Bi 1: Nhp mn sinh hc phn t _ tp 1 Bi 2: Mng sinh cht _ tp 1 Bi 3: Vn chuyn vt cht qua mng _ tp 1 Bi 4: Tiu th v Peroxisomes _ tp 2 Bi 5: Li ni sinh cht v Golgi_

tp 2 Bi 6: Ty th _ tp 2 Bi 7: Nhn t bo gian k _ tp 3 Bi 8: B xng t bo _ tp 3 Bi 9: S phn bo _ tp 3

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NHP MN SINH HC T BO - PHN T

1. Lch s - Phng php nghin cu - Knh hin vi quang hc - Knh hin vi in t - Phng php phn tch & ti lp rp - Phng php ti t hp ADN 2. i tng - Mc tiu mn hc - nh ngha t bo - Hc thuyt t bo - Cc thuc tnh ca t bo - i tng - Mc tiu 3. Cc vn chung - Thuyt tin ho - Cc dng vn ng: Vt cht Nng lng Thng tin
MC TIU: 1. Nu c nguyn tc ca cc phng php nghin cu t bo. 2. Nu c nh ngha t bo. 3. Nu c cc hc thuyt t bo. 4. K c cc thuc tnh ca t bo. 5. Phn tch c mc tiu mn hc. 6. Hiu v k c cc vn ca sinh hc phn t. I. S LC LCH S - CC PHNG PHP NGHIN CU:
* T bo trong c th chng ta c kch thc nh gp 5 ln ngng nhn thy ca mt thng. Kch thc ca cc bo quan trong t bo cn nh hn nhiu ln, do , lch s nghin cu t bo gn lin vi s pht trin ca cc phng tin k thut cng cc ngnh vt l v ho hc. Ngng thy ca mt ngi: Hnh 1, Hnh 2

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* Knh hin vi quang hc (KHVQH) (Hnh: knh hin vi quang hc) c sng ch khong gia th k XVII v Robert Hook (1655) quan st c cc hc trng trong thn cy bc, ng gi l "khoang" (cell = khoang = t bo). Gia th k XIX, "Hc thuyt t bo " ra i (Schleiden v Schwann) vi lun im c bn "c th ng v thc vt c cu to bi cc t bo c nhn". Vo lc ny, cc nh khoa hc ch bit rng t bo gm c mng, nguyn sinh cht v nhn.

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* Knh hin vi in t (KHVT) ra i (gm KHVT xuyn (Hnh: vi nhung mao t bo rut non di knh hin vi in t xuyn) v KHVT qut (Hnh: Hng cu ngi hnh a lm hai mt di knh hin vi in t qut)) trong nhng nm 30 th k XX, cho php quan st cc bo quan nh bn trong t bo. Tuy nhin, c KHVQH v KHVT ch cho php nghin cu cc cu trc bt ng m thi. Trong thc t, cu trc sinh hc lun vn ng v ch c trong vn ng chng mi th hin y chc nng sinh hc ca mnh.

* My siu ly tm ra i (1926- Svedberg) cng cc phng php ho l khc nh sc k, in di, lc phn t ... cho php phn tch, tch chit cc thnh phn ring bit ca t bo. Nh c th nghin cu khi lng, kch thc, thnh phn ho hc ca chng v thc hin cc m hnh trn chng tr li vi nhau xc nh cc sn phm ca chng (h thng ti lp rp). Vic theo di, xc nh cc sn phm sinh hc c thc hin nh cc qu trnh nh du qua vic s dng cc cht ng v phng x v phng php phng x t chp hoc bng phng php ho m min dch.
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* Cui thp nin 70, phng php ti t hp ADN ra i, nh , ngi ta c th lp c "bn gen", gii m mt gen l nhanh chng hoc a mt gen mong mun vo t bo, sn xut ra cc sn phm ca gen vi mt khi lng ln ... Cc lnh vc ny to thnh ngnh di truyn, ngnh cng ngh gen phc v trong y hc, dc hc, nng nghip, mi sinh ...

* Nh vy, sinh hc pht trin c l nh nhiu phng php. Mi phng php c nhng u v khuyt im ring, nhng tt c chng cho php ta c mt ci nhn tng qut v s sng.

II. I TNG V MC TIU MN HC:

nh ngha: T bo l n v cu to v chc nng nh nht ca c th sng (Hnh: T bo v cc bo quan).

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Cc hc thuyt v t bo: * Tt c cc c th sng c cu to bi t bo v cc thnh phn ca n * Tt c cc t bo u ging nhau v mt cu trc ho hc * Cc t bo mi c thnh lp t s phn chia ca cc t bo c sn * Hot ng ca c th sng l tp hp cc hot ng v tng tc gia cc t bo ca c th. T bo c cc thuc tnh ch yu sau: * Tnh vn ng: s vn ng sinh hc bao gm c ngha c hc ln chuyn ho, thay i khng ngng cc cu trc; * Tnh sinh sn: l kh nng to ra nhng phin bn mi (copy) vi tnh cht tng t chnh mnh; * Tnh cm ng: l kh nng thu nhn thng tin t tng tc vi mi trng xung quanh v t mnh bin i, p ng vi iu kin ca mi trng; * Tnh phc hp: t bo c cu to t cc phn t polyme sinh hc, trong cc monome phn b khng tun hon trn chui polyme. * Nh vy, i tng ca t bo hc l nghin cu cc thuc tnh trn ca t bo, nhng l mt cng vic nghin cu tng hp tt c cc thuc tnh m khng phi l vic nghin cu tng thuc tnh ring r. Thng qua nghin cu, khoa hc v t bo tm hiu bn cht ca s sng, do n cung cp nn tng cho cc khoa hc v sc kho v bnh tt ca con ngi v mi trng. Mc tiu mn hc: 1. M t c t bo nh n v cu trc v chc nng c bn nht ca s sng; 2. M t c cc bo quan vi cu to v chc nng bnh thng ca chng; 3. Gii thch c mi quan h cht ch gia cu trc v chc nng ca cc bo quan, nu c s bin i ca cu trc hay chc nng s dn n cc ri lon ca c th; 4. Nm c nguyn tc - ngha ca cc phng php nghin cu v sn phm ca khoa hc v t bo trong iu kin c th c th ch nh hoc trin khai cc phng php v sn phm .

III. MT S VN CHUNG CA SINH HC T BO-PHN T:

3.1. S tin ha t bo t Procaryot n Eucaryot * Ton b sinh gii c hai loi t bo l Procaryot (t bo nhn s) v Eucaryot (t bo nhn hon chnh). Mt s tnh cht phn bit gia hai loi t bo ny nh sau: So snh t bo Procaryot v Eucaryot
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c im Mng nhn Nhim sc th S lng Cu trc Histon, nucleosom Trng thi ca a s gen Tng hp ARN v protein Cc bo quan c mng (Ti th, Li ni Bo, Golgi, Tiu th...) Trung th, thoi v sc B xng t bo Ribosom

Procaryot t Vng M ng thi 70S

Eucaryot + Nhiu Tuyn tnh + ng Tch ri + + + 80S

* T Procaryot n Eucaryot l mt bc tin ha rt di ca sinh gii, trong vn then cht nht l cht liu thng tin di truyn. Cc cu trc mi nh mng nhn, thoi v sc, nhim sc th tuyn tnh c s ngng t ADN-protein vv... u nhm chung mc ch l gip cho t bo Eucaryot duy tr c mt b my thng tin di truyn khng l so vi Procaryot. Theo thuyt tin ha, t bo Eucaryot c hnh thnh t nhng t bo tng t Procaryot. t bo Procaryot, ta thy mt s on mng bo tng lm vo t bo cht, nh nhng cu trc s khai ca h thng cc mng ni bo. Cn t bo Eucaryot, cc phn t ADN hnh vng vn cn tn ti trong cc bo quan ln nh ty th v lp th c hai lp mng bao quanh. Cc bo quan ny cng cha ribosom 70S v h thng tng hp protein kiu Procaryot. Mt gi thuyt c a ra rng trong qu kh, xy ra s cng sinh gia t bo thy t ca Eucaryot vi cc t bo Procaryot. S cng sinh ny ngy cng cht ch n mc cc thnh phn Procaryot tr thnh bo quan ca t bo kiu mi - Eucaryot, nhng mi bn vn c b my thng tin di truyn tng i c lp. * Trong chng trnh hc, chng ta s i su nghin cu cu trc v chc nng ca cc bo quan v bn thn t bo Eucaryot. 3. 2. Vt cht - Nng lng v Thng tin T bo c ba dng vn ng ln l vt cht, nng lng v thng tin. Ni cho chnh xc hn, th qu trnh sng c th c nghin cu mt cch tng i ring r trn ba bnh din ni trn. y cng l thuc tnh ca bt k cu trc v chc nng sinh hc no. a. Vt cht hay vt liu xy dng: bao gm cc phn t nh v cc i phn t. Chng ta s phn tch k hn thnh phn cu to ha hc ca cc t bo trong bi tip theo. y ch gii thiu vn tt cc nhm phn t vt cht ch yu ca t bo. * ng (sugars). Cc phn t ng hiu theo ngha rng bao gm cc loi ng n (monosaccharides) v phn t bao gm t hai gc ng tr ln (di-, oligo- v polysaccharides). Cc phn t ng ng vai tr cung cp nhin liu (nng lng) cho t bo, nhng cng tham gia vo cu trc i phn t AND, ARN, cu trc mng, cc glycoprotein th th, enzym v.v...
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* Lipid. Lipid (phospholipid, cholestrrol, glycolipid...) l thnh phn quan trng nht ca cc mng sinh cht (mng bo tng v mng ni bo). Ngoi ra, lipid c th tn ti di dng cc ht nh phn tn trong bo tng v trong mu, v tham gia vo vic cung cp nng lng cho t bo. * Acid amin v protein. C 20 loi acid amin c bn cu to nn cc i phn t protein. * Nucleotid v acid nucleic. Nucleotid bao gm gc baz nit, gn vi gc ng riboz hay deoxyriboz, cc gc ng ny c phosphoryl ha (gn gc phosphat). Khc vi acid amin trong protein, nucleotid trong ADN hay ARN ch gm 4 loi (A, T, G, X hay A, U, G, X). * Nc v cht in gii v c. Phn t nc c tnh cht rt c bit. Chng l cc lng cc, c th to ra lin kt hydro vi nhau. Cc lin kt ny mc d ch mang tnh tm thi v lun lun ph v v ri li hnh thnh li, nhng nh s lng rt ln, chng lm cho nc trong t bo duy tr trng thi tinh th lng. Tinh th lng ny c cu trc kh xp, vi nhiu khong trng. Kch thc cc khong trng ny kh ph hp vi kch thc chui xon alpha ca protein. Nh vy, ch c trong mi trng nc cc phn t protein mi c n nh v thc hin cc chc nng sinh hc ca chng. Trn b mt ca cc i phn t, cc cu trc mng v bo quan khc, cc phn t nc lng cc c phn cch kh cht. Lp nc ny, thng qua cc lin kt hydro, li nh hng n cu trc lng ca mi trng nc xa hn, cng nh ti cc cu trc i phn t, mng v bo quan khc. Mc d cc tng tc ny rt yu, nhng nh s lng nhiu, chng c th ng vai tr quan trng trong cc p ng ca t bo nh mt th thng nht. b. Nng lng. Mi phn ng sinh ha v qu trnh sinh hc mc t bo, cn t bo u km theo tiu tn nng lng t do v dn n xu hng ta nhit, ph hy cc trt t vt cht trong t bo. Do vy t bo lun lun cn c b sung nng lng. Nng lng ny c cung cp di dng ha nng ca adenozin triphosphat (ATP) l phn t nh c tng hp ch yu trong ty th. ng v m c t bo nhp khu t mu v dch ngoi bo, chng c ngun gc t thc n, hoc t cc m d tr (gan, c, m m). c. Thng tin. Cc cu trc cng nh qu trnh sinh hc trong t bo thng tun theo mt trt t khng gian v thi gian cht ch. Trt t ny phn nh c tnh thng tin ca h thng sng. "Kho tng thng tin" chnh ca t bo nm trong nhn, "bn gc" ca thng tin lun c ct gi trong nhn, ch c cc "bn sao" (mARN c chp ra nhiu bn) l c gi ra bo tng iu khin cc qu trnh sinh hc ca t bo. Thng tin t cc bn sao ny c nhn tip ln nhiu ln na di dng sp xp cc acid amin trn chui polypeptid ca protein. Cc protein c th thc hin rt nhiu chc nng khc nhau. Khc vi bn gc thng tin (AND), cc bn sao v bn dch thng tin (ARN v protein) ch tn ti trong mt thi gian gii hn. duy tr hot ng t bo, mt dng thng tin phi c duy tr thng xuyn t nhn ra bo tng. iu ny c phn nh trong "Hc thuyt trung tm" ca Sinh hc phn t nh sau: ADN ---> ARN ---> Protein * T bo cn thu nhn thng tin t mi trng xung quanh (v d cc hormon, cc dc phm...) v qua iu chnh hot ng ca mnh. Thng tin ny khng tn ti di cc dng vn bn quy nh cu trc ca i phn t, m l cc tn hiu tc ng chn lc ln cc th th c hiu. Cc th th ny c th thng qua h thng cc cht truyn tin th hai (c-ATP, proteinkinaza...) tc ng tr li cc thnh phm ca t bo, k c ti nhn. Nhn p ng cc thng tin ngc dng ny khng phi bng cch thay i cu trc gen ca mnh, m bng cch ng hay m cc gen khc nhau.
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C ba dng vn ng vt cht, nng lng v thng tin c lin quan mt thit vi nhau.

IV. TM TT:
T bo l n v cu to v chc nng c bn ca c th sng. Vic nghin cu t bo thc hin da trn s pht trin ca ca cc khoa hc khc. S pht trin ca khoa hc v t bo li phc v cho cc khoa hc v y t, nng nghip, mi sinh ... Cc vn thng c t ra trong nghin cu l: (1) Qu trnh tin ho t Prokaryote n Eukaryote; (2) S vn ng ca ba dng vt cht, nng lng v thng tin i tng nghin cu ca khoa hc v t bo l 4 thuc tnh ca n: vn ng, sinh sn, cm ng v s phc hp trong cu to. Mc tiu chnh ca mn hc l nm vng cu trc v chc nng ca tng bo quan qua nu bt mi lin h cht ch gia cu trc v chc nng. T kho: Knh hin vi quang hc - Knh hin vi in t (xuyn, qut) - Hc thuyt t bo Phng php phn tch v ti lp rp - Phng php phng x t chp - Phng php ho m min dch - Phng php ti t hp AND - Tnh vn ng - Tnh sinh sn - Tnh cm ng - Tnh phc tp - Prokaryote - Eucaryote - Vt cht - Nng lng - Thng tin Ti liu tham kho:

1. James D.Watson: Molecular Biology of The Cell, 1995, third edition, Garland Publishing Inc., p.477-506 2. Baudhuin: Cytologie,1997, Universit Catholique de Louvain 3. W. E. Ganong: Review of Medical Physiology, thirteen edition, Lange medical book, 1991, p.10-26

MNG SINH CHT

1. NH NGHA 2. LIPID CA MNG 2.1. CU TO: PHOSPHOLIPID, CHOLESTEROL, GLYCOLIPID 2.2. TNH CHT: TNH T KHP KN TNH LNG 3. PROTEIN CA MNG: 3.1. PROTEIN XUYN MNG 3.2. PROTEIN CN MNG Ebook created by CLB195 Bi ging ca cc BS : Nguyn Quc Dng _ Trn Cng Toi_L Quc S

4. GLUCID CA MNG 4.1. GLYCOPROTEIN 4.2. PROTEOGLYCAN 5. CC TNH CHT CA MNG TNH T KHP KN, TNH LNG, TNH BT I XNG

MC TIU: 1. Thuc v phn tch c nh ngha 2. K tn cc loi lipid ca mng v phn tch c im cu trc ca chng 3. K tn cc loi protein ca mng v phn tch c im cu trc ca chng 4. K tn cc loi glucid ca mng v phn tch c im cu trc ca chng 5. Gii thch c hai tnh cht ca mng 6. Gii thch c cu trc bt i xng ca mng I. NH NGHA: (Hnh: S hnh nh 3 chiu ca mng t bo)

* Mng v cng quan trng i vi s sng ca t bo. Mng bo tng bao bc t bo, xc nh gii hn ca n v duy tr s khc bit r rt gia mi trng bn trong t bo (cn gi l bo tng) v mi trng bn ngoi t bo (cn gi l mi trng ngoi bo). Mng ni bo li ni sinh cht, Golgi, ty th v cc bo quan c mng khc ca t bo Eukaryote duy tr s khc bit gia mi trng bn trong mi bo quan v dch bo tng. * nh ngha: D c nhiu chc nng khc nhau, mi mng sinh cht u c mt cu trc chung: chng u gm mt lp cc mng cc phn t lipid v protein gn vi nhau bng cc lin kt phi ho tr cng mt t glucid mt ngoi t bo. Cc mng t bo u l nhng cu trc ng v lng, trong phn ln cc phn t c th di ng khp b mt ca mng. Hnh: Mng sinh cht vi cc phn t lipid, protein v glucid

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Hnh: Mng sinh cht di knh hin vi in t xuyn

II. LIPID CA MNG:

* Lipid to thnh cu trc c bn ca mng v quyt nh nhng tnh cht quan trng nh tnh t khp kn, tnh lng v tnh bt i xng ca mng. C th ni, lipid l thnh phn ch yu thc hin chc nng v cu trc ca mng. * Lipid mng bao gm ba loi chnh: phospholipid, cholesterol v glycolipid. * Cc phn t lipid mng c c im lng tnh, cha c hai thnh phn a nc v k nc. Chng sp xp thnh hai lp, quay ui k nc vo nhau, hng u a nc v hai pha b mt ca mng. Hnh: S sp xp cc phn t lipid , hnh 3 chiu

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2.1. Phospholipid * Phn t phospholipid gm ba thnh phn: (1) Gc amin v gc acid phosphoric c th in ly to thnh ion dng v m (thnh phn "a nc"); (2) Hai gc acid bo khng tan trong nc (thnh phn "k nc"); (3) Gc glycrin lin kt thnh phn a nc v k nc. Hnh: Phn t phospholipid * Mng t bo Eukaryot cha nhiu loi phospholipid khc nhau, bao gm 4 loi chnh, chng khc nhau ch yu gc amin: + Phosphatidyl-cholin + Phosphatidyl-ethanolamin

Hnh: S phn t Phosphatidyl-ethanolamin + Phosphatidyl-serin + Sphingomyelin (gc amin cng l cholin) * Cc gc amin u tch in dng, in tch ny c trung ho bi in tch m ca gc phosphat. Ring serin c cha thm mt in tch m, do phn t phosphatidyl-serin khng trung ho v in m li tch in m.

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2.2. Cholesterol * Ch c mng t bo Eukaryot, t l c khi chim ti trn 40% lipid ca mng (tng ng phospholipid). * Cholesterol c kch thc nh hn phospholipid, thnh phn a nc ch mc phn cc m khng phn ly, do vy cholesterol d dng hn trong vic chuyn ng i ch gia hai lp lipid. Hnh: Phn t cholesterol

2.3. Glycolipid * Ch chim t l vi phn trm trong thnh phn lipid ca mng, c nhiu loi t bo khc nhau, c bit t bo thn kinh. 2.4. Tnh t khp kn ca mng * Mng c cu trc lin tc, khi b phn on th cc b ra lun lun c xu hng t khp li vi nhau. Tnh cht ny l do thnh phn lipid quyt nh. * Do c cha thnh phn k nc nn mi phn t lipid ring bit khng th tan trong mi trng nc, nhng khi c nhiu phn t lipid th chng c th lin kt vi nhau thnh cc cu trc c th tan trong nc: l micelle v liposom.
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* Trong cu trc micelle, cc ui k nc lin kt vi nhau thnh mt khi, bao bc bn ngoi l cc u a nc. Nh c b mt phn cc, ht micelle phn tn c trong mi trng nc nh nhng git m nh li ti. y chnh l kiu cu trc ca cc ht lipoprotein vn chuyn trong mu. * Cu trc liposom to ra cc nang, bn trong cha mi trng nc, ton b nang ny cng tn ti bn vng trong mi trng nc. Nang c bao bc bi mt lp mng lipid gm hai lp phn t, sao cho b mt ca mng c t hai pha u c che ph bi cc u a nc. y chnh l m hnh cu to mng lipid kp ca t bo. ngha sinh hc: * Nh tnh t khp kn m mng lun c xc nh ranh gii r rng vi mi trng; * Mng c th thc hin c cc qu trnh thc bo v xut bo; * Mng phi c tng hp trn c s mng sn c. 2.5. Tnh lng * Tnh cht lng cng do thnh phn lipid quyt nh. Mc d khi lin kt vi nhau, cc chui k nc c xu hng hnh thnh mt cu trc tinh th, nhng l mt "tinh th lng". Tnh cht th hin trc ht qua chuyn ng ca cc phn t lipid: * Chuyn ng i ch cho phn t lipid bn cnh, cng lp. chuyn ng ny c tn sut khong 1/107 giy. * Chuyn ch sang lp i din (flip-flop): mun thc hin iu ny, khng nhng phn t phi quay ui k nc 1800 m cn phi a phn a nc chuyn ng xuyn qua lp "m" (acid bo). V vy, tn sut chuyn ng ny rt thp (1 ln/thng). Mng t bo cha enzym c bit xc tc cho chuyn ng ny. * Chuyn ng quay quanh trc * Tnh lng ph thuc vo: * Nhit : nhit tng th tnh lng tng theo

Hnh: S minh ho tnh lng ph thuc vo nhit

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* Thnh phn ho hc ca cc lipid: ui k nc ngn th tnh lng tng, t l cholesterol tng th tnh lng gim. ngha sinh hc: * Nh c tnh lng, mng bo tng c tnh mm do, n hi v bn vng, n c th bin dng, gp np trong cc chuyn ng (chuyn ng gi tc). * C th t tng hp v thc hin cc qu trnh hp mng nh nhp bo, xut bo ... * Nhiu qu trnh enzym: din ra trn b mt mng vi hot tnh cao nhng vi trt t nht nh. Hnh: Chuyn ng gi tc trong qu trnh thc bo: ton cnh, th gi tc, bt mi, a vo TB

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III. PROTEIN MNG:

* T l v mt khi lng gia protein/lipid mng khong 1/1, nhng protein c phn t ln hn lipid nhiu, nn xt v mt s lng th trung bnh trn mng c 50 phn t lipid mi c 1 phn t protein. Do , c th v protein nh nhng "hn o" trn "i dng"cht bo m cc "hn o" ny c kch thc v gm nhiu loi khc nhau. Chng c th c phn thnh hai loi chnh: protein xuyn mng v protein cn mng.

Hnh: S phn b ca cc "hn o" prtin trn "i dng" cht bo

3.1. Protein xuyn mng * Protein xuyn mng bao gm thnh phn k nc v thnh phn a nc. * Thnh phn k nc khng phn cc nn c th nm chm trong lp lipid mng, v 1 protein c th cha mt hay nhiu on k nc nn c th "xuyn mng" mt hay nhiu ln. * Thnh phn a nc th phn cc v hay tch in, do c th "ni" trn b mt mng (trong hoc ngoi). Hnh: Protein xuyn mng

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3.2. Protein cn mng * Protein cn mng khng c thnh phn k nc nn ch nm trn b mt, n c th gn vi mng thng qua: - Lin kt phi ho tr vi protein xuyn mng, trng hp ny n c th d dng tch khi mng nn cn c gi l protein ngoi mng. - Lin kt ho tr vi mt phn t lipid (proteolipid), do c xp vo loi protein cu trc mng ging nh protein xuyn mng.

IV. NG CA MNG:
* Chim khong 2-10% khi lng mng v c tnh cht phn cc nn khng th nm chm trong lp lipid m ch c trn b mt v gn vi mng thng qua lin kt ho tr vi lipid mng. * Hp cht ng ca mng gm hai loi chnh: - Glycoprotein - Proteoglycan thc ra cng l mt loi glycoprotein c bit, n c t bo tit ra mi trng xung quanh to thnh cht nn quanh cc t bo lin kt (v d t bo sn).

V. CC TNH CHT CA MNG:

5.1. Tnh lng 5.2. Tnh t khp kn 5.3. Tnh bt i xng: * Tnh bt i xng ca mng th hin c v cu trc ln chc nng. Xt v mt cu trc, tnh bt i xng ca mng c lin quan c ba thnh phn lipid, protein v ng. * Lp trong v ngoi mng c cc thnh phn lipid khc nhau. mt ngoi, c nhiu phostatidyl-cholin v sphingomyelin cha t l acid bo bo ho cao nn lp ngoi t lng hn so vi lp trong. Mt trong li cha nhiu phosphatidyl-serin tch in m nn in tch trong mng m hn so vi ngoi. * ng ch tp trung ngoi mng v to thnh lp o t bo (glycocalix hay cell coat) * Protein xuyn mng mt hoc nhiu ln to cho mng cu trc bt i xng ng thi c th to thnh knh cho cc phn t nh a nc lt qua mng hoc hnh thnh nhng tm hot ng enzym cho mng. Protein cn mng gm cc protein hoc lun nm pha bo tng hoc lun nm hng v pha ngoi bo. T kho: Mng bo tng - Mng ni bo - Bo tng - Ngoi bo - Phospholipid Cholesterol - Glycolipid - Protein xuyn mng - Protein cn mng - Glycoprotein Proteoglycan - Tnh lng - Tnh lin tc - Tnh bt i xng - Micelle - Liposom Ti liu tham kho: 1. James D.Watson: Molecular Biology of The Cell, 1995, third edition, Garland Publishing Inc. 2. Baudhuin: Cytologie, 1997, Universit Catholique de Louvain.
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3. W. E. Ganong: Review of Medical Physiology, thirteen edition, Lange medical book, 1991

VN CHUYN VT CHT QUA MNG

1. i cng Gradient nng v vn chuyn th ng - tch cc n vn v hip vn: ng vn v i vn Vn chuyn phn t cht tan v khi ln 2. Vn chuyn phn t cht tan: Khuch tn n gin Protein trung gian vn chuyn qua mng Protein ti: Th ng Tch cc: bm Na+/K+, bm H+, gradient Na Tn thng do ri lon VC tch cc Protein knh So snh protein ti v knh 3. Vn chuyn phn t khi ln: Xut bo: lin tc hoc iu khin Nhp bo: m bo Nhp bo qua trung gian th th Thc bo I. I CNG:
* Cc mng sinh cht ang hot ng u c thuc tnh l gy ra mt s chnh lch v nng ca cc cht ho tan gia hai pha ca mng. y l iu kin cn thit cho vic thc hin mi chc nng ca t bo sng v l kt qu cn bng ng gia cc qu trnh trao i, vn chuyn vt cht (VCVC) lin tc qua mng. Chnh lch nng c th c trng bng mt vect gi l gradient nng v c hng i t pha nng cao n pha c nng thp. Gradient nng to ra mt th nng. Vt cht c xu hng t nhin tr v th nng ti thiu, ngha l cc cht ha tan c xu hng vn ng xui chiu vect gradient nng v trit tiu gradient.
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Mng sinh cht (MSC) ch c th VCVC ngc chiu gradient v lm tng tr s cc gradient nng qua mng khi c cung cp nng lng. Nng lng t do do ATP cung cp c chuyn thnh th nng ca gradient nng vt cht qua mng. Th nng ny cng l mt dng nng lng t do v c th c t bo s dng trong vic thc hin cc cng khc nhau. V vy, VCVC qua mng c th c phn bit thnh vn chuyn tch cc (ngc chiu gradient, cn cung cp nng lng) v vn chuyn th ng (xui chiu gradient, khng cn cung cp thm nng lng).

* VCVC qua mng cn c phn bit thnh n vn hay hip vn. n vn l vn chuyn mt cht qua mng mt cch c lp, khng km vn chuyn cht khc. Hip vn l hin tng hai phn t khc nhau c ng thi qua cng mt v tr trn mng, trong c mt cht c vn chuyn ngc chiu gradient, nng lng cn thit cho qu trnh ny c cung cp t s vn chuyn ca cht th hai xui chiu gradient. Nh vy, hip vn cng l mt dng c bit ca vn chuyn tch cc nhng s dng nng lng khng do ATP cung cp. Hip vn phn bit thnh ng vn v i vn. Nu hai phn t vn chuyn cng chiu (u t trong ra ngoi hoc t ngoi vo trong), ta c hin tng ng vn. Khi hai phn t c vn chuyn theo kiu trao i, mt t trong ra, mt t ngoi vo, l i vn.

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* Vt cht c th qua mng di dng phn t hay ion ho tan, nhng cng c th di dng ht khng tan hoc khi kh ln. VCVC khi ln chia thnh nhp bo v xut bo (nut vo hay tng ra). Nhp bo chia thnh m bo (ung, nhp bo khng c hiu) v nhp bo qua th th. Mt trng hp c bit ca nhp bo qua th th l hin tng thc bo (n). Vn chuyn khi ln thc hin qua c ch hp mng, v c coi l vn chuyn tch cc v km theo tiu tn nng lng do ATP cung cp. * Tm tt phn loi s vn chuyn vt cht qua mng: 1. Tiu th nng lng t do hay khng? - C 1.1. Vn chuyn tch cc - Khng 1.2. Vn chuyn th ng, khuch tn: 1.2.1.- Khuch tn n gin 1.2.2.- Khuch tn qua protein trung gian 2. Vn chuyn c lp hay kt hp hai cht? - c lp 2.1.n vn - Kt hp 2.2.Hip vn: 2.2.1.- ng vn 2.2.2.- i vn 3. Vn chuyn tng phn t (ion) hay khi nhiu phn t? 3.1.- Vn chuyn phn t 3.2.- Vn chuyn khi ln: 3.2.1.- Xut bo 3.2.2.- Nhp bo: 3.2.2.1.- m bo 3.2.2.2.- Nhp bo th th 3.2.2.3.- Thc bo

II. VN CHUYN PHN T CHT TAN:

1. Khuch tn n gin: Phn t nh ho tan c th khuch tn n gin qua mng lipid kp t pha nng cao sang nng thp. Tc khuch tn ph thuc vo ho tan ca phn t trong mng lipid kp v vo gradient nng gia hai bn mng (n cn ph thuc vo din tch khuch tn v on ng i). ho tan trong mng lipid kp ph thuc vo mc k nc. Ion v phn t phn cc c mc a nc cao, do khuch tn qua mng kh khn hn so vi cc cht khng phn cc. S khuch tn t do
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2. Protein trung gian vn chuyn qua mng: Phn t phn cc v ion khng tan trong lipid nn khng th lt qua mng bng khuch tn n gin. Chng ch c th qua mng nh protein trung gian trong mng.

y l cc protein xuyn mng v c c hiu nht nh vi cht m chng vn chuyn. iu ny c ngha l mi loi protein ch vn chuyn mt loi phn t hay ion no . C hai kiu hot ng ca protein trung gian vn chuyn:

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- Protein ti (carrier, transporter): l protein xuyn mng c tm gn phn t cn vn chuyn v sau khi gn, chng thay i cu hnh khng gian sao cho tm gn ny c "m" v pha bn kia ca mng.

- Protein knh (chanel): to thnh mt l hng xuyn qua mng lm phn t phn cc hay ion c th i qua m khng cn phi tip xc vi lipid.

Nh protein trung gian vn chuyn nn tc khuch tn xui chiu gradient c th tng vt nhiu ln. Tuy nhin, khi nng cht cn vn chuyn qu ln so vi s lng protein trung gian, ta c hin tng bo ho, ngha l tc khng th tng thm hn na.

a. Protein ti: - Trong vn chuyn th ng: V d qu trnh vn chuyn v trao i ion HCO3- v H+ hng cu: Trong mao mch cc m, nng CO2 cao, cht ny khuch tn t do qua mng hng cu. Trong hng cu, n kt hp vi nc to ra HCO3- v H+ (di tc dng ca enzym carbonic anhydraza). H+ lin kt vi Hemoglobin (Hb), cn HCO3Ebook created by CLB195 Bi ging ca cc BS : Nguyn Quc Dng _ Trn Cng Toi_L Quc S

 mao mch phi, HCO3- c vn chuyn qua protein ti vo hng cu v di tc dng ca enzym carbonic anhydraza (theo chiu ngc li, v lc ny nng CO2 gim thp) ion ny kt hp tr li vi H+ v to thnh CO2 + H2 O. Nng CO2 trong huyt tng thp, do CO2 khuch tn qua mng ra khi hng cu. Nng HCO3- trong hng cu tip tc h xung, ion ny li c protein ti vn chuyn t huyt tng vo hng cu. H+ th c cung cp nh s phn ly khi Hb.O2. Nh vy, qu trnh thi kh carbonic khi m l nh khuch tn (vn chuyn th ng) t ni c nng cao n ni thp hn, nhng nh enzym xc tc v protein ti m din ra vi hiu qu cao. - Trong vn chuyn tch cc: Protein ti tham gia vo vn chuyn th ng khi c cung cp nng lng t do cng c th vn chuyn c cht tan ngc chiu gradient nng . Nng lng c th ly t phn ng thy phn ATP, hoc t mt gradient khc. Trng hp vn chuyn ngc chiu gradient bng nng lng gii phng ra do phn t khc vn chuyn xui chiu gradient c gi l hip vn. Tuy nhin a s c cung cp nng lng t do bi phn ng thy phn ATP, nhiu protein ti ng thi cng c hot tnh ATPaza. Khi c cung cp ATP v vn chuyn cc cht ngc chiu gradient nng , protein ti c gi l bm ATPaza: * Bm Na/K - ATPaza:

Nng Na+ ni bo thp hn nhiu so vi ngoi bo, cn K+ ni bo cao hn ngoi bo (10-20 ln). S chnh lch ny c duy tr thng xuyn bi phc hp protein gi l bm Na+/K+ - ATPaza nm trn mng bo tng. y l protein xuyn mng c tm gn K+ pha ngoi bo v cc tm gn Na+ pha ni bo , tm ATPaza pha ni bo. Mi ATP b phn hy bm c 3 Na+ ra v 2 K+ vo. Qu trnh bm Na+/ K+ ngc chiu gradient nng v thy phn ATP lun lun song hnh vi nhau, v c th b c ch bi ouabain khi ho cht ny c mt dch ngoi bo.

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* Bm H+ - ATPaza: y l protein ti vn chuyn tch cc H+ qua mng km theo thy phn ATP. Bm H+ tham gia vo vic duy tr mi trng acid trong cc tiu th nh vn chuyn lin tc H+ t dch bo tng qua mng tiu th. N cng c mt trn mng bo tng v vn chuyn H+ t bo tng ra khoang gian bo, nh vy, d t bo trong khi trao i cht lun sinh ra acid (CO2, acid lactic...) nhng vn duy tr c pH trung tnh. Mng t bo d dy tit acid chlohydric c bit c bm H+ rt hot ng. * Gradient Na+ cung cp nng lng cho vn chuyn cht khc: Trong t bo, nng Na+ thp hn nhiu so vi ngoi t bo. Gradient Na+ (gradient in ho) nh mt ngun th nng vn chuyn mt s phn t v ion khc nhau nh hip vn: Na+ xui chiu gradient (t ngoi vo bo tng) km theo vn chuyn mt cht khc ngc chiu gradient ca cht . Chng hn glucoza t khoang rut c hp thu vo t bo nh c ch ng vn vi Na+, trong c Na+ v glucoza c vn chuyn t ngoi vo trong qua mt protein ti. Trng hp t bo c tim vn chuyn Ca++ ngc chiu gradient t bo tng ra gian bo v vo li ni c tng da trn i vn vi Na+ chy xui chiu gradient: Ca++ v Na+ chy ngc chiu nhau qua cng mt protein ti nm trn mng bo tng hoc mng li ni c tng. - Tn thng t bo do ri lon vn chuyn tch cc: T bo sng lun vn chuyn tch cc duy tr cc gradient nng qua mng. Trong iu kin thiu nng lng, chng hn thiu oxy, ty th khng sn xut y ATP, bm ATPaza ngng hot ng dn n tn thng v cht t bo: - Bm Na+/K+ ngng hot ng, Na+ xui chiu gradient lt vo t bo, in tch m gim i, mng bo tng b kh cc, Cl- d dng lt vo t bo. Nng mui bo tng tng ln, lm tng p lc thm thu, ht nc t ngoi vo, t bo trng n, thm ch c th v.
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- Bm H+ ngng hot ng lm cho bo tng b axit ho (pH gim) trong khi pH tiu th tng ln. Enzym trong tiu th c th lt ra bo tng gy ph hy t bo v.v.... Cc t bo tiu th nhiu ATP nh c tim v no li cng nhy cm vi cc ri lon thiu oxy. Thiu oxy nng (vi chc pht) c th gy ra nhng hoi t trong c tim hay no (nhi mu). b. Protein knh: Protein knh l protein xuyn mng lm nhim v trung gian vn chuyn ging protein ti. Tuy nhin, c nhng im khc bit so vi protein ti l: - Ch c protein knh vn chuyn ion, khng c knh cho phn t trung ho. V vy, chng thng c gi l knh ion. - Knh ion ch vn chuyn th ng m khng th va th ng, va tch cc nh protein ti. - Tc vn chuyn qua knh ln hn nhiu so vi protein ti (mi giy mt knh c th cho trn 106 ion i qua!) - Knh ion thng khng m lin tc, m c c ch m v ng knh. Bnh thng trng thi ng, khi c tc nhn tc dng ln mng, knh m ra tm thi ri ng li ngay. Tc nhn m knh c th l in th mng, kch thch c hc, hoc ho hc nh cht trung gian dn truyn thn kinh, nucleotid, protein (protein-G), hay mt ion khc. Knh ion ng vai tr quan trng trong qu trnh kh cc mng t bo, dn truyn xung ng thn kinh, thng tin, iu khin lin t bo.

III. VN CHUYN KHI LN:

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* Cc i phn t nh protein, polysaccharid, polynucleotid, hoc cc ht ln nh vi sinh vt, khng qua mng c bng khuch tn n gin hay protein trung gian vn chuyn. Qu trnh vn chuyn t bo tng ra ngoi bo gi l xut bo, cn ngc li gi l nhp bo. Trong c hai trng hp, cht c vn chuyn bao gm mt khi kch thc ln, c bao bc bi mng lipid kp. S vn chuyn i hi mt c ch gi l kt hp mng, bao gm giai on dnh mng v giai on ha nhp mng (membrane fusion = adherence + joining)

xy ra gia mng bao quanh khi c vn chuyn vi mng bo tng. Trong qu trnh trn, tnh lin tc v bt i xng ca mng lun c duy tr. Khi vt cht khng bao gi ha nhp trc tip vo bo tng m lun c ngn cch qua mng. Lp lipid hng v pha khi vt cht c vn chuyn lun tng ng vi lp ngoi (lp hng v pha ngoi bo). 1. Xut bo: Xut bo l qu trnh vn chuyn khi vt cht c ngn cch vi dch bo tng t trc bi mt lp mng ni bo, ra khoang gian bo. i tng xut bo gm cc ti ch tit do Golgi hnh thnh v th cn b to ra t tiu th th cp. Trc ht, xy ra s dnh gia hai lp lipid hng v dch bo tng. Sau , mng ca ti ha nhp vi mng bo tng, nh khong khng bn trong ti c m thng vi khong gian bo. Xut bo c hai kiu: (1) Ch tit lin tc thy mi t bo, trong ti c chuyn ra mng v xut bo ngay; (2) Mt s t bo c c ch ch tit c iu khin, trong cc nang kt hp mng vi nhau to thnh ti d tr c kch thc ln hn. Ti chuyn ng v pha mng, nhng ch xy ra kt hp mng khi c tn hiu iu khin tc ng ln mng. Tn hiu iu khin thng tc dng thng qua knh ion Ca++. Ion ny c th t ngoi bo lt vo bo tng, hoc c gii phng t nhng cu trc d tr Ca++ trong t bo. Nng Ca++ tng t ngt nh knh ion to thnh tn hiu kt hp mng, gy xut bo cc cht cha bn trong ti ch tit (hormon, enzym tiu ho, cht trung gian dn truyn thn kinh v.v...). Tn hiu iu khin c th ch tc dng ln mt khu vc hn ch ca mng bo tng, v phn ng ch tit cng c th ch xy ra trn khu vc ny ca mng.
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S lng mng ni bo c nhp vo mng bo tng do c ch xut bo c th rt cao. V d mi t bo cc ngn ca ty c din tch mng nh l 30 micromt 2, nhng khi c tn hiu ch tit enzym tiu ho, mng phi tip nhn thm n 900 micromt2 mng ca cc ti ch tit. Mng ny sau c thu hi vo h thng mng ni bo nh nhp bo. 2. Nhp bo: Nhp bo l qu trnh vn chuyn t gian bo vo bo tng, trong khi vt cht sau khi vo bo tng vn c ngn cch bng mt lp mng: khi vt cht c cch li nh s dnh mng gia hai lp lipid pha ngoi bo, sau c chuyn hn vo bo tng. C hai kiu nhp bo: m bo v thc bo. m bo c th thy hu ht cc t bo, trong khi thc bo ch xy ra mt s loi t bo. a. m bo: m bo l s nut vo mt cch khng c hiu v thng xuyn cc cht dch v phn t ho tan t dch ngoi bo vo bo tng. Mng bo tng lm xung thnh mt cu trc gi l lm mc o (coated pit), sau bt vo bn trong nh kt hp mng, to thnh nang mc o (coated vesicle). Lm v nang mc o c kch thc chng 150 nm. Pha di mng c mt lp li protein clathrin. Chnh li ny to ra lc ko mng bo tng lm xung v xy ra kt hp mng.

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Mt phc hp clathrin gm 3 chui polypeptid gn vi nhau thnh mt hnh qut 3 cnh, gi l triskelion. Nhiu triskelion lin kt nhau to thnh mng li hnh cu lt bn di mng bo tng, ko mng lm xung ri bt ra thnh nang mc o. Lc ko do qu trnh polyme ho cc triskelion vi nhau to ra. Nhng khi nang mc o tch khi mng, li xy ra s phn ly phc hp clathrin, to thnh nang trn (uncoated vesicle). Nang ny gi l nang nhp bo (endosom), c vn chuyn hng n v nhp vo tiu th tiu ho.

Lm mc o ch tn ti khong mt pht, cn nang mc o ch trong vi giy. m bo l cch m t bo lin tc hp thu vt cht t dch ngoi bo. Mi pht, mt nguyn bo si nui cy c th nut vo n 2500 nang. Nh vy, mng ngoi bo b lin tc chuyn thnh mng ni bo v c mt qu trnh ngc li (xut bo) cn bng.
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b. Nhp bo qua trung gian th th: Lp lipid ngoi ca mng bo tng thng cha nhiu protein-th th. T bo c th cha hng chc loi th th khc nhau trn mng. Do tnh lng ca mng, cc th th di ng c trn mng, c th trong trng thi gn vi phi t (ligand) ca chng, hoc l t do cha gn. Nh li protein clathrin, mng thng xuyn hnh thnh cc lm v ti mc o hp thu vt cht ngoi bo. Cc lm ny cng bao gm lun c th th cng vi phi t ca chng (mi lm c th cha khong 1000 th th). Nh th th m ngoi thnh phn c nut vo mt cch khng c hiu (m bo), t bo cng hp thu lun cc cht tp trung trn b mt. Bng cch ny, nhiu cht trong ngoi bo ch vi nng rt thp nhng t bo vn c th tp trung trong bo tng vi nng cao hn bn ngoi hng ngn ln. Sau khi lm mc o hnh thnh ti mc o, mng li clathrin b phn ly, cn li nang trn (endosom). Trong endosom, cc phi t phn ly khi th th, c hp thu mt phn vo bo tng, hoc nm li ti cc tiu th th cp. Cc th th c th tp trung vo mt khu vc ring bit ca mng endosom, vng ny sau tch ra thnh nang vn chuyn n nhp mng vi mng bo tng. Nh th th v mng ni bo li c chuyn ra mng ti s dng. Endosom dch chuyn dn vo pha trong bo tng, ng thi cht dch bn trong dn dn b axit ho do cc bm H+ trn mng. Cui cng, endosom nhp vi tiu th s cp, hnh thnh tiu th th cp. Bn trong tiu th th cp, cc vt cht nhp bo s b cc enzym "tiu ho" v hp thu vo dch bo tng.

V d: Hp thu LDL-cholesterol (LDL = low-density lipoprotein = lipoprotein t trng thp).nh nhp bo qua trung gian th th. Cholesterol l vt cht m t bo cn hp thu xy dng mng. Chng c c tng hp trong gan v vo mu di dng ht kch thc khong 22 nm, c cha c lipid v protein. Mng cc t bo trong m c cha th th ca LDL, nh t bo hp thu c ht LDL bng nhp bo. Khi LDL c chuyn vo tiu th, cc ester ca cholesterol phn hy, to thnh cholesterol t do xy dng mng. mt s ngi do di truyn, gen ca th th LDL b ri lon. T bo hoc b thiu th th LDL, hoc c nhng b mt kh nng tp trung vo cc lm mc o. Kt qu l LDL khng c hp thu m li lu thng trong mu. LDL trong mu tng cao nn lng ng trn thnh mch, gy x va v thu hp lng mch. c. Thc bo:
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Thc bo l mt dng nhp bo c bit, trong nhng ht kh ln v kch thc (vi sinh vt, mnh xc t bo...) c chuyn vo bn trong bo tng qua c ch gi tc v nhp mng. Ch c hai loi t bo thc hin c chc nng ny l i thc bo v bch cu ht trung tnh. i thc bo lu thng trong mu hoc tp trung mt s c quan hng ro ngn chn vi sinh vt xm nhp. Trong gan, tp trung to thnh nhng cu trc hnh ng, dng mu chy qua ng s c lc khi cc ht l. Thc bo c thc hin vi cc ht c kch thc ln (250 nm hoc hn na). Trc ht, cc khng nguyn trn b mt ca ht c gn vi khng th tng ng. Ht kch thc ln c rt nhiu khng th bao bc xung quanh. Mi khng th

(immunoglobulin) u cha u bin ng v ui hng nh (chui Fc).

Khng th nhn bit v gn vi khng nguyn thng qua u bin ng.

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Chui Fc khng tham gia tng tc vi khng nguyn nn trng thi t do v hng ra pha ngoi phc hp khng nguyn-khng th. Sau Fc tng tc vi th th tng ng (th th Fc) trn b mt i thc bo hoc bch cu ht trung tnh.

Tng tc Fc-th th c th bt u t mt im tip xc gia ht vi mng, sau lan rng ra v bao trm ht b mt ca ht. Mng bo tng cng vi dch bo tng vn ra v bao trm ton b b mt ca ht l c gi l gi tc, hin tng to thnh gi tc cn c gi mt cch hnh nh l c ch khuy ko mng (membranezippering mechanism).

Ngoi vai tr ca Fc-th th theo c ch khuy ko, trong s chuyn ng ca mng bo tng to thnh gi tc cn c vai tr ca mng li protein si actin nm pha di mng. Ngoi actin, clathrin cng c tm thy c nhiu trong gi tc. Tuy nhin vai tr ca clathrin trong chuyn ng gi tc cha c sng t.
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Cc ht c thc bo thnh ti thc bo (phagosom). Ti ny nhp vi tiu th to thnh tiu th th cp (khng bo tiu ho). Bn trong tiu th th cp cc ht c tiu ho, cht ho tan c chuyn vo dch bo tng. Mng ca khng bo tiu ho cng tch ra cc ti vn chuyn nh hon tr vt liu lipit cho mng bo tng. Cui cng tiu th th cp vi vt cht khng tiu ho to thnh th cn b v c xut bo. C ch thc bo ng vai tr rt quan trng trong s khng ca c th i vi cc tc nhn gy bnh t bn ngoi xm nhp. Ngoi vic bt hot v tiu hy vi sinh vt, i thc bo cn lm nhim v trnh din khng nguyn ca vi sinh vt cho lympho T nhn bit. Lympho T sau khi "nhn din khng nguyn" s hot ho v sn xut khng th chng li khng nguyn . i thc bo cn ng vai tr trong vic tiu hy cc t bo gi c trong c th. Trung bnh mi ngy, trong c th ngi c hn 1011 hng cu b thc bo v tiu hy.

T kha
Gradient - Vn chuyn tch cc, th ng - Hip vn, ng vn, i vn - Khuch tn n gin - Protein knh - Protein ti - Nhp bo - Xut bo - ? bo - Thc bo Bm ATPaza: Bm Na/K, Bm H+, Gradient Na - Ch tit lin tc, ch tit c iu khin - Lm mc o - Nang mc o - Clathrin - Triskelion - Nang trn - Nang nhp bo - Th th - Phi t - Khng nguyn - Khng th

Ti liu c thm
1. James D.Watson: Molecular Biology of The Cell, third edition, Garland Publishing Inc., 1995. 2. P. Baudhuin: Cytologie, Universit Catholique de Louvain, 1997. 3. W. E. Ganong: Review of Medical Physiology, thirteen edition, Lange medical book, 1991. 4. S. Silbernagl, A. Despopoulos: Atlas de poche de Physiologie, 2e dition, Flammarion, 1996

Ebook created by CLB195 Bi ging ca cc BS : Nguyn Quc Dng _ Trn Cng Toi_L Quc S