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SINH L HC T BO
I. Vn chuyn vt cht qua mng t bo
1. Cu to c bn ca mt t bo ng vt Mt t bo ng vt in hnh c th chia lm 4 phn c bn (1) Mng bo tng: mng ngn cch thnh phn ni bo vi thnh phn vt cht v mi trng bn ngoi t bo. (2) Dch t bo (cytosol): l mt dch keo cha nhiu loi protein, enzym, cht dinh dng, cc ion v cc phn t nh ha tan khc nhau, tham gia vo cc qu trnh chuyn ha khc nhau ca t bo. Cc bo quan v th vi nm l lng trong dch t tng. T bo tng (cytoplasm) dng bao hm c dch t bo, tt c cc bo quan (tr nhn) v cc th vi. (3) Cc bo quan: gm cc cu trc c hnh dng v chc nng c trng, bao gm c nhn. (4) Cc th vi (inclusions): Cc cu trc c mt khng thng xuyn trong dch bo tng, cha cc sn phm bi tit hoc cc cht d tr ca t bo. 2. Mng bo tng

Hnh 1: Cu trc ca mng bo tng

1: knh; 2: l; 3: cholesterol; 4: protein ngoi vi; 5: protein xuyn mng; 6: lp kp phospholipid; 7: phn a nc ca phospholipid; 8: glycoprotein; 9: glycolipid; 10: protein ngoi vi; 11: dch ngoi bo; 12: bo tng; 13: phn k nc ca phn t phospholipid;

2.1. Cu trc - Cu trc ca mng bo tng (hnh 1) l mt cu trc dng khm lng (fluid mosaic model) vi cc phn t protein nm xen k trn mt mng kp lipid. - Mng bo tng ca cc t bo ng vt in hnh c t l v mt khi lng gia protein v lipid xp x 1: 1 v t l v mt s lng phn t gia chng l 1 protein: 50 lipid.

- Thnh phn lipid rt t thay i gia cc loi mng bo tng khc nhau nhng thnh phn protein c s thay i rt ln v ng vai tr quyt nh trong hot ng chc nng ca t bo. 2.1.1. Thnh phn lipid ca mng - Phospholipid: Chim 75% thnh phn lipid ca mng. Cc phn t phospholidid vi c im cu trc mt u phn cc (u a nc do c cha phosphat) v mt u khng phn cc (u k nc do cha 2 ui acid bo) to thnh mt lp lipid kp vi 2 u k nc quay vo nhau to thnh b khung ca mng bo tng. Cc phn t phospholipid c th di chuyn d dng gia 2 lp v thay i ch cho nhau to nn tnh linh hot cho lp lipid kp. Mng ny c kh nng t hn gn khi b thng. - Glycolipid: Chim khong 5% thnh phn lipid ca mng, cng c cu trc phn cc nhng ch c phn tip xc vi dch ngoi bo ca mng bo tng. Chc nng cha r, c l lin quan n vic ghi nhn v truyn t thng tin gia cc t bo, tham gia vo cc c ch iu ha s sinh trng v pht trin ca t bo. - Cholesterol: Ch chim 20% thnh phn lipid ca mng bo tng, loi lipid ny khng c mng bo tng ca t bo thc vt. Cu trc dng vng ca nhn steroid trong cu trc ha hc ca cholesterol tng tnh vng chc nhng li lm gim i tnh mm do ca mng t bo ng vt. 2.2. Thnh phn protein ca mng 2.1.2.. Phn loi - Da vo cch thc phn b trn mng m cc protein c chia lm 2 loi: + Protein xuyn mng (integral protein): Nm xuyn qua chiu dy ca lp lipid kp, hu ht l cc glycoprotein vi thnh phn ng nm quay ra pha ngoi ca mng t bo. + Protein ngoi vi (peripheral protein). Ch gn lng lo vi mt ngoi hoc mt trong ca mng lipid kp. 2.1.3. Chc nng

Hnh 2: Cc chc nng ca protein trn mng

a: knh; b: cht vn chuyn; c: receptor; d: enzyme; e: neo khung xng t bo; f: du nhn dng t bo. 1: dch ngoi bo; 2 mng bo tng; 3: bo tng; 4:ligand; 5:c cht; 6: sn phm; 7:vi si; 8: protein MHC

- Cc protein trn mng bo tng c nhng vai tr nh sau trong hot ng sng ca t bo (hnh 2): + Cc knh: l nhng l nm xuyn qua cc protein xuyn mng cho php mt s cht nht nh i ra ngoi hoc vo bn trong t bo. + Cht vn chuyn: l nhng protein xuyn mng thc hin vic vn chuyn cc cht t pha ny sang pha khc ca mng t bo. + Cc receptor: l cc protein xuyn m ng c vai tr xc nh cc phn t c hiu nh horrmon, cht dn truyn thn kinh v.v..., gn vi chng qua khi ng mt s cc hot ng chc nng ca t bo. + Cc enzyme: c th l protein xuyn mng hay protein ngoi vi, xc tc cho cc hot ng sinh ha din ra trn mng. + Cc neo khung xng t bo: l cc protein ngoi vi mt trong ca mng bo tng, y l v tr gn ca cc vi si lm hnh thnh nn khung xng ca t bo. + Cc du nhn dng t bo (cell identity markers: CIM): ng vai tr ca cc du nhn dng t bo, thng c cu to glycoprotein hoc glycolipid. Gip t bo ca c th nhn bit c t bo cng loi trong qu trnh to m cng nh nhn dng v p ng vi cc t bo l. 2.3. Chc nng ca mng bo tng 2.3.1. Thng tin

Hnh 3: Gradient in - ha
a: m hnh chi tit; b: m hnh n gin 1: dch ngoi bo; 2: mng bo tng; 3: bo tng

- Mng bo tng c chc nng thng tin t bo, bao gm vic tng tc vi cc t bo khc trong c th, vi cc t bo l v cc ligand nh cc horrmon, cc cht dn truyn thn kinh, cc enzyme, cc cht dinh dng v cc khng th trong dch ngoi bo 2.3.2. Duy tr gradient in - ha - Mng bo tng duy tr mt s khc bit v nng ca cc cht ha hc v cc ion gia hai bn mng to nn mt gradient in - ha gia bo tng v dch ngoi bo (hnh 3)(bng 1).
Bng 1: Cc thnh phn ion chnh trong dch ni bo v ngoi bo Ion Na+ K+ Ca2+ Mg Cl
2+

Nng ni bo (mEq/l) 15 135 2.10-4 40 4 10 20 4 152

Nng ngoi bo (mEq/l) 142 4 4 2 106 24 4 1 1

HCO3 HPO4 2SO4

2-

Cc protein- , cc amino acid , urea v.v...

- Trong dch ngoi bo cation chnh l Natri (Na+) v anion chnh l Clo (Cl-) trong khi trong bo tng cation chnh l Kali (K+) v 2 loi anion chnh l cc phosphat hu c (cc nhm PO43- gn vo cc phn t hu c nh ATP) v cc acid amin mang in tch m trong cu trc ca cc protein. - S khc bit v nng ca cc ion lm cho mt trong ca mng m hn so vi pha ngoi mng. 2.3.3. Tnh thm chn lc - Mng bo tng cho php mt s cht i qua nhng li khng cho hoc hn ch s vn chuyn qua mng ca mt s cht khc, tnh cht ny c gi l tnh thm chn lc. Tnh cht ny ph thuc vo cc yu t sau ca cht c vn chuyn: + Kh nng tan trong lipid: cc cht tan trong lipid (khng phn cc, cc phn t k nc) d dng i qua lp phospholipid kp ca mng bo tng. + Kch thc: Hu ht cc phn t c kch thc ln nh cc protein u khng th i qua mng bo tng. + in tch: Lp phospholipid kp ca mng bo tng khng thm vi tt c cc phn t phn cc. Tuy nhin mt s cc cht mang in tch c th qua mng nh cc knh xuyn mng hoc thng qua cc cht vn chuyn. in th m hn bn trong mng lm tng dng chy ca cc cation vo pha trong mng v cn tr s i vo ca cc anion.

+ S c mt ca cc knh v cc cht vn chuyn c hiu trn mng: Cc knh v cc cht vn chuyn trn mng gip cc cht phn cc hoc mang in tch nh cc ion c th i qua mng, hu ht chng u c tnh chn lc rt cao, mi loi ch phc v cho mt cht nht nh. + Nc l mt phn t c bit, c th i qua mng bo tng mt cch d dng hn tt c cc cht khc. 3. S vn chuyn cc cht qua mng bo tng - S vn chuyn qua mng c thc hin thng qua 3 hnh thc chnh: (1) vn chuyn th ng (passive transport), khng tiu tn nng lng, (2) vn chuyn ch ng (active transport), cn tiu tn nng lng v (3) hnh thc vn chuyn bng cc ti (vesicular transport). 3.1. Cc hnh thc vn chuyn th ng 3.1.1. Khuch tn n gin (simple diffusion) - Khuch tn n gin l hnh thc khuch tn trong cc phn t vt cht c vn chuyn t ni c nng cao n ni c nng thp v khng tiu tn nng lng. - S khc bit v nng ca mt cht 2 bn mng bo tng to nn mt gradient nng . S khc bit ny lm cho cc phn t cht i t ni c nng cao n ni c nng thp cho ti khi t ti s cn bng ng hai bn mng m khng i hi phi cung cp nng lng.

Hnh 4: S khuch tan qua mang bao tng

a: khuch tan qua lp lipid kep; b: khuch tan qua knh 1: dch ngoai bao; 2: mang bao tng; 3: bao tng; 4: lp phospholipid kep; 5:H20, O2, CO2, N2, cac steroid, cac vitamintan trong m, glycerol, ru, ammonia; 6: knh; 7: l; 8: protein xuyn mang

- Sau khi t c cn bng, s khuch tn ca cc phn t vn c tip tc duy tr tuy nhin nng ca chng hai bn mng khng thay i. - Hin tng ny ph thuc vo ng nng (kinetic energy) ca cc phn t nn s

khuch tn s xy ra nhanh hn khi (1) nhit tng, (2) gradient nng ln v (3) vt th c kch thc nh. - Cc phn t tan trong lipid nh oxygen, doxide carbon, nitrogen, cc steroid, cc vitamin tan trong lipid nh A, D, E v K, glycerol, ru v ammonia c th dng i qua lp phospholipid kp ca mng bo tng theo c 2 pha bng hnh thc ny (hnh 4). Tc khuch tn ca chng t l thun vo kh nng tan trong lipid ca cc phn t. - Cc phn t c kch thc nh khng tan trong lipid cng c th khuch tn qua mng theo hnh thc ny thng qua cc knh (hnh 4), nh cc ion natri (Na+), ion kali (K+), ion calci (Ca2+), ion clo (Cl-), ion bicarbonate (HCO3-) v ur. Tc khuch tn ca chng t l thun vi kch thc phn t, hnh dng v in tch ca cc phn t. - Nc khng nhng d dng i qua lp phospholipid kp m cn khuch tn qua cc knh ny. 3.1.2. Hin tng thm thu (hnh 4)(osmosis) - Hin tng thm thu l hin tng vn chuyn th ng ca cc phn t nc t ni c nng nc cao (c nng cht ha tan thp) ti ni c nng nc thp (c nng cht ha tan cao). Mt dung dch c nng cc cht ha tan cng cao th p lc thm thu cng ln v ngc li. - Gradient p lc thm thu c hnh thnh hai bn mng do s c mt ca cc cht ho tan vi cc nng khc nhau mi bn. - Di tc ng ca p lc thm thu nc s di chuyn t ni c p lc thm thu thp n ni c p lc thm thu cao t n s cn bng p lc thm thu. - Bnh thng p lc thm thu trong t bo cn bng vi p lc thm thu trong dch ngoi bo nh th tch ca t bo duy tr c s hng nh mt cch tng i, trong khi p lc thm thu ca huyt tng li cao hn so vi dch k bao quanh cc thnh mao mch, s khc bit ny lm nc s di chuyn t pha m k v trong lng mao mch. Cc tnh hung lm gim p lc thm thu ca huyt tng s lm tr nc trong dch k v dch ngoi bo. 3.1.3. Hin tng khuch tn qua trung gian (facilitated diffusion) - Hin tng khuch tn qua trung gian (hnh 5) l hin tng khuch tn ca cc cht t ni c nng cao n ni c nng thp nh vai tr trung gian ca cc protein ng vai tr cht vn chuyn trn mng bo tng. Tc ca kiu khuch tn ny ph thuc vo s khc bit v nng ca cht c vn chuyn hai bn mng v s lng ca cc cht vn chuyn c hiu. - Trong c th cc ion, ur, glucose, fructose, galactose v mt s vitamin khng c kh nng tan trong lipid i qua lp phospholipid kp ca mng s di chuyn qua mng theo hnh thc ny. - V d: Glucose l mt trong nhng cht quan trng i vi hot ng sng ca t bo c vn chuyn vo theo hnh thc khuch tn qua trung gian i vo trong t bo, qu trnh ny din ra theo cc bc trnh t nh sau:

Hnh 5: S khuch tan qua trung gian

1: dch ngoai bao; 2: mang bao tng; 3: bao tng; 4: protein xuyn mang; 5: gradient nng

+ Glucose gn vo cht vn chuyn c hiu pha bn ngoi mng, cc cht vn chuyn ny khc nhau ty theo tng loi t bo. + Cht vn chuyn thay i hnh dng. + Glucose i qua mng v gii phng vo trong t bo, ti y enzyme kinase s gn mt nhm phosphat vo phn t glucose to thnh glucose 6-phosphate. Phn ng ny gip duy tr nng glucose trong t bo lun lun mc thp to iu kin cho glucose lun lun c vn chuyn vo bn trong. 3.2. Cc hnh thc vn chuyn ch ng - Hnh thc vn chuyn ch ng l hnh thc vn chuyn tiu tn nng lng ATP nhm a cc cht i ngc li chiu gradient nng ca chng. - Hnh thc vn chuyn ny c thc hin qua vai tr ca cc protein xuyn mng c hiu ng vai tr nh cc bm hot ng nh ATP y cc ion nh Na+, K+, H+, Ca2+, I-, Cl- hoc cc phn t nh nh cc acid amin, cc monosaccharide i ngc li chiu gradient nng ca chng. - Hnh thc vn chuyn ny c chia lm hai loi (1) vn chuyn ch ng nguyn pht v (2) vn chuyn ch ng th pht ty theo nng lng ATP c s dng trc tip hay gin tip trong qa trnh vn chuyn cc cht. 3.2.1. Vn chuyn ch ng nguyn pht (primary active transport) - Vn chuyn ch ng nguyn pht l hnh thc vn chuyn trong nng lng t ATP c s dng trc tip "bm" mt cht qua mng theo chiu ngc vi chiu gradient nng . - T bo s s dng nng lng ny thay i hnh dng ca cc protein vn chuyn trn mng bo tng qua thc hin vic vn chuyn. Khong 40% ATP ca t bo phc v cho mc ch ny.

Hnh 6: Hoat ng cua Bm natri - kali

- Bm natri (hnh 6) l mt v d in hnh cho hnh thc vn chuyn nguyn pht: + Qua hot ng ca bm natri, cc ion natri (Na+) s c "bm" ra khi t bo (ni c nng ion natri cao hn) v ion kali (K+) s c "bm" vo trong t bo (ni c nng ion kali cao hn). + Bng cch ny bm natri s duy tr c nng n nh ca ion natri v kali trong v ngoi t bo, iu ny rt quan trng cho hot ng sng ca t bo. + Tt c cc t bo u c bm natri, trn mi micro mt vung mng bo tng c ti hng trm bm nh vy v chng phi hot ng lin tc duy tr s n nh ca cc ion Na+ v K+ do cc ion Na+ v K+ lin tc khuch tn qua mng thng qua cc knh lm ph v trng thi n nh ca cc ion ny. + Bm natri i khi cn c gi l bm Na+/K+ ATPase do protein thc hin vn chuyn hot ng nh mt enzyme tch nng lng t ATP. Trong cu trc ca phn t

ATPase gm c 4 tiu n v (2 n v v 2 n v ). Cc tiu n v c hot tnh enzym chuyn ATP thnh ADP gii phng nng lng v trn chng c c cc v tr gn vi cc ion pha trong v ngoi t bo. Pha trong t bo c cc v tr gn 3 ion Na+ v ATP, pha ngoi t bo c cc v tr gn vi 2 ion K+. + Qu trnh hot ng ca bm c th chia lm hai giai on: (1) Khi ba ion Na+ v ATP gn pha mt trong ca bm, mt nhm phosphate c chuyn t phn t ATP ti gc acid aspartic ca tiu phn . S c mt ca nhm phosphate giu nng lng s lm thay i cu trc ca bm lm chuyn 3 ion Na ra pha ngoi t bo. (2) Khi 2 ion K+ gn vo pha mt ngoi t bo, lin kt gia nhm phosphate v acid aspartic b thu phn. Nng lng c gii phng t qu trnh dephosphoryl (dephosphorylate) ny s lm thay i cu trc ca bm ln th hai lm cho 2 ion K+ c a vo bn trong t bo. + S c ch hot ng ca bm: Bm s khng hot ng nu nng ca cc ion Na+, K v ATP qu thp. Tc dng ca digitalis, mt loi thuc c s dng trong vic iu tr suy tim, da trn kh nng kt hp vi tiu phn pha mt ngoi t bo v qua can thip vo qu trnh dephosphoryl ca bm lm c ch hot ng ca bm.
+

- Ngoi bm Na+/K+, hin tng vn chuyn ch ng nguyn pht cn c thy trong hot ng ca bm K+/H+ trn mng t bo nim mc d dy, iu khin vic bi xut ion H+ vo d dy trong qu trnh tiu ho, bm Ca2+ c trn h li ni sinh cht ca cc t bo c duy tr nng ion Ca2+ trong t bo lun lun di mc 0,1(mol/L. 3.2.2. Vn chuyn ch ng th pht (secondary active transport) (hnh 7) - Trong hnh thc vn chuyn ny nng lng tn tr do s khc bit v gradient nng ca ion Na+ c s dng vn chuyn cc cht i ngc li chiu gradient nng ca chng qua mng. - Bm natri duy tr mt s khc bit ln v nng ion Na+ hai bn mng bo tng, nu c mt con ng qua cho php cc ion Na+ i t ni c nng cao n ni c nng thp th nng lng tn tr do s khc bit v nng ca Na+ s c chuyn thnh ng nng gip vn chuyn mt cht khc i ngc li chiu gradient nng ca cht . - V s khc bit nng ca ion Na+ c thit lp qua hnh thc vn chuyn ch ng nguyn pht, i hi ATP mt cch trc tip nn c th coi hnh thc vn chuyn th pht s dng ATP mt cch gin tip thc hin vic vn chuyn ch ng qua mng. - Nhiu loi ion v cht dinh dng c vn chuyn bng hnh thc ny: + S vn chuyn glucose, galactose v cc acid amin cng vi ion Na+ i qua mng t bo lp mt trong rut non v cc t bo ca ng thn din ra theo cch ny, qua cc cht dinh dng trong thc n c hp thu mt cch trit ti rut non v c ng thn ti hp thu a tr li vo mu. + S vn chuyn ion Ca2+ ra ngoi bo tng ca cc t bo tm tht v cc loi t bo c khc (s vn chuyn ny ca ion Ca2+ phi hp vi hot ng bm Ca2+ trn li ni sinh cht ca cc t bo c s gy ra tnh trng gin c)

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+ Cc ion H+ hnh thnh trong qu chuyn ha t bo c bm ra khi t bo theo hnh thc vn chuyn ny. C ch ny ht sc quan trng duy tr pH n nh trong t bo v trong lng ng ln gn ca thn ( gip ti hp thu bicarbonate).

Hnh 7: Hin tng ng vn v i vn

a: hin tng ng vn; b: hin tng i vn 1: dch ngoi bo; 2: mng bo tng; 3: bo tng; 4: protein ng vn; 5: amino acid; 6: ion Natri; 7: ion calcium; 8: protein i vn; 9:khuch tn th ng theo chiu gradient nng ; 10: vn chuyn ch ng th pht.

- Nng lng tn tr do gradient in ha ca ion Na+ s lm thay i cu hnh ca protein vn chuyn + Khi ion Na+ gn vi protein vn chuyn s lm tng i lc ca protein ny vi cht c vn chuyn. + Khi c ion Na+ v cht c vn chuyn gn vo protein vn chuyn s lm thay i cu trc ca protein ny gip ion Na+ v cht c vn chuyn c a qua mng. + Khi hai cht c vn chuyn theo cng mt hng qua mng th qu trnh ny c gi l hin tng ng vn (symport) nh s vn chuyn ca glucose, cc acid amin qua nim mc rut v ng thn. + Khi hai cht c vn chuyn theo hai hng khc nhau qua mng th qu trnh ny c gi l hin tng i vn (antiport) nh s vn chuyn ch ng ca ion Ca2+ ion H+ qua mng. - S chnh lch v nng ion Na+ hai bn mng cng ln th s vn chuyn ch ng th pht xy ra cng nhanh. 3.3. Hnh thc vn chuyn bng cc ti - y l hnh thc vn chuyn cho php cc phn t c kch thc ln c th i qua c mng t bo, hnh thc ny gm c (1) Hin tng nhp bo (endocytosis) bao gm hin tng thc bo (phagocytosis), hin tng m bo (pinocytosis), hin tng nhp bo qua trung gian receptor. (2) Hin tng thi bo (exocytosis)

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3.3.1. Hin tng nhp bo - Thnh phn vt cht ngoi bo c a vo trong cc ti c to thnh t s lm vo ca mng t bo - Trong bo tng cc ti nhp bo s ho ln vi lysosome, cc thnh phn trong ti nhp bo s b thy phn bi cc enzyme v cc n phn s c a vo trong dch ni bo. Hin tng thc bo (hnh 8)

Hnh 8: Hin tng thc bo

1: vi khun hoc vt th l; 2: gi tc; 3: ti thc bo; 4: mng bo tng.

- Bo tng v mng t bo to thnh cc gi tc m ly vt th bn ngoi t bo vi vt th ny vo trong lng bo tng, ti y vt th c bc trong lp mng xut pht t mng bo tng v c gi l ti thc bo (phagocytic vesicle) hay phagosome. - Ch c mt s t bo trong c th thc hin chc nng thc bo. Cc thc bo quan trng nht l cc bch cu trung tnh v i thc bo (macrophage). Hin tng thc bo gip a cc vi khun, cc mnh vn t bo vo bn trong cc t bo c kh nng thc bo. Hin tng m bo: - Hin tng qua cc dch ngoi bo v cc phn t ha tan pha ngoi t bo c a vo bn trong t bo. y l chc nng c thy mi loi t bo ca c th. - m bo c thc hin n gin qua s lm vo ca mng bo tng to nn ti m bo (pinocytic vesicle) mang cc ht dch vo trong lng bo tng. Hin tng nhp bo qua trung gian receptor (hnh 9): - Hin tng ny din ra tng t nh hin tng m bo nhng c tnh chn lc cao, trong t bo la chn cc phn t hay cc vt th c hiu a vo trong bo tng, nh mc du nng ca chng trong dch ngoi bo rt thp nhng chng vn c th i c vo bn trong t bo thng qua cc protein xuyn mng ng vai tr receptor c hiu cho chng trn mng bo tng. - Cht c i lc vi mt receptor c hiu c gi l ligand, nhng cht ny c th l cholesterol, st v cc vitamin rt cn thit cho cc qu trnh chuyn ha trong t bo hoc cc

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horrmon v.v... nhng i khi y cng l ng xm nhp ca mt s loi virus trong c virus HIV (human immuno- deficiency virus).

Hnh 9: Hin tng nhp bo qua trung gian receptor

a: ligand; b: receptor; c: enzyme tiu ha;d: ligand ging ha

- Qu trnh nhp bo din ra theo cc bc sau: (1) Ligand gn vi receptor c hiu ca n pha ngoi ca mng bo tng. (2) S kt gn ny lm vng mng bo tng v tr phc hp ligand - receptor lm xung to thnh ti nhp bo (endocytosis vesicle) mang phc hp ni trn. (3) Trong lng bo tng cc ti nhp bo ha ln vi nhau to nn mt cu trc ln hn gi l endosome. (4) Trong endosome cc receptor tch khi cc ligand v chia lm hai phn, phn endosome ch cha cc receptor v phn endosome ch cha cc ligand. (5) Phn endosome cha cc receptor s ho nhp tr li vi mng bo tng tip tc nhim v.

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(6) Phn endosome cha cc ligand s ha nhp vi cc lysosome v cc enzyme ca bo quan ny s phn hy cc ligand s dng cho cc hot ng sng khc ca t bo. 3.3.2. Hin tng thi bo - Hin tng thi bo l hin tng cc cu trc c gi l ti tit (secretory vesicle) c to thnh trong lng bo tng tin ti v ha nhp mng ca ti vo mng bo tng a cc thnh phn bn trong ti vo dch ngoi bo.