .h'nh aî. +ucînic *' aî. g(yî(ic ngng ;%. *Ii Kc:.y(cfK .hN hai nhG -a(a.+in.:.a+: )1 .\ .h'nh aî. -a(ic. T5 aî. -a(ic chu .r4nh .i%= .{c&i%n )hi nh chu .r4nh fr:&+.So d chu trnh duc trnh by nhu sau:fhu .r4nh g(iînic c6 k ngh!a ùan .rJng .rng c .h1 .hTc *H., )_c &i>. , cay chNanhi3u Ueu.c fhu .r4nh g(iî(ic (' ceu n2i gi[a cc ù .r4nh .ra )hi g(uî. *Ii ù .r4nh .ra)hi (i=iU*'ngOc(\i. H\. gi'u(i=iU;hi n0y-e-chu.r4nhg(iî(ich\. )ng)1chuy1n (i=iU .h'nh g(uî. (' c ch/. ch ù .r4nh nay -e-.c fhu .r4nh g(iî(ic (' &i%n U\ng c@a chu .r4nh fr:&+ nn " chu .r4nh c6 .h1 h~ .rO(dn nhau .rng ù .r4nh hD h/= , cay c6 Ueu.c S0n =hz- .rung gian c@a chu .r4nh .ha- gia chu~i hD h/= =h{ .rng .% &' c fc +0n =hz- .rung gian c@a chu .r4nh g(iî(ic .ha- gia *' ùang hD h/=.c S0n =hz- .rung gian c@a chu .r4nh g(iî(ic con .ha- gia nhi3u cn )Gng .ra )hich/.;hc, ('- nguyn (i>u .hng hO= nn nhi3u ch/. ùan .rJng .rng c .h1 .hTc *H. nhch(r=hy(..... 5.2.1.3. H hphiu kh theo chu trnh pentozo-P.Phan huQ g(uc+: ùa )Gng =han ;hDng =h0i (' cn )Gng Uuy nh/. -' con c6 cccn )Gng ;hc .rng )6 =hh &i%n nh/. (' cn )Gng =:n.|cP. fn )Gng =:n.|cP)Oc =h. hi>n )eu .in , n/- -:n, +au c6 , )ng *H. *' cu2i cgng , .hTc *H. cPng .h/yc6+Thi>nUi>nc@acn)Gngn'yWar&ung, fri+.ian, $Y#, Gri+:, $mY, i(::n,$Y7....Z.ihc*Ii )Gng=han, cn)Gng=:n.|cP;hDng=hangi0i g(uc+:.h'nhhai.ri:+ -' g(uc:+ & î h *' U:cac&^y( h )1 .\ ra cc =:n.|cP. T5 cc=:n.|cP .i .\ (\i g(uc|cP. fn )Gng =:n.|cP ^0y ra .rng .% &' ch/. cgng *Ii )Gng=han. VHy y%u .2 n' ùy%. )nh g(uc+: &i%n )2i .h: )Gng =han hay .h: =:n.|cP T5 g(uc|.7P n%u )Oc :n|i-: g(uc|.7P. S|-:ra+: ^?c .c +} &i%n g(uc| 7P.h'nh uruc.| 7P *' )Gng =han +} ^0y ra. fon n%u :n|i-: g(uc| 7P. U:hyUrg:na+:h\. )ng +} î h g(uc| 7P .h'nh aî. 7P c g(ucnic *' cn )Gng =:n.|cP ^0yra.fhu .r4nh =:n.|cP ^0y ra ùa " =henbc Phan huQ g(uc+: .h'nh fw" *' NKPH".c NKPH" .hTc hi>n chu~i hD h/= .\ H"w *' KTP.So d tng qut cua chu trnh nhu sau: i%. ù0 c@a chu .r4nh =:n.|cP ('bf7H$"w7 x 7 H"w => 7fw" x $"H"W=hen $Z$"H" x 7 w"=>$"H"w WPhen "Zccccccccccccccccccccccccccccccccccccccccccccccccccccccccccccci%. ù0 chung (' f7H$"w7 x 7w" => 7fw" x uH"wV3 -_. nng (Ong chu .r4nh =:n.|cP .\ ra $" NKPH". lua chu~i hD h/= .\ raY7 KTP chu .r4nh +t U{ng $ KTP )1 h\. h g(uc+: .h'nh g(uc| 7P. Nh *Hy ;hi =hanhuQ $ g(uc+: ùa chu .r4nh =:n.|cP .\ ra ch c .h1 Ym KTP.fhu .r4nh =:n.|cP c6 k ngh!a nh/. )nh )2i *Ii .hTc *H.bc Eay (' ù .r4nh =han huQ .ri>. )1 f7H$"w7 .hay ch cn )Gng )Gng =han chu.r4nh fr:&+.c Nng (Ong U chu .r4nh cung c/= .ng )ng cn )Gng )Gng =han chu .r4nhfr:&+ nn g6= =hen ùan .rJng .rng *i>c .\ nng (Ong ch c .h1 h\. )ng.cfhu.r4nhh=:n.|cP .\ranhi3u+0n=hz-.runggianùan.rJng, )6('cc)Gng =h.=ha. WfY, fu, fm, f7, f8Z. Nh[ng +0n =hz- .rung gian n'y ('- c ch/. chnhi3u ù .r4nh .ra )hi ch/.;hc c@a c .h1 .hTc *H., )_c &i>. (' chu .r4nh fa(*in .rngluang hO=. fhu .r4nh fa(*in *' chu .r4nh =:n.|cP c6 nhi3u c ch/. gi2ng nhau nn +0n=hz- .rung gian c@a cn )Gng n'y c6 .h1 (Di ;s +ang ('- c ch/. ch cn )Gng ;ia.Ng'i ra cc +0n =hz- .rung gian c@a =:n.|cP con .ha- gia .hng hO= cc hO= ch/. .hNc/=WT5fu.hnghO=cu-arin, (i+u(in, aî. &:n|ic, =(a*Dnic....Z, -. +2aî. a-inWTyr|in .... Z. .hTc *H. chu .r4nh =:n.|cP )Oc .i%n h'nh ch@ y%u , cc -D gi', -D .r,ng.h'nh )< =han h, , cc *gng & &>nh, & .hn .hng c6 hD h/= *%. .hng ^0y ra. fon ,cc -D nn, cac -D &4nh .hGng g(uc+: )Oc =han huQ ch@ y%u .h: cn )Gng )Gng=han chu .r4nh fr:&+.5.2.2. H hpky kh - ln men.HD h/=; ;hV (' ù .r4nh =han huQ g(uc+: .rng )S3u ;i>n ;hDng c6 w" .ha- gia.Giai )\n )eu c@a hD h/= ; ;hV (' )Gng =han. Tuy nhin .rng hD h/= ; ;hV )Gng=han chq ^0y ra giai )\n =han huQ g(uc+: .h'nh Kî. =yru*ic *' NKH" con giai )\nNKH".hTc hi>n chu~i hD h/= ;hDng ^0y ra U ;hDng c6 w". B,i *Hy ;%. ù0 )Gng=han .rng hD h/= ; ;hV ('bf7H$"w7 => "fHYfwfwwH x"NKH"Giai )\n hai c@a hD h/= ; ;hV (' &i%n )hi aî. =yru*ic .h'nh cc +0n =hz- nh:.an(, aî. (ac.ic, .... Eay (' ù .r4nh (n -:n. Tu .h: +0n =hz- c@a ù .r4nh -' c6cc ù .r4nh (n -:n ;hc nhau nh (n -:n rOu, (n -:n (ac.ic ....5.2.2.1. Ln men ruu.ST (n -:n rOu ^0y ra ùa Y giai )\n chVnhbcThuQ =han .inh &. .h'nh g(uc+: Wn%u c ch/. (' .inh &.Z.cEGng =han g(uc+: .h'nh aî. =yru*ic *' NKH".cLn -:n rOu .hH. +T.Giai )\n (n -:n rOu ^0y ra " =h0n Nngb "fHYfwfwwH=> fHYfHw x fw"fHYfHw xNKH" => fHYfH"wH x NK.Nh *Hy ;%. ù0 chung c@a .'n & ù .r4nh (n -:n rOu ('f7H$"w7x "NK => "fHYfwfwwH x"NKH""fHYfwfwwH =>"fHYfHw x "fw""fHYfHw x"NKH" =>"fHYfH"wH x "NK.cccccccccccccccccccccccccccccccccccccccccccccccccccccccccccf7H$"w7 => "fHYfwfwwH x"fw" V3 -_. nng (Ong (n -:n rOu chq .\ ra )Oc "KTP .rng giai )\n )Gng =hannn hi>u ù0 nng (Ong r/. .h/=. T5 $ g(uc+: chuy1n .h'nh " :.an( nng (Ong .T Ugi0- "m7 ica( chq .\ ra )Oc "KTP W.ng )ng $u,7 ica( .rng )S3u ;i>n chuznZ.Hi>u +u/. nng (Ong chq )\. "7n. S *Ii hD h/= hi%u ;hV chq &vng mn. W"KTP c@a (n-:n + *Ii YX KTP c@a hD h/= hi%u ;hVZ. S, U! nh *Hy *4 +0n =hz- (n -:n rOu conchNa nng (Ong ;h (In, =hen nng (Ong gi0i =h6ng ra .5 g(uc+: )1 .\ :.an( chq ('-. =hen nh W$u ica(78u ica(Z. Nh *Hy )1 c6 nng (Ong ch c .h1 h\. )ngnh hD h/= hi%u ;hV cung c/= .h4 (n -:n rOu =h0i +t U{ng (Ong c ch/. nhi3u g/= u#cm# (en. *Hy nn hD h/= ; ;hV W(:n -:nZ ;s U'i +} ('- ch cay &i )6i, -D & +u/. ccch/. .Vch (u &,i cc ù .r4nh ;hc Wluang hO=, HD h/= hi%u ;hVZ.5.2.2.2. Ln men lactic.fPng nh (n -:n rOu, (n -:n (ac .ic (' ù .r4nh hD h/= ; ;hV ;h =hh &i%n ,.hTc *H..lu .r4nh(n -:n (ac .ic ^0y ra .h: " cn )Gng ;hc nhaubc Trnggiai)\n)Gng=han+au;hi .\ K(PG,K(PG;hDng&^yh.h'nhK$YPG -' &i%n )hi .rTc .i%= .h'nh aî. (ac .icbNh *Hy .h: cn )Gng n'y .5 g(uc+: .\ ra " aî. (ac .ic *' .iu .2n -/. " KTP.rng giai )\n )eu c@a )Gng =han.c EGng =han .\ ra fHYfwfwwH *' NKH", NKH" ;ht aî. =yru*ic .h'nh aî.(ac .ic.f7H$"w7x "NK => "fHYfwfwwH x"NKH""fHYfwfwwH x "NKH" =>"fHYfHwHfwwH x "NKccccccccccccccccccccccccccccccccccccccccccccccccccccccccccccccccf7H$"w7=> "fHYfHwHfwwHV3 nng (Ong cn )Gng n'y .\ ra )Oc " KTP nh .rng (n -:n rOu.5.2.3. H hp sng.5.2.3.1. D}c dim.:c;:r W$mmZ, :(i..ch W$7Z )< =h. hi>n ra hi>n .Ong .h0i fw"+au -. .hGigian chi%u +ng , -. +2 cay. Nh *Hy , nh[ng cay n'y cc +0n =hz- + c/= c@a ùanghO= )< & =han huQ .h'nh fw" ng'i +ng. ST h/= .h{ w" cgng *Ii +T .h0S fw" ^0y ra =h{.huc *' nh +ng nn )Oc gJi (' hD h/= +ng Wùang hD h/=Z. Nh[ng cay n'y hD h/=)Fng h'nh *Ii ùang hO=.f6 .h1 =han &i>. hD h/= +ng *Ii hD h/= .2i nhG .Vnh nh\y c0- c@a ùang hD h/=*Iicc y%u .2 -Di .rGng.c HD h/= (uDn )Fng &i%n *Ii cGng ) nh +ng, con hD h/= .2i ;hDng chu 0nhh,ng c@a nh +ng. nh +ng *Ii m#c8##n- c6 hi>u ù0 ca *Ii hD h/= +ng.c HD h/= gi0- ;hi .Q (> ^y .h/= W "nZ ;hi h'- (Ong w"c'ng ca hD h/= +ngc'ng -\nh. ihi .ng h'- (Ong w" .5 "$n )%n $##n hD h/= +ng .ng g/= "cY (en. c Tng h'- (Ong fw" +} h\n ch% hD h/= +ng, ;hi h'- (Ong fw" ca hn #,$nhD h/= +ng gi0- -\nh *' c6 .h1 ng5ng ;hi h'- (Ong fw" )\. $c"n. fon h'- (Ongfw" ca V. 0nh h,ng )%n hD h/= .2i.c HD h/= +ng nh\y *Ii nhi>. ) hn + *Ii hD h/= .2i. fc nh6- .hTc *H. ;hc nhau c6 -Nc ) hD h/= +ng ;hDng gi2ng nhaubc fay fYc6 hD h/= +ng -\nh. VV U{ , (?a, )Hu, c0i )Gng, hIng Ung, .hu2c( ... c6 hD h/= .2i ;h0ng $cY-g fw"U-"h. fon hD h/= +ng -\nh g/= "cY (en h/= .2i)6.c fay fu nhb ngD, -Va, ca (ng .... ;hDng c6 hD h/= +ng h_c ^0y ra y%u ;hDng.h1 ^c )nh )Oc. *Hy nh6- cay n'y c6 nng +u/. ca hn cay fY.c fay fKM c6 ùang hD h/= y%u *' .hay )2i nn ;h6 ^c )nh.NgGi .a ch rvng nguyn nhan ('- ch ù .r4nh hD h/= +ng , nh6- .hTc *H. fuy%u hay ;hDng c6 (' U h\. .Vnh c@a îg:na+: , nh6- cay n'y y%u U .Q (> fw"w".rng .% &' &a &6 -\ch ca )i3u )6 gi?= ch h\. .Vnh cac&^y( h -\nh hn h\..Vnh ^y h. M_. ;hc ;hi .h0i fw" .5 .% &' &6 -\ch (H= .Nc )Oc KTP .5 .% &'.h.(.i%= nhHn, U )6 ('- gi0- hD h/= +ng.5.2.3.2. Co ch.luang hD h/= ^0y ra .\i Y &' ùan ;hc nhaub ({c (\=, =:rî^- *' .y .h1. T% &'ch/. (' -Di .rGng )1 cc ch/. )i ùa .5 &' ùan n'y +ang &' ùan ;hc.` Lc lp: T\i ({c (\= Uin ra ù .r4nh ^y h 9i&u(| $,m UiP U 9i&u(| $,mUiPc^yUa+: ^?c .c .\ nnaî. g(yc:ric *'aî. g(yc(ic. EFng .hGi aî. g(yî(ic .5 .y.h1 )a +ang cPng )Oc ;ht .h'nh aî. g(yc(ic. K.g(ic(ic chuy1n +ang =:rî^- )1.i%= .{c &i%n )hi .h: hD h/= +ng.` Peroxixom:)ay (' &' ùan &i%n )hi H"w"nn )Oc gJi (' =:rî^-. T\i )ayK.g(yc(ic & î h .h'nh K.g(yî(ic nhG g(yc(a.c^yUa|a. H"w")Oc .\ ra U îh aî. g(ic(ic +} & =han huQ &,i ca.a(a|a .h'nh H"w *' w". Ti%= .h: (' cc =h0n Nngchuy1n a-in )1 .\ g(ycin. G(ycin ùay

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