Ly Sinh Hoc 4901

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ON SUY NGH (ch bin) L VN TRNG
L SINH HC
Hu 2005
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LI NI U
Vit Nam v trn th gii L sinh (biophysics) l mn hc c s c dy trong cc trng i hc Khoa hc, ai hc Y - Dc, i hc Nng nghip, i hc S phm, i hc Thu sn v.v... Hiu c L sinh, cng vi mt s mn khoa hc c bn khc s hiu c cc nguyn l ca cc qu trnh sinh hc, c bit l c ch ho l v bn cht vt l ca cc hin tng sng. vit c gio trnh L sinh va bao hm nhng kin thc c bn va cp nht nhng thng tin mi, tht l mt cng vic ht sc kh khn. Nhng vi mong mun phc v kp thi kin thc c bn v L sinh trong khun kh ca d n Gio dc i hc mc B, chng ti mnh dn vit gio trnh ny. Gio trnh L sinh khng ch phc v cho cc sinh vin ca cc trng thuc i hc Hu m cn l ti liu cn thit cho nhng ai quan tm n chuyn ngnh L sinh. Phn cng ni dung: - M u, chng 1, 2, 3, 7, 8, 9 do Tin s on Suy Ngh bin son. - Chng 4, 5, 6 do Tin s L Vn Trng bin son. vit c gio trnh ny, chng ti xin chn thnh cm n Ban d n Gio dc i hc ca i hc Hu, GS. TS. Nguyn Th Kim Ngn v GS. TSKH. L Don Din c v ng gp nhng kin qu bu cho gio trnh ny. gio trnh ngy cng hon thin, chng ti mong nhn c s gp ca ngi c gn xa. Ch bin Tin s: on Suy Ngh
M U L SINH, S HNH THNH V PHT TRIN

S p dng kin thc vt l vo nghin cu sinh hc c thc hin vo cui th k XVIII. Nm 1780 hai nh khoa hc Php l Lavoadie v Laplace tin hnh th nghim kho st tnh ng n ca nh lut I nhit ng hc khi p dng vo h thng sng. Nm 1791, Galvani, gio s gii phu trng i hc Bolon (Italia) cng b kt qu nghin cu trong quyn sch "Bn v cc lc in ng vt trong co c", khng nh c tn ti dng in sinh vt. Nm 1859, Raymond pht hin phn trc v phn sau cu mt ng vt c xng sng tn ti mt hiu in th v o c gi tr t 10 n 38mV, gi l in th tnh (hay in th ngh ngi). Nm 1865, Holgreen pht hin c gi tr hiu in th gia phn trc v phn sau cu mt ng vt c xng sng s tng ln khi mt c chiu sng. Sau ny cc nh khoa hc xc nh, chnh l in th hot ng (hay in th hng phn). Nm 1875, Calton khng nh khi mt c chiu sng, khng nhng in cu mt tng ln nh Holgreen pht hin m in vng th gic trn bn cu i no cng tng ln. Sau ny cc nh khoa hc xc nh chnh l dng in hng phn xut hin khi mt c chiu sng, lan truyn theo dy thn kinh th gic ti vng th gic trn bn cu i no, dn ti hiu ng sinh hc l cm nhn c nh sng. Nm 1922, Erlanger v Gasser dng dao ng k m cc o dng in hng phn xut hin trong dy thn kinh. Nm 1922,Vin L sinh Lin X c c thnh lp. Nm 1929, Berger ghi c in no ca ng vt. Lch s hnh thnh L sinh c Taruxop, gio s trng ai hc tng hp Lomonoxop khng nh: "L sinh c xem nh l mt khoa hc bt u c hnh thnh t th k XIX". Th k XX l th k pht trin mnh m nhng nghin cu khoa hc v L sinh trong cc lnh vc: Nhit ng hc, ng hc ca cc qu trnh sinh vt, vn chuyn cht qua mng t bo, quang sinh hc v phng x sinh hc v.v... Thi k u L Sinh c xc nh nh l mt ngnh khoa hc nghin cu cc hin tng vt l trong h thng sng. Sau L sinh c xc nh nh l mt ngnh khoa hc nghin cu cc c ch vt l, c bit l c ch ho l ca cc qu trnh xy ra trong h thng sng mc phn t, t bo, m v c th. Bc sang th k XXI, hng lot vn ang c t ra cho cc nh L sinh cn phi nghin cu. l nng lng sinh hc, s chuyn ho nng lng v s dng nng lng ca h thng sng? Bn cht v c ch hnh thnh in th sinh vt? Hin tng phn cc trong h thng sng xy ra nh th no v c g khc so vi h vt l ? Bn cht ca qu trnh hng phn l vn cn phi tip tc nghin cu. Cc ch s c trng v vt l v ho l i vi t bo, m, c quan, c th c mi lin quan nh th no trong h thng tin ho ? Vn t iu chnh cc qu trnh sinh hc ca c th sng trc nhng thay i ca yu t mi trng cng ang c cc nh L sinh quan tm nghin cu. Sinh hc phng x hin ang thu ht nhiu nh khoa hc i su nghin cu nhm phc v cho cng tc chn ging mi, bo qun lng thc, thc phm, cng cuc chinh phc v tr, s dng nng lng ht nhn v mc ch ho bnh
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com v khng loi tr kh nng c cuc chy ua v trang trong vic nm gi "n ht nhn u tin" vi tham vng b quyn th gii ?
Chng 1
NHIT NG HC H SINH VT
I. Nhit ng hc h sinh vt v hng nghin cu
Nhit ng hc h sinh vt l lnh vc nghin cu hiu ng nng lng, s chuyn ho gia cc dng nng lng, kh nng tin trin, chiu hng v gii hn t din bin ca cc qu trnh xy ra trong h thng sng. C th sng trong qu trnh sinh trng v pht trin u c s dng nng lng v vy nhit ng hc h sinh vt l lnh vc cn c nghin cu. i tng nghin cu ca nhit ng hc h sinh vt l c th sng, l mt h m do lun xy ra s trao i vt cht v nng lng vi mi trng xung quanh, c kh nng t iu chnh, t sinh sn... nn khc vi h vt l nh cht rn, cht lng hay cht kh... Hin nay nhit ng hc h sinh vt c cc hng nghin cu ch yu sau: - Nghin cu s chuyn bin nng lng mc phn t, t bo, m, c quan hay ton b c th khi trng thi sinh l bnh thng v trng thi ang hot ng. Xc nh hiu sut s dng nng lng ca cc qu trnh sinh vt v nng lng lin kt trong cc lin kt ca cc cao phn t sinh hc. - Nghin cu tnh cht nhit ng ca cc qu trnh din ra trong c th sng nh qu trnh khuych tn, thm thu, vn chuyn tch cc... - Nghin cu c ch tc ng ca s thay i cc yu t mi trng ln qu trnh chuyn ho nng lng v s trao i nng lng gia c th sng vi mi trng.
II. Mt s khi nim v i lng c bn

- H: H l mt vt th hay mt nhm vt th c dng lm i tng nghin cu. V d khi chn c th nghin cu th c th l mt h cn khi chn qun th nghin cu th qun th l mt h. - H c lp: L h khng c s trao i vt cht v nng lng gia h vi mi trng xung quanh. Trn thc t kh xc nh c mt h c lp hon ton nhng qui m th nghim cc nh khoa hc c th thit k c h c lp nh bom nhit lng dng nghin cu hiu ng nhit ca cc phn ng oxy ha. - H kn: L h khng trao i vt cht vi mi trng xung quanh nhng c trao i nng lng vi mi trng xung quanh. - H m: L h c trao i c vt cht v nng lng vi mi trng xung quanh. V d: c th sng l mt h m. - Tham s trng thi: L cc i lng c trng cho trng thi ca mt h, v d nh nhit , p sut, th tch, ni nng, entropi... - Trng thi cn bng: L trng thi trong cc tham s trng thi t mt gi tr nht nh v khng i theo thi gian. - Qu trnh cn bng: L qu trnh trong cc tham s trng thi thay i vi tc chm ti mc sao cho ti mi thi im c th xem nh trng thi ca h l trng thi cn bng. - Qu trnh ng nhit, ng p, ng tch l qu trnh din ra trong nhit , p sut v th tch lun khng i trong sut qu trnh din ra.
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com - Qu trnh thun nghch: L qu trnh bin i m khi tr v trng thi ban u khng km theo bt c mt s bin i no ca mi trng xung quanh. - Qu trnh bt thun nghch: L qu trnh bin i m khi tr v trng thi ban u lm thay i mi trng xung quanh. - Hm trng thi: Mt i lng c xem l mt hm trng thi, c trng cho trng thi ca h, khi s bin thin gi tr ca n trong bt c qu trnh no cng ch ph thuc vo gi tr u v gi tr cui m khng ph thuc vo con ng chuyn bin. Ni nng (U), nng lng t do (F), th nhit ng (Z hay G), entanpi (H), entropi (S) l nhng hm trng thi. - Nng lng: Nng lng l i lng c th o c, c th bin i mt cch nh lng lun theo cng mt t l thnh nhit lng. Nng lng phn nh kh nng sinh cng ca mt h. n v dng o nng lng l Calo (Cal) hay Joule (J). - Cng v nhit: l hai hnh thc truyn nng lng t h ny sang h khc. Nu nh s truyn nng lng t h ny sang h khc gn lin vi s di chuyn v tr ca h th s truyn c thc hin di dng cng. V d khi chy 100 mt th nng lng tiu tn c dng vo thc hin cng di chuyn v tr. Nu s truyn nng lng t h ny sang h khc lm tng tc chuyn ng ca phn t h nhn nng lng th s truyn c thc hin di dng nhit. Cng v nhit l hm s ca qu trnh v chng u ph thuc vo cch chuyn bin. - Ni nng: Ni nng ca mt vt th bao gm ng nng ca cc phn t chuyn ng v th nng tng tc do s ht v y ln nhau gia cc phn t cng vi nng lng ca ht nhn nguyn t v nng lng ca cc in t. III. nh lut I nhit ng hc v nhng h qu ca n nh lut I nhit ng hc c hnh thnh qua cc cng trnh nghin cu ca cc tc gi nh M. V. Lomonoxob (1744), G. I. Heccer (1836), R. Majo (1842), Helmholtz (1849), Joule (1877)... nh lut I nhit ng hc c pht biu nh sau: "Trong mt qu trnh nu nng lng dng ny bin i th nng lng dng khc s xut hin vi lng hon ton tng ng vi gi tr ca nng lng dng ban u". nh lut I nhit ng hc bao gm hai phn: - Phn nh tnh khng nh nng lng khng mt i m n ch chuyn t dng ny sang dng khc. - Phn nh lng khng nh gi tr nng lng vn c bo ton (tc gi nguyn gi tr khi qui i thnh nhit lng) khi chuyn t dng nng lng ny sang dng nng lng khc. Gi tr nng lng ch c bo ton khi qu trnh xy ra l qu trnh thun nghch v hiu sut ca qu trnh t 100%. i vi qu trnh bt thun nghch, hiu sut ca qu trnh nh hn 100% th ngoi phn nng lng truyn cho h phi cng thm phn nng lng to ra mi trng xung quanh. Biu thc ton hc ca nh lut I nhit ng hc: Mt h c lp trng thi ban u c ni nng U1, nu cung cp cho h mt nhit lng Q th mt phn nhit lng h s dng thc hin cng A, phn cn li lm thay i trng thi ca h t trng thi ban u c ni nng U1 sang trng thi mi c ni nng U2 (U2>U1). T nhn xt trn ta c biu thc: Q = U + A (1.1)
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com Trong U = U2 - U1 Cng thc (1.1) c th vit di dng: U = U1 - U1 = Q - A (1.2) i vi qu trnh bin i v cng nh, phng trnh (1.2) c th vit di dng: dU = Q - A (1.3) dU: Ch s bin i ni nng, l hm s trng thi Q v A: Ch s bin i nhit v cng, l hm s ca qu trnh. T biu thc (1.2), nh lut I nhit ng hc c th pht biu nh sau: "S bin thin ni nng ca h bng nhit lng do h nhn c tr i cng do h thc hin". T nh lut I nhit ng hc dn n cc h qu sau y: - Nu h bin i theo mt chu trnh kn (c trng thi u v trng thi cui trng nhau) th ni nng ca h s khng thay i (U2 = U1U = 0). - Khi cung cp cho h mt nhit lng, nu h khng thc hin cng th ton b nhit lng m h nhn c s lm tng ni nng ca h. Theo (1.2) U = U2 - U1 = Q - A, nu A = 0 U2 - U1 = Q. H nhn nhit nn Q > 0 U2 - U1 = Q > 0 U2 > U1. - Khi khng cung cp nhit lng cho h m h mun thc hin cng th ch c cch l lm gim ni nng ca h. Theo (1.2) U = U2 - U1 = Q - A, nu Q = 0 U2 - U1 = -A A = U1 - U2. H mun thc hin cng, tc A > 0 U1 - U2 > 0 U1 > U2. Sau khi thc hin cng (tc A > 0), ni nng ca h gim t U1 xung U2 nh hn. - H thc hin theo chu trnh kn, nu khng cung cp nhit lng cho h th h s khng c kh nng sinh cng. Theo (1.2) U = Q - A, nu h thc hin theo chu trnh kn, theo h qu 1 th U = 0 Q-A=0Q=A Do vy, nu Q = 0, tc khng cung cp nhit lng cho h th h cng khng c kh nng sinh cng, tc A = 0. H qu ny, c th pht biu di dng: "Khng th ch to c ng c vnh cu loi mt, l loi ng c khng cn cung cp nng lng nhng vn c kh nng sinh cng". IV. nh lut Heccer c Heccer tm ra nm 1836, sau ny c cc nh khoa hc xp thuc vo h qu ca nh lut I nhit ng hc. nh lut Heccer pht biu nh sau: "Hiu ng nhit ca cc phn ng ho hc ch ph thuc vo dng v trng thi ca cht u v cht cui m khng ph thuc vo cch chuyn bin". V d: Phn ng to kh CO2 t than nguyn cht l cacbon (C) c th tin hnh theo 2 cch sau: Cch 1: t trc tip than nguyn cht thnh kh CO2 s gii phng nhit lng l Q1. Phn ng xy ra:
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com C + O2 CO2 + Q1 Cch 2: Chuyn than nguyn cht thnh CO theo phn ng: C+
1 O2 CO + Q2 2
T CO chuyn tip thnh CO2 theo phn ng: CO +
1 O2 CO2 + Q3 2
Q1 C Q2 CO Q3 CO2
S minh ho:
Theo nh lut Heccer, cht u tham gia phn ng (C) v sn phm ca phn ng (CO2) ging nhau nn c hiu ng nhit ging nhau: Q1 = Q2 + Q3 Trong thc nghim, hiu ng nhit ca qu trnh t than thnh CO khng th o trc tip c v khi than chy khng bao gi ch cho CO m cn cho mt t CO2. Nhng thc nghim li o trc tip c Q1 = 97 KCal/M v Q3 = 68 KCal/M. T d dng suy ra gi tr Q2 = Q1 - Q3 = 97 KCal - 68 KCal Q2 = 29 KCal/M. nh lut Heccer c ngha rt quan trng i vi h sinh vt. Trong h sinh vt din ra nhiu phn ng phc tp, cho n nay vn cn nhiu phn ng trung gian cha c th o trc tip c hiu ng nhit. Da vo nh lut Heccer c th gii quyt c kh khn ny. V. nh lut I nhit ng hc p dng vo h sinh vt Ngi u tin tin hnh th nghim chng minh tnh ng n ca nh lut I nhit ng hc khi p dng vo h thng sng l hai nh khoa hc Php Lavoisier v Laplace vo nm 1780. i tng th nghim l chut khoang. Th nghim cch ly c th khi mi trng bn ngoi bng cch nui chut trong nhit lng k nhit 0oC. Dng mt lng thc n xc nh trc nui chut th nghim. Trong c th chut s din ra cc phn ng phn hu thc n ti sn phm cui cng l kh CO2 v H2O, ng thi gii phng ra nhit lng Q1. Nu coi iu kin 0oC, chut ng yn, khng thc hin cng m ch s dng nhit lng gii phng ra do oxy ho
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com thc n cung cp nhit lng cho c th v ta nhit ra mi trng, qua nhit k o c s tng nhit , theo cng thc s tnh c nhit lng Q1. ng thi ly mt lng thc n tng ng vi lng thc n cho chut n trc khi th nghim em t chy trong bom nhit lng k cng ti kh CO2 v H2O, gii phng ra nhit lng Q2. So snh hai kt qu th nghim thy gi tr Q1 tng ng vi Q2. iu ny chng t nhit lng gii phng ra t cc phn ng ho sinh din ra trong c th sng hon ton tng ng vi nhit lng gii phng ra t cc phn ng xy ho din ra ngoi c th sng. Ni cch khc, hiu ng nhit ca qu trnh xy ho cht din ra trong c th sng v hiu ng nhit ca qu trnh xy ho cht din ra ngoi c th sng l hon ton tng ng. tng chnh xc ca th nghim, sau ny c nhiu m hnh th nghim ca nhiu nh nghin cu c tin hnh nhng ng ch nht l ca Atwater v Rosa vo nm 1904. i tng th nghim l ngi v thi gian th nghim l mt ngy m (24 gi). Trong thi gian th nghim, cho ngi tiu th mt lng thc n nht nh, thng qua o lng kh xy ht vo (hay kh CO2 th ra), nhit thi ra t phn v nc tiu... s tnh c hiu ng nhit ca cc phn ng phn hu thc n din ra c th ngi trong 24 gi. ng thi t lng thc n tng ng vi lng thc n m ngi tiu th trong bom nhit lng k s o c nhit lng to ra. Kt qu th nghim: Hiu ng nhit ca cc phn ng din ra c th ngi trong 24 gi: Nhit lng to ra xung quanh : 1374 KCal Nhit lng to ra do th ra : 181 KCal Nhit lng to ra do bc hi qua da: 227 KCal Nhit do kh thi ra : 43 KCal Nhit to ra t phn v nc tiu : 23 KCal Hiu nh (do sai s) : 31 KCal Tng cng nhit lng thi ra : 1879 KCal Nhit lng do thc n cung cp: 56,8 gam Protein : 237 KCal 79,9 gam Gluxit : 335 KCal 140,0 gam Lipit : 1307 KCal Tng cng : 1879 KCal Lu : Khi xy ho 1 gam Protein trong bom nhit lng k ti kh CO2 v H2O, gii phng ra 5,4 KCal cn trong c th sng phn gii 1 gam Protein ti ur ch gii phng khong 4,2 KCal. Khi oxy ho hon ton 1 gam Gluxit, gii phng khong 4,2 KCal cn xy ho hon ton 1 gam Lipit gii phng t 9,3 n 9,5 KCal. Kt qu th nghim ca Atwater v Rosa khng nh nng lng cha trong thc n sau khi c th tiu th chuyn thnh nng lng gii phng thng qua qu trnh phn gii bi cc phn ng ho sinh din ra trong c th sng. Nng lng cha trong thc n v nng lng gii phng ra sau khi c th phn gii thc n l hon ton tng ng. Nhit lng trong c th ngi c chia lm hai loi l nhit lng c bn (hay nhit lng s cp) v nhit lng tch cc (hay nhit lng th cp). Nhit lng c bn xut hin ngay sau khi c th hp th thc n v tiu th xy thc hin phn ng xy ho
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com ng thi gii phng ra nhit lng. V d khi c th hp th 1 phn t gam (tc 1M) glucose, lp tc xy ra phn ng xy ho ng v gii phng ra 678 KCal (nhit lng c bn). C th s s dng nhit lng c bn vo cc hot ng sng, nu cn d s c tch lu vo ATP. Phn nhit lng tch lu vo cc hp cht cao nng gi l nhit lng tch cc. Trong c th sng, nhit lng c bn v nhit lng tch cc c lin quan vi nhau. Nu nhit lng c bn nhiu m c th s dng t th nhit lng tch cc s tng ln. Nu nhit lng c bn khng c th khng nhng nhit lng tch cc bng khng m c th phi phn gii ATP, gii phng ra nng lng cung cp cho cc hot ng sng. trng thi sinh l bnh thng, c th sng s duy tr mi tng quan nht nh gia nhit lng c bn v nhit lng tch cc. mc t bo, c khong 50% nng lng ca cht dinh dng c tch lu vo ATP.
VI. Phng php nhit lng k gin tip v nguyn tc hot ng ca c th sng
Phng php o nhit lng ca Lavoadie v Laplace dng trong th nghim chng minh tnh ng n ca nh lut I nhit ng hc khi p dng vo h sinh vt, gi l phng php nhit lng k gin tip. C s ca phng php ny l da vo lng kh xy tiu th hoc lng kh CO2 do c th thi ra ng vt mu nng (ng vt c v v ngi), c lin quan cht ch vi nhit lng cha trong thc n. V d: Qu trnh xy ha glucose, phn ng din ra nh sau: C6H12O6 + 6O2 = 6CO2 + 6H2O + 678 KCal (180gam) (134,4l) (134,4l) T phn ng trn cho thy c xy ho hon ton 1 phn t gam glucose th cn phi tiu th 6 phn t gam xy ng thi thi ra 6 phn t gam kh CO2 v gii phng ra 678 KCal. iu kin tiu chun, mi phn t gam cht kh u cha 22,4 lt. Do vy 6 phn t gam xy hoc CO2 u cha: 6 x 22,4 lt = 134,4 lt. T suy ra, c th c tiu th 1 lt O2 xy ho hon ton mt phn t gam glucose ng thi thi ra 1 lt CO2 th km theo gii phng mt nhit lng l: 678 KCal: 134,4 lt = 5,047 KCal/lt v gi l ng lng nhit ca xy. Da vo phng php nhit lng k gin tip, c th xc nh c s thi nhit ca bt k ng vt mu nng no thng qua s lt xy tiu th (hoc s lt CO2 thi ra). T phn ng xy ha glucose trn v sau ny p dng chung cho Gluxit khi xy ho hon ton s gii phng ra nhit lng c tnh theo cng thc: Q(KCal) = s lt O2 ( hoc s lt CO2) x 5,047 (1.4) Khi xy ha Protein, nhit lng gii phng ra c tnh theo cng thc: (1.5) Q(KCal) = s lt O2 x 4,46 Khi xy ho Lipit, nhit lng gii phng ra c tnh theo cng thc: Q(KCal) = s lt O2 x 4,74 (1.6) Mi quan h gia thc n, s lt O2 tiu th v s lt CO2 thi ra cng ng lng nhit ca xy c th hin qua bng 1.1.
Bng 1.1: ng lng nhit ca xy i vi cc loi thc n.
Thc n
S lt O2 cn xy ho 1 gam thc n
s lt CO2 thi ra sau khi xy ho 1g thc n
ng lng nhit ca xy
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com Gluxit Protein Lipit 0,83 0,97 2,03 0,83 0,77 1,42 5,047 4,46 4,74
i vi thc n hn hp gm c Gluxit, Protein v Lipit khi b xy ho, nhit lng gii phng ra c tnh theo cng thc: Q(KCal) = s lt O2 x 4,825 (1.7) Phng php nhit lng k gin tip cn c th xc nh c nhit lng gii phng ra khi xy ho thc n thng qua: Thng s h hp l t l kh CO2 trn kh O2. Thng s h hp cng thay i tu thuc vo loi thc n c xy ho. - i vi phn ng xy ho glucose C6H12O6 + 6O2 = 6CO2 + 6H2O S lt kh CO2 Thng s h hp = S lt O2 = 6 x 22,4 lt 6 x 22,4 lt =1
Thng s h hp ca glucose c s dng cho c Gluxit. - i vi phn ng xy ha Lipit c thng s h hp bng 0,7, i vi Protein bng 0,8 cn vi thc n hn hp c gi tr nm trong khong t 0,85 n 0,9. Thng s h hp c lin quan vi ng lng nhit ca xy, th hin qua bng 1.2.
Bng 1.2: Thng s h hp (TS h hp) v ng lng nhit ca xy (LN ca xy)
TS h hp
0,7
0,75
0,8
0,85
0,9
0,95
1 5,05
LN ca xy 4,686 4,739 4,801 4,862 4,924 4,985
Khi xy ho thc n, bng cch o lng kh O2 tiu th v lng kh CO2 thi ra (n v l lt), tnh c thng s h hp. Da vo bng 1.2, ly gi tr ng lng nhit ca xy tng ng vi thng s h hp nhn vi s lt O2 tiu th s bit c nhit lng gii phng (cn gi l lng nhit trao i hay tr s trao i nng lng). V d: Nu thng s h hp l 0,85 th c ng lng nhit ca xy l 4,862 v bit c th tiu th 20 lt O2 th tr s trao i nng lng s l: 4,862 x 20 lt O2 = 97,24 KCal
VII. Phn bit nguyn tc hot ng ca c th sng vi my nhit

My nhit l ng c dng nhin liu t nh du Diedel, xng sinh cng nh my n t, xe my, my bm nc... Trong c hc, hiu sut s dng nng lng ca mt ci my nhit, c tnh theo cng thc: T T = 2 1 (1.8) T2 : Hiu sut ca my nhit (%) T1: Nhit Kelvin trng thi u (oK) T2: Nhit Kelvin trng thi cui (oK)
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com Cng thc chuyn i gia nhit Kelvin v nhit bch phn (oC): T(oK) = t(oC) + 273 (1.9) Gi s hiu sut s dng nng lng ca mt my nhit t trung bnh l 33% = 33/100 1/3 Nu ta cng gi s c th sng hot ng ging nh mt my nhit, tc l cng c hiu sut s dng nng lng l 33% ? Nhit ban u ca c th ngi l 37oC, theo cng thc (1.9) tnh ra: T1 = 37 + 273 = 310oK 1 Thay = 33% v T1 = 310oK vo cng thc (1.8) s c: 3
1 T2 310 o K 3.(T2 - 310oK)=T2 = 3 T2 2T2 = 930oK T2 = 465oK p dng cng thc (1.9) ta c: 465 = t2(oC) + 273 t2 = 465 - 273 = 192oC Kt qu trn cho thy c th sng hot ng khng ging nh mt my nhit. i vi c th sng, Protein b bin tnh ngay nhit t 40oC n 60oC cn 192oC th khng c mt sinh vt nhn chun no c th sng c. iu khng nh c th sng hot ng khng ging nh mt my nhit m hot ng theo nguyn l ca cc qu trnh sinh hc.
VIII. nh lut II nhit ng hc

nh lut I nhit ng hc ch cho bit v s bin i gia cc dng nng lng khc nhau, cho php xc nh biu thc ch r s lin quan v lng gia cc dng nng lng khc nhau khi xut hin trong mt qu trnh cho trc. Song nh lut I nhit ng hc khng cho bit qu trnh khi no c th xy ra hoc khng xy ra v chiu hng din bin ca qu trnh nu xy ra th theo chiu hng no? nh lut II nhit ng hc xc nh c chiu hng t din bin ca mt qu trnh cng nh cho bit qu trnh t din bin n khi no th dng li v cho php nh gi kh nng sinh cng ca cc h nhit ng khc nhau. nh lut II nhit ng hc c ba cch pht biu. Cch pht biu th nht cn gi l tin Clausius a ra 1850: "Nhit khng th t ng truyn t vt lnh sang vt nng". T suy ra rng nhit ni ring cn nhng qu trnh nhit ng ni chung ch c th t din ra nu xy ra s truyn nng lng t mc cao n mc thp, tc l theo chiu gradien. Gradien ca mt thng s c trng cho mt tnh cht no v trng thi ca h (nh nng ) c xc nh bng hiu s gi tr ca thng s ti hai im chia cho khong cch gia hai im . V d: Gi s ti v tr 1 c nng l C1 cn v tr 2 c nng l C2 v C1 > C2 th hiu s nng s l C1 - C2 cn hiu s khong cch l x2 - x1. t C = C1 - C2 v x = x2 - x1 th gradien nng c xc nh: C C1 C 2 = x x 2 x1 (1.10)
C1 > C2
x1
B
x2
B
Gradien l mt i lng vect, c gi tr v hng v gi tr ln. Khi so snh mt t bo sng vi mt vt v sinh nh mt ht ct ta thy r ngay rng trong t bo sng duy tr nhiu loi gradien khc nhau. Gradien mng duy tr in th tnh v in th hot ng, gradien nng duy tr nng , gradien p sut thm thu duy tr lng nc trong t bo... Nu t bo cht th cc loi gradien cng b trit tiu. Nu xt mc gradien th s sng ca t bo lun km theo s tn ti ca cc loi gradien. Cch pht biu th hai do Thomson pht trin tin ca Clausius "Khng th c mt qu trnh bin i chuyn ton b nhit lng thnh cng". Theo cch pht biu ca Thomson th hiu sut hu ch ca qu trnh bao gi cng nh hn 1 (tc < 1). iu ny c ngha trong t nhin khng c mt qu trnh no c th chuyn ton b nhit lng c cung cp thnh cng hu ch. i vi cc qu trnh din ra trong h thng sng c tun theo cch pht biu ca Thomson hay khng? Vn ny s cp n phn sau. Cch pht biu th ba trn c s kin ca Planck, cho rng Entropi l mt tiu chun y v cn thit xc nh tnh thun nghch v khng thun nghch ca bt c qu trnh vt l no din ra trong thin nhin. nh lut II nhit ng hc pht biu nh sau: "i vi h c lp, mi qu trnh trong t nhin u din bin theo chiu tng ca entropi". Vy entropi l g? hiu r i lng ny ta xt v d v nguyn l hot ng ca my nhit. Theo hnh 1.1, nguyn l hot ng ca my n nh sau: My ch c kh nng sinh cng A (tc bnh quay) khi c cung cp nng lng l xng. Khi xng b t cp nhit T Ngun cung 1 chy c nhit l T1 v gii phng nhit lng l Q1. Mt phn ca nhit lng Q1 dng sinh cng, phn Q 1 cn li truyn cho ngun nc lm lnh my l Q2, dn n lm tng nhit ca nc l T2. y Q1>Q2 My sinh v T1>T2 cng Theo (1.8) th hiu sut hu ch ca qu trnh thun nghch c xc nh theo cng thc: Q2 A Q1 Q 2 T1 T2 = = = (1.11) T2 Ngun thu nhit Q1 Q1 T1 Cn bng phng trnh ta c: T1(Q1-Q2)=Q1(T1-T2) T1Q1-T1Q2=Q1T1-Q1T2 Hnh 1.1: Nguyn l hot ng ca my n Q1 Q 2 = Gin c ta c: T1Q2=Q1T2 hay (1.12) T1 T2 T vt l hc cho bit s thay i entropi ca mt h c xc nh theo cng thc: Q (1.13) S = T S: s thay i entropi ca h.
Q: Nhit lng cung cp cho h (calo) T: Nhit Kelvin (oK) ca h i vi qu trnh bin thin v cng nh, ta c: Q (1.14) dS = T n v ca entropi l Cal/M. Entropi l mt hm trng thi nn n ch ph thuc vo trng thi u v trng thi cui cng ca h. Cng thc (1.12) c th biu din qua hm entropi nh sau: Q Q S1 = 1 ; S2 = 2 T1 T2 S1: Entropi trng thi u S2: Entropi trng thi cui i vi qu trnh thun nghch theo cng thc (1.12) ta c: S1=S2 S = Const (hng s) (1.15) Trong mt h nu ch xy ra cc qu trnh thun nghch th h lun duy tr trng thi cn bng nn entropi ca h l khng i. i vi qu trnh khng thun nghch th
S >
lm thay i entropi ca mi trng xung quanh do s ma st v ta nhit. Thc nghim xc nh i vi mt qu trnh khng thun nghch th entropi ca h trng cui (tc S2) bao gi cng ln hn so vi entropi ca h trng thi u (tc S1). Do vy: S2-S1>0 (1.16) Trong mt h xy ra cc qu trnh khng thun nghch th entropi ca h bao gi cng tng ln. Do vy, nu l h c lp th cc qu trnh xy ra trong h s tin trin theo chiu tng ca entropi v entropi ca h s t gi tr cc i trng thi cn bng nhit ng. Tnh chung cho c qu trnh thun nghch v khng thun nghch th s thay i entropi ca h c th vit nh sau: S 0 (1.17) i vi qu trnh thay i entropi v cng nh (gi l qu trnh vi phn) th: dS 0 (1.18) (Du bng dng cho qu trnh thun nghch cn du ln hn dng cho qu trnh khng thun nghch). Vt th H c lp V d 1: H c lp gm vt th A v vt th B. Vt A th A c nhit l TA cn vt th B c nhit l TB. Gi s TA>TB th theo nh lut II nhit ng hc vt A s truyn cho vt B mt nhit lng l Q. Q S thay i entropi ca vt A do mt nhit l: Q dSA= cn s thay i entropi ca vt B do TA B
B B
Q v nhit lng cung cp cho h khng ch lm thay i entropi ca h m cn T
Vt th
nhn nhit l dSB=

B
Q (du tr ch entropi ca vt TB
Cc nc Lp cch V TA>TB nn TA-TB>0 suy ra dS>0 nhit hon (0oC) ton V d 2: Ngm cc nc c nhit l 0oC (tc tnh ra nhit Kelvin l: T1=273+0oC=273oK) vo trong thng du 100oC Thng du c nhit l 100oC. H c lp (tc T2=273+100=373oK). Nhit s t ng truyn t du sang cc nc qui ra nhit lng l Q. Thc nghim xc nh c Q=80cal. Du cung cp nhit nn s thay i entropi ca du l: 80 dSD= = 0,214 cal/ (b gim i). 373 Cc nc nhn nhit nn s thay i entropi ca nc l: 80 dD= =0,293cal/. 273 S thay i entropi ca h (gm du v cc nc ) s l: dSh= dSD+dS = 0,293cal/ - 0,214cal/d = 0,079cal/ > 0 Nhn xt: i vi h c lp, qu trnh truyn nhit t din bin theo chiu tng ca entropi. iu ny minh chng cho tnh ng n ca nh lut II nhit ng hc.
B B
A b gim). S thay i entropi ca ton h c xc nh: Q Q Q(TA TB ) dS=dSA+dSB= = TB TA TA .TB

B
IX. Tnh cht thng k ca nh lut II nhit ng hc

nh lut II nhit ng hc c tnh cht thng k nn chnh xc ph thuc vo s phn t c trong h. Trng thi t trt t (tc c entropi ln) c xc sut cao hn trng thi c trt t cao (tc c entropi nh). Gia entropi v xc sut nhit ng c mi quan h nh th no? Nm 1896 Boltzmann l ngi u tin tm c mi lin quan gia entropi v xc sut nhit ng qua phng trnh: S = klnW (1.19) S: Entropi ca h k: Hng s Boltzmann bng 1,38.10-16ec/ W: Xc sut nhit ng hiu r tnh cht thng k ca nh lut II nhit ng hc, ta xt v d: Trong 1 bnh c 6 phn t c nh s t s 1 n s 6. Ta ngn bnh thnh 2 phn: pha bn phi khng c phn t no c. - Thi im ban u: Bn tri Bn phi 1 2
3 5 4 6
Thnh ngn
Sau khi b thnh ngn do chuyn ng nhit ca cc phn t nn cc phn t s chuyn ng trong ton b th tch ca bnh to nn tt c c 26 = 64 - Kiu phn b 5 phn t bn tri bnh cn 1 phn t bn phi bnh (c 6 kiu phn b). 2 2 1 1 1 2
3 5 4 6 3 5 4 6 5 4 6 3
Kiu 1
Kiu 2
Kiu 3
1 3 5
2 4 6
1 3
2 4 6 5
1 3 5
2 4 6
Kiu 4 Kiu 5 Kiu 6 Xc sut phn b ca phn t trong bnh c trnh by bng 1.3. Xc sut nhit ng l s kiu c th xy ra ca mi kiu phn b cho nn xc sut nhit ng lun lun ln hn 1 hoc nh nht l bng 1. Xc sut ton hc (k hiu l w) bao gi cng nh hn mt v bng 1 l ln nht. Xc sut ton hc l kh nng xy ra s kiu trn ton b s kiu phn b.
Bng 1.3: S phn b ca cc phn t gia hai phn bnh v xc sut phn b ca chng.
S phn t Xc sut Xc sut nhit ng W ton hc w Bn tri bnh Bn phi bnh 6 0 1 1/64 5 1 6 6/64 4 2 15 15/64 3 3 20 20/64 2 4 15 15/64 1 5 6 6/64 0 6 1 1/64 Kt qu bng 1.3 cho thy s phn b cn bng mi bn c 3 phn t l c xc sut nhit ng ln nht, tc W=20 nn theo cng thc (1.19) th entropi cng c gi tr ln nht. Nh vy, trng thi cn bng l trng thi c xc sut nhit ng ln nht ng thi cng c entropi ln nht nn l trng thi d xy ra nht. Trong 64 kiu phn b ch c 20
kiu tun theo nh lut II nhit ng hc, tin trin theo chiu hng tng ca entropi v t ti entropi ln nht cn 42 kiu (gm 6+15+15+6) cng tin trin theo chiu tng ca entropi nhng cha t ti gi tr entropi ln nht. c bit chiu hng tin trin c 6 phn t cng mt pha ca bnh c xc sut nhit ng v entropi ging vi trng thi ban u (tc entropi khng tng ln). iu ny khng ng vi nh lut II nhit ng hc l qu trnh lun din ra theo chiu tng ca entropi. Khi s phn t trong bnh cng tng ln, v d l 50 phn t th tng s kiu phn b s l 250, l 1 con s cc k ln. Khi kh nng tin trin c c 50 phn t mt pha ca bnh cng ch chim 1 kh nng (tc W=1) so vi 250 kh nng v nh vy trng hp xy ra khng ng vi nh lut II nhit ng hc s cng t i
1 . chnh l tnh thng k ca nh lut II nhit ng 50 2
hc, tc l khi s phn t trong h cng ln th tnh ng n ca nh lut II nhit ng hc s cng cao.
X. nh lut II nhit ng hc p dng vo h sinh vt

H thng sng l mt h thng m, lun xy ra qu trnh trao i vt cht v nng lng vi mi trng ngoi. Theo cch pht biu ca Thomson: "Khng th ch to c ng c vnh cu loi hai" l ng c c hiu sut hu ch l 100%, khi p dng vo h thng sng l hon ton ng n. Thc nghim xc nh mi qu trnh din ra trong h thng sng u c hiu sut hu ch nh hn 100% (xem bng 1.4).
Bng 1.4: Hiu sut ca mt s qu trnh sinh vt
TT 1 2 3 4 5
Qu trnh sinh vt Qu trnh glicoliz (tiu glucoza) Qu trnh co c Qu trnh oxy ha photphorin ha Qu trnh quang hp Qu trnh pht quang ca vi khun
Hiu sut () 36% 30% 55% 75% 90%
Qu trnh quang hp ca thc vt c hiu sut 75% c ngha l cy xanh c hp th 100calo t nng lng nh sng mt tri th c 75 calo c s dng vo tng hp cht (phn nng lng c ch) cn 25 calo ta nhit si m c th hay pht tn nhit ra mi trng xung quanh (phn nng lng v ch). * Vai tr ca entropi i vi h c lp, nh lut II nhit ng hc khng nh mi qu trnh din bin u din ra theo chiu tng ca entropi v t gi tr cc i khi t n trng thi cn bng nhit ng th dng hn. C th sng l mt h m cho nn khng th p dng nh lut II nhit ng hc trc tip ln c th sng. nh lut II nhit ng hc ch c th p dng vo h sinh vt nu xem h bao gm c c th sng v mi trng sng. Ta hy xt s dS thay i entropi ca h (k hiu l ) c lin quan ti s thay i entropi ca c th dt

dS C dSm ) v s thay i entropi ca mi trng (k hiu l ) nh th dt dt no? Entropi l mt hm trng thi v c tnh cht cng tnh (tc l entropi ca h bng tng entropi ca cc thnh phn nm trong h) nn s thay i entropi ca h s c xc nh theo cng thc: dS dSC dS m (1.20) = + dt dt dt dSC = 0 khi cc qu trnh din ra trong c th sng u l qu trnh thun Thnh phn dt nghch. Song trn thc t phn ln cc qu trnh din ra c th sng l nhng qu trnh dSC bt thun nghch, do vy >0 dt dS m = 0 khi gia c th sng v mi trng khng xy ra qu trnh trao i Thnh phn dt vt cht v nng lng. iu ny khng xy ra v tri vi thc t. Qu trnh trao i vt cht v nng lng gia c th sng vi mi trng l qu trnh bt thun nghch (nht l dSC dS m dSm dS qu trnh trao i nhit) do vy >0. Nh vy, khi >0 v >0 th >0. dt dt dt dt iu ny l ph hp vi nh lut II nhit ng hc. Trong c th sng xy ra trng hp khi qu trnh ng ha din ra vi cng mnh dSC hn so vi qu trnh d ha th <0. Trng hp ny gp qu trnh quang hp ca dt dSC <0. B li, qu trnh trao i cht gia thc vt vi mi trng thc vt, khi c dt nc, khng kh v nht l s hp thu nng lng t nh sng mt tri lm tng entropi ca mi trng ln rt cao nht l trung tm mt tri xy ra cc phn ng ht nhn gii phng nng lng nh sng chiu xung tri t. Cc phn ng ht nhn bao dSm gi cng c entropi v cng ln. Do vy, gi tr s ln hn khng rt nhiu nn s dt dS vn ln hn khng, ph hp vi nh lut II nhit ng thay i entropi ca h l dt hc. Tm li khi p dng nh lut II nhit ng hc vo h thng sng th ta khng nn tch ri c th sng ra khi mi trng sng m phi xem c th sng v mi trng sng l cng nm trong mt h. V mi lin quan gia entropi v trt t cu trc ca c th sng, Schrodinger cho rng: "S sng l s hp th entropi m". Gii thch quan im ny, theo tc gi l c th sng lun lun duy tr trt t cao ca mnh bng cch hp th cht dinh dng c trt t cao nh protit, Gluxit, Lipit qua thc n. Thc ra, khi tiu th cht dinh dng, c th sng khng s dng chng nh mt ngun trt t ( nguyn v dng lm nguyn liu xy dng nn c th sng) m cht dinh dng sau khi hp th c phn gii thnh cht t bo c th hp thu c. Chng hn Protein ca tht g c trt t cao (tc entropi thp) khi c c th hp th n s b phn gii thnh cc axit amin nn c
sng (k hiu l
trt t km hn (tc entropi cao). Do vy, quan im ca Schrodinger l hon ton khng phn nh ng bn cht ca qu trnh tiu ha v hp thu ca c th sng. trt t cu trc v trt t ca cc qu trnh sinh hc din ra trong c th sng khng phi do entropi quyt nh m do c th s dng ngun nng lng t do t ngun thc n duy tr s tn ti v pht trin ca c th sng. Trong qu trnh pht sinh v hnh thnh s sng trn tri t, tri qua thi gian tin ha vi s chn lc ca t nhin hnh thnh nn cc loi sinh vt c s thch nghi cao vi tng loi mi trng sng. Do cc nguyn l ca cc qu trnh sinh hc quyt nh lm cho c th sng thch nghi c v mt cu trc cng nh thch nghi v mt chc nng ch khng phi hon ton do entropi quyt nh nh trong h l ha. Tuy nhin mi qu trnh sinh l, sinh ha din ra trong c th sng u km theo s thay i ca entropi. Khi xt entropi ring ca mt c th sng m khng gn vi entropi ca mi trng sng th khi c th trng thi cn bng dng, entropi c mt gi tr xc nh nhng khng phi l cc i v khng i. Khi c th sng nhim phng x, nhim cht c hi, nhim virut th entropi s tng v c gi tr ln hn so vi entropi khi trng thi cn bng dng. Khi c th sng c qu trnh sinh tng hp cht (nh qu trnh quang hp thc vt) din ra mnh hn so vi qu trnh phn hy cht th entropi s gim v c gi tr nh hn so vi entropi khi trng thi cn bng dng l trng thi c tc phn ng tng hp cn bng vi tc phn ng phn hy. S thay i entropi din ra trong c th sng khng vi phm nh lut II nhit ng hc v c th sng l mt h m ch khng phi l mt h c lp.
XI. Nng lng t do

T nh lut I nhit ng hc thit lp c cng thc (1.3) l: dU = Q- A T nh lut II nhit ng hc thit lp c cng thc (1.14) l: Q Q = T.dS dS = T Thay Q = T . dS vo cng thc (1.3) ta c: dU = T . dS - A -A = dU - T . dS (1.21) i vi qu trnh ng nhit (l qu trnh din ra nhit lun khng i) ta c th vit: -A = d(U - T.S) (1.22) Ngi ta t F = U - T.S v gi F l nng lng t do. Nng lng t do l mt hm trng thi . Thay F vo (1. 22) ta c: - dF = A (1.23) Du tr th hin qu trnh s dng nng lng t do thc hin cng. i vi qu trnh thun nghch, nng lng t do, c chuyn ton b thnh cng. T vt l xc nh c i vi qu trnh ng p (l qu trnh din ra p sut lun khng i) nu cng sinh ra ch chng li p sut bn ngoi th cng c tnh theo cng thc: A = P . dV (1.24) A: Cng thc hin
P: p sut dV: S thay i th tch Thay (1.24) vo (1.3) ta c: dU = Q - P . dV Q = dU + P . dV (1.25) i vi qu trnh ng p (P=hng s), cng thc (1.25) c th vit: Q = d(U + P . V) (1.26) Ngi ta t H = U + P . V v gi H l hm entanpi. Entanpi l mt hm trng thi. Cng thc (1.26) c th vit li nh sau: Q = dH (1.27) i vi qu trnh din ra iu kin nhit v p sut khng thay i, ngi ta a vo hm th nhit ng, k hiu l Z v c tnh theo cng thc: dZ = dH - T . dS (1.28) Hm th nhit ng ca phn ng c tnh theo cng thc: Z = Zo + R.T.lnK (1.29) Zo: Th nhit ng chun khi phn ng din ra nhit l 25oC. R: Hng s kh T: Nhit Kelvin (oK) K: H s cn bng ca phn ng Da trn nghin cu s ph thuc h s cn bng ca phn ng vo nhit s xc nh c s thay i ca entanpi theo cng thc: k ln K Q (1.30) = dT RT 2 d ln K (1.31) Q = R .T 2 . dT Xc nh c Q, da vo cng thc (1.27) s tnh ra dH. 1. Qu trnh hnh thnh nng lng t do trong c th sng Nng lng t do c hnh thnh trong c th sng l do qu trnh phn hy cc cht dinh dng. Theo Crebs v Gonberg qu trnh hnh thnh nng lng t do chia lm 3 giai on chnh sau y: - Phn hy cc cao phn t sinh hc ti monome (n phn t), nh t Protein ti axit amin, t Gluxit ti glucose, t Lipit ti glixerin v axit bo. Nng lng t do c gii phng ra giai on ny ch chim t 0,1% n 0,5% nng lng d tr c trong cao phn t. - S chuyn ha ca cc monome k trn ti axit Piruvic v axetyl coenzim A (l axit Axetic hot ha) v mt s hp cht nm trong chu trnh Crebs gii phng ra nng lng t do t t 15% n 30% nng lng d tr c trong monome. - Qu trnh oxy ha axetyl coenzim A ti kh CO2 v H2O trong chu trnh Crebs, nng lng t do c gii phng ra t t 70% n 80% nng lng d tr c trong axetyl coenzim A. Hay qu trnh oxy ha axit palmatic, nng lng t do c gii phng ra chim 60%.
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com 2. S dng nng lng t do ca c th sng - C th sng s dng nng lng t do cung cp nhit cho c th (n nh nhit c th vo ma h cng nh ma ng). - C th sng s dng nng lng t do thc hin cng c hc (co c), cng thm thu (hp th hay bi tit nc v cc sn phm cht dinh dng), cng h hp, cng in (duy tr in th tnh hay pht xung in th hot ng)... - Quan trng hn c l c th sng c kh nng tch ly nng lng t do dng cc hp cht cao nng (ATP). Hp cht cao nng ATP chnh l ngun nng lng vn nng ca mi c th sng nn c v l "tin t nng lng". 3. Trng thi cn bng nhit ng v trng thi cn bng dng Trng thi cn bng nhit ng: L trng thi ch c trng cho h c lp. Khi h trng thi cn bng nhit ng s c nng lng t do t gi tr cc tiu v khng i do vy h khng c kh nng sinh ra cng. Khi h trng thi cn bng nhit ng s c entropi t gi tr cc i, do vy h c mt trt t cao nht. Trn thc t kh bt gp trng thi cn bng nhit ng v kh tm thy h c lp hon ton. Trng thi cn bng dng: L trng thi c trng cho h m ni chung v h sinh vt ni ring. Khi h trng thi cn bng dng th s thay i nng lng t do lun xy ra nhng vi mt tc khng i. S d nh vy l do h lun nhn nng lng t do t bn ngoi qua con ng thc n. Khi h trng thi cn bng dng, entropi ca h t gi tr xc nh v nh hn gi tr cc i. C th sng lun c xu hng duy tr trng thi cn bng dng. V d nh ng vt n nhit lun duy tr thn nhit n nh theo thi gian ( ngi l 37oC).
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Chng 2
NG HC CA CC PHN NG SINH VT
ng hc ca cc phn ng sinh vt l nghin cu nhng c ch v cc qui lut tin trin theo thi gian ca cc phn ng ha sinh din ra trong c th sng. Trong c th sng thng xuyn din ra cc phn ng ca qu trnh trao i cht. Phn ng din ra c th l phn ng bc khng, bc mt, bc 2..., phn ng t xc tc, phn ng ni tip, phn ng song song, phn ng vng, phn ng dy chuyn...Ty iu kin ban u v vi s tham gia ca cht xc tc hay cht c ch m cc phn ng ha sinh xy ra vi tc khc nhau v bng cc con ng khc nhau.
I. Bc phn ng v tc phn ng
Bc phn ng: Bc ca phn ng bng s lng cc loi phn t (hay nguyn t) tham gia trong mt n v c bn ca chuyn bin ho hc m nng hay p sut ca chng quyt nh tc phn ng. Theo Gunberd v Vaag th tc c ca phn ng: A = M+ N c xc nh bng cng thc: dC =k.C v= dt v: Tc phn ng k: Hng s tc phn ng C: Nng cht A Du tr th hin s gim nng cht A theo thi gian din ra phn ng. Vi phn ng tng qut: aA + bB = mM + nN Tc phn ng c xc nh theo cng thc: (2.1) v = k. [A]aB]b [A] v [B] l nng cht A v nng cht B cn a v b l cc h s ca cht A v ca cht B cn k l hng s tc phn ng. Bc ca phn ng (n) l tng cc s m ca nng cc cht trong biu thc tnh tc phn ng (2.1), tc l n = a + b Da vo , ngi ta chia ra phn ng bc mt, phn ng bc hai, phn ng bc ba. V d 1: Phn ng:N2O = N2 + 1/2 O2 c v = k [N2O]l phn ng bc mt.
V d 2: Phn ng: 2HI=H2 + I2 c v = k [HI]2 l phn ng bc hai. V d 3: Phn ng: 2NO + H2 = N2O + H2O c v = k [NO]2 [H2] l phn ng bc ba.
II. Phn ng bc mt
Phn ng bc mt l phn ng: A k P Tc phn ng c xc nh theo cng thc: dC = kC (2.2) v = dt C: Nng cht A k: Hng s tc phn ng Gi thit vo thi im ban u trc khi phn ng din ra l to = 0, ng vi nng cht A l Co cn vo thi im t nng cht A l C. Phng trnh (2.2) c th vit di dng sau: dC = kdt C Ly tch phn 2 v ta c:
dC C = k.dt C0 0
C ln C C o t o
Nng C Co
= k.t lnC - lnCo = - kt
ln
C C =-k.t =e-kt Co Co
C = Co e-kt (2.3) Phng trnh (2.3) biu din di dng th: (hnh 2.1) T th th hin nng cht A gim theo thi gian sau khi phn ng xy ra v gim theo qui lut hm s m.
t Thi gian Hnh 2.1: S thay i nng cht A theo thi gian ca phn ng bc 1. O
III. Phn ng bc hai

Phn ng bc hai l phn ng: A+B k P Theo nh ngha, tc phn ng bc 2 c xc nh theo cng thc: dC dP = =k.Ca.Cb (2.4) v= dt dt Gi thit thi im ban u trc khi phn ng din ra l to=0, ng vi nng cht A l a cn nng cht B l b. Sau mt thi gian l t, phn ng xy ra, to thnh sn phm c nng l x, th tc phn ng c th vit di dng sau: dx v = = k(a-x) (b-x) (2.5) dt dx = k.dt Suy ra: (a x ) ( b x )
gii phng trnh trn, ly tch phn 2 v:
k.dt =
o
dx (a x ) ( b x ) o (2.6)
k.t =
(a x ) 1 . ln +C (b x ) (a b )
tm hng s C, da vo iu kin khi to=0 th x = 0 Thay vo (2.6) s c: a a b 1 k.0 = . ln +C C = -ln C = ln (a b ) a b b Thay hng s C vo phng trnh (2.6) s xc nh c hng s tc phn ng: b(a - x) 1 (2.7) . ln k= (a b).t a(b - x)
IV. Phn ng bc ba
Phn ng bc 3 l phn ng:
A + B + C k P Theo nh ngha, tc phn ng bc 3 c xc nh theo cng thc: dC dP = = k.Ca.Cb.Cc (2.8) v= dt dt
Gi thit thi im ban u trc khi phn ng din ra l to=0, ng vi nng cht A l a, nng B l b, nng cht C l c. Sau mt thi gian t phn ng xy ra, to thnh sn phm c nng l x, th tc phn ng c th vit di dng sau: dx v = = k(a-x) (b-x) (c-x) (2.9) dt dx = k.dt hoc: (a x ) ( b x ) (c x ) gii phng trnh trn, xt trng hp n gin: a = b = c Phng trnh khi s c dng: dx k.dt = (a x ) 3 Ly tch phn hai v phng trnh trn s c:
t x o
k.dt = (a x)3
o
dx
k.t =
1 1 1 2 +C 2 2 (a x ) a
(2.10)
tm hng s C, da vo iu kin khi to=0 th x=0 Thay vo (2.10): 1 1 1 2 +C C=0 2 a2 a Vy hng s tc ca phn ng bc 3 khi nng cc cht tham gia vo phn ng bng nhau, c xc nh theo cng thc: k.0 =
k= 1 1 1 2 2 2.t (a x ) a
(2.11)
V. Phn ng thun nghch

Phn ng thun nghch n gin nht c dng: A k1: Hng s tc phn ng thun k2: Hng s tc phn ng nghch
1 k
k2
Gi thit thi im bt u xy ra phn ng to=0, cht A c nng l a, sau thi gian t phn ng xy ra, to sn phm P c nng l x, khi nng cht A s l (a-x). Tc phn ng thun nghch c xc nh theo phng trnh: dC dx (2.12) v= = = k1 (a x) k 2 .x = v1 v2 dt dt v1=k1(a-x) l tc phn ng thun v2=k2.x l tc phn ng nghch Khi tc phn ng thun bng tc phn ng nghch (v1=v2) th tc ca qu trnh s bng khng (v=0) v h trng thi cn bng. Khi h trng thi cn bng, phng trnh (2.12) c vit nh sau: k1(a*-x*)-k2x*=0 (2.13) * a : Nng cht A khi h trng thi cn bng x*: Nng cht P khi h trng thi cn bng T (2.13) suy ra:
k 1 (a * x * ) (2.14) x* Thay k2 vo phng trnh (2.12) s c: k2 =
( a* x* ) dx .x = k1 (a x) k1 dt x* dx (ax * a * x ) .k1 = dt x*
t x o
k1 .dt =
x *dx (ax * a * x )
(2.15)
Ly tch phn hai v phng trnh (2.15) ta c:
k1dt = x
o
dx (ax * -a * x )
k 1 .t = k1.t =
x* x ln(ax * a * x ) o + C * a
x* ln ax * ln(ax * a * x ) + C (2.16) a* tm hng s C, da vo iu kin khi to=0 th x=0. Thay vo phng trnh (2.16) tnh c C=0
Khi hng s tc phn ng thun c xc nh theo cng thc: k1 = x* ax * ln a * t (ax * a * x ) (2.17)
Thay k1 vo phng trnh (2.14) s tnh c hng s tc ca phn ng nghch:

ax * (a * x * ) x * k2 = . * ln x* a t (ax * a * x)
k2 =
(a * x * ) ax * ln a *t (ax * a * x )
(2.18)
VI. Phn ng song song

Trong c th sng c nhiu cht tham gia vo cc phn ng song song. Th d nh glucose c th b oxy ha theo con ng oxy ha kh ca chu trnh Crebs hoc theo chu trnh hecxozamonophotphat. Phn ng song song n gin c dng: k1 A k2 C B
Gi thit thi im bt u xy ra phn ng to=0, cht A c nng l ao, sau thi gian t phn ng xy ra to ra sn phm B c nng l b v sn phm C c nng l c cn nng cht A khi l a. Tc phn ng c xc nh theo phng trnh: dc db = k1a v v 2 = = k 2a (2.19) v1 = dt dt v1: Tc phn ng cht A chuyn thnh cht B v2: Tc phn ng cht A chuyn thnh cht C Tc ca c qu trnh: da db dc + = k1.a + k 2 .a = (k1 + k 2 ).a (2.20) v= = dt dt dt Ly tch phn 2 v ta c:
da a = (k1 + k 2 ).dt ao o ln a = (k 1 + k 2 ).t a = a o .e ( k1 + k 2 ).t ao (2.21)
Thay a vo phng trnh (2.19) ta c: db = k 1 .a o .e ( k1 + k 2 ).t .dt v1 = dt db = k1. a o .e ( k1 + k 2 ).t .dt

Ly tch phn hai v ta c:
b t
db = k1.a o e
o o
(k1 k 2).t
.dt
b=
k 1 .a o . 1 e ( k1 + k 2 ) t (k1 + k 2 )
(2.22)
T phng trnh (2.19) suy ra: v1 db dc k1.a db k1 = = = k1.dc=k2.db v 2 dt dt k 2 .a dc k 2 Ly tch phn 2 v ta c:
(2.23)
k1dc = k 2 db k1c =k 2 .b c =
o o
k2 .b k1
(2.24)
Thay gi tr b t phng trnh (2.22) vo (2.24) tnh c nng cht C k .a k k c = 2 .b = 2 . 1 o . 1 e ( k1 + k 2 ).t k1 k1 (k1 + k 2 )
c=
k 2 .a o . 1 e ( k1 + k 2 ).t (k1 + k 2 )
(2.25)
T phng trnh (2.23) cho bit trong phn ng song song, nu hai phn ng bc 1 xy ra cng mt lc th t s hng s tc phn ng s bng t s nng sn phm c to nn. Trng hp nu c 1 trong 2 phn ng l phn ng bc khng (l phn ng c tc phn ng khng ph thuc vo nng cht tham gia vo phn ng) th phn ng bc 1 s c ngha quyt nh n tc gim nng cht A theo thi gian.
Trn c s Hinshenvut rt ra mt kt lun quan trng: "i vi c th sng khi nng c cht c nng thp th phn ng do men xc tc s xy ra theo phn ng bc 1 cn khi nng c cht bo ha th mt s phn ng do men xc tc tin trin theo phn ng bc khng cn mt s phn ng do men xc tc vn tin trin theo phn ng bc 1 v chnh phn ng bc 1 s xc nh tc ca ton b qu trnh". Th d: Trong c th thng xy ra phn ng song song. Axit hipyric Glicocol k1 Axit benzoic k2 Glucuronic Glucuronit Khi nng axit benzoic nh th c hai phn ng u tin trin theo phn ng bc mt. Song trn thc t k1>k2 nn phn ng ch yu xy ra theo hng axit benzoic kt hp vi glicocol to thnh axit hipuric, cho nn trong nc tiu ch pht hin c axit hipuric. Khi nng axit hipuric ln th phn ng chuyn axit benzoic thnh axit hipuric l phn ng bc khng. Do vy phn ln axit benzoic kt hp vi axit glucuronic to thnh glucuronit, nn xt nghim thy nng axit hipuric khng tng cn nng glucuronit tng ln rt nhanh.
VII. Phn ng ni tip

Phn ng ni tip c dng:
k2 A k1 B C Tc ca phn ng t cht A to thnh cht B c xc nh theo phng trnh ca phn ng bc nht: dCa v= = k1Ca (2.26) dt Gii phng trnh ny thu c kt qu phn phng trnh bc nht:
Ca = Co. e k1.t (2.27) Co: Nng cht A ti thi im ban u (to=0) Ca: Nng cht A ti thi im t Tc s to thnh cht B s bng hiu s gia tc hnh thnh cht B t cht A v tc phn hy cht B thnh cht C, xc nh theo phng trnh:
dC b = k1.C a k 2 .C b (2.28) dt Cb: Nng cht B thi im t gii phng trnh trn cn da vo iu kin bin: Khi to=0 th nng cht A l Co cn cht B cha c to thnh nn Cb=0. Nghim ca phng trnh (2.28) theo cch gii t phng trnh vi phn c tm di dng: v= Cb= 1.e k1.t + 2 .e k 2 .t (2.29) Trong 1, 2 l nhng h s cn phi xc nh. Ly o hm nghim Cb theo thi gian t ta c: dC b = 1.k1.e k1.t 2 .k 2 .e k 2 .t (2.30) dt dC b t (2.30) vo phng trnh (2.28) Thay Ca=Co. e k1.t , Cb t (2.29) v dt ta c: 1.k1e k1.t 2 .k 2 e k 2 .t = k1 C o .e k1.t k 2 1.e k1.t + 2 .e k 2 .t Gin c cn li: k1.1 = k1.Co k21 1 =
k1.Co k2 k1
Thay gi tr 1 vo phng trnh (2.29) s thu c: kC C b = 1 o .e k1 .t + 2 e k 2 .t k 2 k1 Khi to = 0 th Cb = 0 2 = k1C o 1 = 2 k 2 k1
Khi bit 1 v 2, thay vo (2.29) s thu c kt qu nng cht B ti thi im t. kC C b = 1 o e k1t e k 2t (2.31) k 2 k1
Tc hnh thnh cht C s c xc nh qua nng cht B v hng s k2 theo phng trnh: dC v = b = k 2Cb dt
dC c = k 2Cb (2.32) dt Cc: nng cht C thi im t Thay gi tr Cb phng trnh (2.31) vo phng trnh (2.32) ta c: dCc k1 .k 2 .Co k1t = .(e e k2t ) dt k 2 k1 hay v= dC c = k 1k 2 C o k I t kk C e dt 1 2 o e k 2t dt k 2 k1 k 2 k1 k1k 2 C o k1t k 1k 2 C o k t e dt k 2 k1 e 2 dt k 2 k1 o o
1 k 1t e 1 k1
t t
Ly tch phn 2 v ta c:
Cc o
dC c =
k 1t
e
o
dt =
e
o
k 2 t
dt =
1 k 2t e 1 k2
)
) ( )]
Kt qu nng cht C ti thi im t s c xc nh: Co Cc = k 2 1 e k1t k1 1 e k 2 t (2.33) k 2 k1 C Biu din cc phng trnh (2.27), (2.31), (2.33) v s thay i nng A cht A, B, C tham gia vo phn ng B ni tip theo thi gian trn th s c dng nh trn hnh 2.2.
[ (
Hnh 2.2: S thay i nng cht A, B, C theo thi gian
Xt mt s trng hp c th: - Nu k1 ln hn rt nhiu so vi k2 th giai on u tin trin nhanh gp bi giai on hai v c hai giai on u tin trin theo ng hc ca phn ng bc nht. Trong trng hp ny tc chung ca phn ng ni tip
c xc nh bng tc ca giai on hai v l giai on tin trin chm nht. - Nu k1 nh hn rt nhiu so vi k2 th cht B c to thnh s khng bn v b phn gii nhanh hnh thnh nn cht C (tc k2 ln). Theo qui nh ca phn ng ni tip, phn ng no c tc chm nht s quyt nh tc ca ton b qu trnh. Trong trng hp ny th k1 s quyt nh tc ca phn ng ni tip.
VIII. Phn ng vng

Phn ng vng l mt chui cc phn ng din ra theo chu trnh nh chu trnh Crebs, chu trnh Glioxilic, chu trnh Ocnitin... gi mt vai tr quan trng trong qu trnh trao i cht ca c th sng. Tc ca chu trnh ph thuc vo nng tng i ca cc cht tham gia vo tng giai on ca chu trnh. C th c kh nng iu chnh nng ca cc cht tham gia vo chu trnh nh vo c ch iu ha tc phn ng. Dng n gin nht ca phn ng vng l phn ng men - c cht (hnh 2.3). M M: Men C P C: C cht MC: Phc cht men - c cht MC P: Sn phm ca phn ng
Hnh 2.3. Phn ng men - c cht
IX. Phn ng bc khng

Phn ng bc khng l phn ng c tc khng thay i v tc phn ng khng ph thuc vo nng cht tham gia vo phn ng. Tc phn ng bc khng c xc nh theo phng trnh: dC v= = k o (2.34) dt Phn ng bc khng in hnh l phn ng men tin trin trong iu kin tha c cht. Trong iu kin , tt c cc phn t men u dng lin kt vi c cht v tc phn ng c xc nh bi tc ca phn ng phn hy phc hp men - c cht to thnh sn phm P. Nh vy, tc hnh thnh sn phm P s khng ph thuc vo nng c cht m ph thuc vo nng men tham gia xc tc cho phn ng. Phn ng bc khng l mt h thng gm nhiu phn ng. Trong c th nu c xy ra phn ng bc khng th phn ng to sn phm phi xy ra t nht qua hai giai on nh kiu phn ng men - c cht, xt trn.
X. Phn ng t xc tc
Phn ng t xc tc l phn ng to thnh sn phm v sn phm li ng vai tr l mt cht xc tc. V d phn ng t xc tc: A k B Tc phn ng t xc tc s ph thuc vo c nng cht A (k hiu Ca) v nng cht B (k hiu l Cb). dC b =k.CaCb (2.35) v= dt Khi sn phm ca phn ng tng th tc phn ng cng tng ln. Mt c trng ring ca phn ng t xc tc l trong khong thi gian tng i di, lng sn phm ca phn ng tng i nh nhng sau chuyn sang giai on tin trin cc nhanh ca phn ng. Trong c th phn ln phn ng chuyn t Proenzyme thnh enzyme l phn ng t xc tc. V d phn ng chuyn pepxinogen thnh pepxin l phn ng t xc tc.
XI. Phn ng dy chuyn

Phn ng dy chuyn l mt h thng cc phn ng v c s xc tc ca sn phm trung gian. iu kin c th xy ra phn ng dy chuyn l phi c cc trung tm hot ng u tin. Cc trung tm hot ng u tin thng l cc gc t do. Cc gc t do c cc in t khng c ghp i nn chng c kh nng tham gia vo phn ng ho hc rt cao do vy chng thng c thi gian sng rt ngn. Gc t do khi tham gia vo phn ng vi cc phn t ngoi to thnh sn phm cui cng cn c kh nng to ra sn phm trung gian l nhng gc t do mi v gc t do mi ny li tip tc tng tc vi cc phn t khc gy ra phn ng tip theo. Nng lng hot ha ca cc phn ng dy chuyn thng rt cao nn trong iu kin bnh thng rt kh xy ra. Di tc dng ca tia phng x s dn ti phn ng dy chuyn, chng hn nh qu trnh chiu x nc gy ra cc phn ng sau:
e hv+H2O H2O+H++OHO (gc t do)
+e hv+H2O H2O-OH-+HO (gc t do) OHo+OHOH2O2H2O+Oo (gc t do) v.v... * Phn ng dy chuyn khng ny nhnh Phn ng dy chuyn khng ny nhnh l phn ng khi mt gc t do tham gia vo phn ng mt i th mt gc t do mi xut hin. Nh vy,
lng gc t do vn gi nguyn cho ti khi xy ra s t mch. S t mch ng ngha vi phn ng dy chuyn chm dt v s khng cn kh nng to ra gc t do mi. C hai nguyn nhn chnh dn ti s t mch. Nguyn nhn th nht l do gc t do b thnh bnh phn ng hp th hoc gc t do tng tc vi tp cht lm chng khng c kh nng to ra gc t do mi. Nguyn nhn th hai l khi nng gc t do cao, chng s tng tc vi nhau nn dn ti s t mch. Phn ng to HCl l phn ng dy chuyn khng ny nhnh: Clo+H2HCl+Ho Ho+ClHCl+Clo v.v... Hai phn ng trn l mt mt xch ca phn ng dy chuyn. S mt xch c to ra do mt gc t do ban u c gi l chiu di ca mch. Nu gi l xc sut t mch, th xc sut tip tc mch s l 1-. Xc sut mch c n mt xch c tnh theo cng thc: Pn=(1-)n. (2.36) Chiu di trung bnh ca mch s bng: 1 l= (2.37) Chiu di ca mch ph thuc vo bn cht ca phn ng dy chuyn, iu kin tin trin ca phn ng nh bnh phn ng, nhit , p sut... Tc ca phn ng dy chuyn khng ny nhnh c xc nh theo cng thc: dn (2.38) =no-g.n dt no: S trung tm hot ng c to thnh g: Tc t mch n: Nng gc t do Nu tc to mch l wo th tc phn ng dy chuyn khng ny nhnh cng c xc nh theo cng thc: (2.39) w = wo.l l: Chiu di trung bnh ca mch. Qua cc cng thc trn cho thy tc phn ng dy chuyn khng ny nhnh ch yu ph thuc vo lng trung tm hot ng c to thnh (tc nng gc t do). Nu cc yu t bn ngoi lm tng lng gc t do nh cc cht b chiu sng hay b chiu x s lm tng lng gc t do dn ti tng tc ca phn ng dy chuyn. Ngc li, cc yu t lm
gim lng gc t do nh cc cht c ch s hnh thnh gc t do s lm gim tc phn ng dy chuyn. * Phn ng dy chuyn ny nhnh Phn ng dy chuyn ny nhnh l phn ng mi mch ca phn ng khi mt gc t do tham gia vo phn ng c th to ra hai hay nhiu gc t do mi. Phn ng xy ha hydro l mt phn ng dy chuyn ny nhnh: H2 + O2 Ho + H O o (to mch) 2
Ho + O2 OHo + Oo (ny nhnh) Oo + H2 OHo + Ho (tip tc mch phn ng) OHo + H2 H2O + Ho (tip tc mch phn ng) Gc t do Ho tng tc vi thnh bnh s lm ngt mch phn ng. Mt s trng hp qu trnh to trung tm hot ng c th khng phi do cc gc t do ban u trc tip gy nn m do sn phm ca phn ng dy chuyn to nn. V d: Ro+O2ROOo ROOo+RHROOH+Ro ROOHROo+OHo tng lng gc t do phn ng dy chuyn ny nhnh c xc nh qua phng trnh: dn =wo+.n (2.40) dt n: Nng gc t do wo: Tc to trung tm hot ng. =f-g, trong f l tc ny nhnh cn g l tc t mch. Nu f>g th nng gc t do s c xc nh theo cng thc: w n= o (2.41) gf Gii phng trnh (2.40) s thu c nghim: w n = o e t 1 (2.42)
Nu tch s .t rt ln hn 1 th nghim c tnh theo cng thc:
n=
w o t e
(2.43)
* c im ca phn ng dy chuyn Phn ng dy chuyn c cc c im sau: - Phn ng dy chuyn c thi gian tim n. Trong thi gian ny ch yu to trung tm hot ng u tin. - Phn ng dy chuyn c hai gii hn nng . gii hn nng gc t do thp th gc t do d tng tc vi thnh bnh hay vi phn t cht c ch nn phn ng khng tin trin c. gii hn nng gc t do qu cao th cc gc t do d tng tc vi nhau gy ra hin tng t mch nn lm cho tc phn ng tin trin chm li. - Tc phn ng dy chuyn ny nhnh khng tun theo nh lut Arrhenius. Khi nhit tng th tc phn ng dy chuyn ny nhnh tng gp bi so vi nh lut Arrenius. Taruxop theo c ch ca phn ng dy chuyn ny nhnh gii thch s tc dng ca tia phng x ln c th sng. Khi chiu x, nu xt v mt nng lng th c gi tr rt thp nhng li gy ra hiu ng sinh hc cao (gi l hiu ng nghch l nng lng). iu ny c gii thch l do tia bc x gy ra hin tng ion ha, to ra cc trung tm hot ng l nhng gc t do v chnh cc gc t do gy ra phn ng dy chuyn ny nhnh, dn ti hiu ng tn thng sinh vt b chiu x. Taruxop cng cho rng cc cht c tc dng hn ch tn thng do tia x gy ra (gi l cc cht bo v phng x) l do n ngn chn c phn ng dy chuyn ny nhnh. Phn ng min dch ngi khi tim khng nguyn, sau thi gian bnh (thng t 3 n 21 ngy) khng th cha xut hin trong mu, sau l giai on nng khng th tng theo hm s m, khng t l thun vi lng khng nguyn a vo c th. Theo E. Manuel, phn ng to khng th din ra theo kiu phn ng dy chuyn ny nhnh. V d khi tim vaccine phng bnh bch cu, a vo c th ngi ch 0,36mg nhng sau thi gian bnh, lng khng th trong mu sut hin ln gp 1 triu ln so vi lng khng nguyn. E. Manuel cng cho rng c th khi b nhim cc c t nh nc c ca rn, b cp, iperit, rixin... th phn ng to khng th xy ra theo kiu phn ng dy chuyn ny nhnh.
XII. Nhit v tc phn ng

Theo l thuyt ng hc ca cc qu trnh ho hc th tc phn ng bao gi cng ph thuc vo nng cht tham gia vo phn ng v ph thuc vo nhit . Khi nhit tng th tc phn ng cng tng. Ban
u c gii thch do tng s ln va chm ca cc nguyn t hay phn t tham gia vo phn ng ho hc. Trn c s tnh ton xc sut va chm, nu va chm no gia cc nguyn t hay phn t cng dn ti phn ng th tc phn ng s tng ln t 102 n 106 ln so vi tc thc t o c nu ta tng nhit trong khong t 0oC n 100oC. T cc nh nghin cu cho rng ch mt phn trong s cc va chm mi c kh nng to phn ng ho hc. Phn ng ch c th xy ra khi cc nguyn t hay phn t t c mt gi tr nng lng nht nh gi l nng lng hot ha. Nng lng hot ha l gi tr nng lng nh nht m nguyn t hay phn t cn phi t c mi c th tham gia vo phn ng. Cc nh nghin cu p dng hm phn b Maxwell - Boltzmann gii thch mi lin quan gia tc phn ng v nng lng hot ha. Maxwell - Boltzmann thit lp c cng thc : N =N. e (2.44) N : S phn t t gi tr nng lng bng hoc ln hn nng lng hot ha. N: Tng s phn t ban u trong mt n v th tch. E: Nng lng hot ha ca phn t v R l hng s kh T: Nhit tuyt i (nhit Kelvin) Cng thc (2.44) biu din trn th (hnh 2.4) gi l hm phn b mv 2 ). Boltzmann. th biu din s phn t N theo vn tc v (E= 2
*
N N
E RT
T1 <
T2
0 a)
Ehh
E0 b)
Ehh
Hnh 2.4: S phn b ca cc phn t theo nng lng
Qua th (hnh 2.4a) cho thy ch c mt s phn t c nng lng bng hoc ln hn gi tr nng lng hot ha (Eh) th mi c kh nng tham gia vo phn ng. Khi nhit tng ln (T2>T1) th ng cong phn
b Boltzmann dch chuyn v pha bn phi, tc l s lng phn t c EEh tng ln (phn gch ngang hnh 2.4b) v vy tc phn ng tng ln. Nhiu phn ng, nht l cc phn ng ha sinh, tuy c nng , nhit v nng lng hot ha ging nhau nhng tc phn ng vn khc nhau. Nh vy, khng phi mi va chm gia cc nguyn t hay phn t c nng lng bng hay ln hn nng lng hot ha u dn ti phn ng. iu ny c gii thch do tc phn ng cn lin quan ti cu trc hnh hc ca cc phn t va chm, gi l yu t lp th (k hiu l p).
XIII. S ph thuc ca tc phn ng ha sinh vo nhit

S ph thuc ca tc phn ng vo nhit c m t qua phng trnh Arrenius. K= p.z. e RT (2.45) K: Tc phn ng p: Yu t lp th Eh: Nng lng hot ha z: S ln va chm R: Hng s kh bng 1,987 KCal/M. T: Nhit tuyt i Phng trnh Arrenius c th vit di dng logarit: E lnK = ln(p.z) - h (2.46) RT E 1 ; a = h ; b = ln(p.z) t y = lnK ; x = T R Phng trnh (2.46) c th vit di dng: y = a.x + b (2.47) y l phng trnh bc nht, th ca n c dng ng thng (hnh 2.5). Da vo th xc nh c: Eh tg = Eh = R.tg R Bit tg s xc nh c nng lng hot ha.
Eh
lnK b
1/T
1 T Nng lng hot ha l i lng nh gi kh nng tham gia vo phn ng ca cc cht. nh gi s ph thuc ca tc phn ng vo nhit , i lng c trng cho s tng (hoc gim) tc phn ng khi tng (hoc gim) nhit l h s VantHoff (cn gi l h s nhit phn ng) k hiu l Q10. K (2.48) Q10= 2 K1
Hnh 2.5: S ph thuc ca lnK vo nhit
K2: Tc phn ng nhit T10 K1: Tc phn ng nhit T Gia nng lng hot ha v h s Q10 c mi lin quan vi nhau. Theo phng trnh Arrenius (2.45), thay K2=p.z. e phng trnh (2.48) ta c: Q10=pz. e Q10= e
Eh R .T1 Eh R .T2
Eh R .T2
v K1=pz. e
Eh R .T1
vo
/ pz. e =e
Eh R .T1
.e
Eh R .T2
Eh 1 1 R T1 T2
Q10 = e
E h T2 T1 R T1 .T2
thay T2 - T1 = 10 ta c:
Q10 = e
E h .10 RT1 .T2
lnQ10 =
10E h RT1 .T2
(2.49) Eh = 0,1 RT1T2. lnQ10 Thay R=1,987KCal/M. v i logarit c s e (ln) thnh logarit c s 10 (lg) ta c: (2.50) Eh=0,46 T1 T2 lg Q10 n v tnh nng lng hot ha l KCal/M. Nhng phn ng c bn cht khc nhau thng c nng lng hot ha khc nhau v nh vy h s Q10 cng khc nhau. Cc phn ng ho hc c nng lng hot ha khong vi chc KCal/M, thng c Q10 bng 2 n 3. Nng lng hot ha ca cc phn ng men khong t 4 n 12KCal/M, Q10 thng nh hn 2. Qu trnh khuych tn nng lng hot ha thp, Q10 xp x bng 1. c trng ca phn ng men l c nng lng hot ha thp nn cc phn t d dng tham gia vo phn ng. Nguyn nhn l do cc phn t men c vai tr lm gim nng lng hot ha ca cc phn t tham gia vo phn ng khi men kt hp vi phn t to phc cht trung gian, lm cho qu trnh phn hy phc cht din ra nhanh hn. V d phn ng phn hy ng saccharose bi axit c nng lng hot ha bng 25,6KCal/M nhng nu thy phn ng saccharose c s xc tc ca men amilase th nng lng hot ha gim xung cn 11 KCal/M. Nng lng hot ha ca a s cc qu trnh din ra trong h sinh vt thng c gi tr thuc vo mt trong ba gi tr l 8; 12; 18 KCal/M. Tuy vy, nng lng hot ha ca cc qu trnh bin tnh li c nng lng hot ha rt cao. V d nh phn ng ng t hemoglobin c nng lng hot ha ti 60 KCal/M hay phn ng tan huyt di tc dng ca nc nng c nng lng hot ha l 64 KCal/M v.v... i vi cc qu trnh sinh vt, ngoi gi tr nng lng hot ha, h s nhit Q10 cn c c trng bi gi tr nhit thch ng. Theo Klein, nhit thch ng l nhit qu trnh xy ra vi tc ln nht. Cn phn bit nhit thch ng vi khong nhit thch ng. Khong nhit thch ng l khong nhit m gi tr Q10 c gi tr khng i. Trong h sinh vt, khong nhit thch ng rng hay hp l ty thuc vo loi. Trong h sinh vt, ng vt n nhit c nhit thch ng khc nhau. V d: ngi l 37oC, ch l 39,2oC, g vt l 42,5oC, b cu 44oC v.v... i vi cc phn ng c s xc tc ca men th ng cong ph thuc ca tc phn ng vo nhit c dng nh trn hnh 2.6.
Tu
Hnh 2.6: S ph thuc ca tc phn ng c men xc tc vo nhit
ng cong trn hnh 2.6 l kt qu ca phn ng trao i ho hc v phn ng phn hy cu trc. Trong khong nhit (0<T<Tu) khi nhit tng th tc phn ng tng l do phn t men vn cn gi c hot tnh xc tc. Tu l nhit ti u. Phn ng din ra nhit ti u c tc phn ng ln nht. Khi nhit tng cao hn gi tr nhit ti u th cc phn t men b ph hy cu trc nn mt hot tnh xc tc, dn ti tc phn ng gim.
XIV. Phng php phc hot ha

Phng php phc hot ha l phng php l thuyt xc nh tc ca phn ng ha sinh n gin, da trn tnh cht ca cc nguyn t hay phn t tham gia vo phn ng v cc tham s nhit ng nh s thay i ca entropi, ca entanpi... C s ca phng php phc hot ha l da trn quan nim v s tn ti ca mt phc cht hot ha (cn gi l trng thi hot ha) v phn ng ch c th xy ra khi cc phn t trng thi hot ha va chm vi nhau. Trong phn ng, s thay i th nng ca cc cht tham gia trc v sau khi phn ng din ra c biu din trn hnh 2.7. Cc nguyn t hay phn t mun tham gia vo phn ng phi t mc nng lng bng hoc ln hn gi tr nng lng hot ha (hay nguyn t hoc phn t c hot ha). Phn t cht ban u (cht A) c gi tr nng lng Eo, nu xy ra va chm s khng dn ti phn ng. Phn t cht A mun tham gia vo phn ng th phi c hot ha, tc l cung cp thm cho n gi tr nng lng E1Eh. Nng lng E1 dng lm gim th nng tng tc (tc lm gim lc ht) gia cc nguyn t hay nhm nguyn t trong phn t lm gim nng lng lin kt bn trong phn t. Nu phn ng xy ra theo chiu thun th Eo l gi tr nng lng ca cht ban u, E1 l nng lng hot ha ca phn ng thun v E1>Eo.
A*
E1 E2 Eo
P
Hnh 2.7: S thay i nng lng ca cc cht tham gia vo phn ng
Khi cc phn t cht A c hot ha (A*) va chm vi nhau s dn ti phn ng to sn phm P c gi tr nng lng l E2. i vi phn ng thun E1<E2 nn E1-E2<0. Do vy, y l phn ng ta nhit. Nu phn ng din ra theo chiu nghch th cht P ban u phi c gi tr nng lng hot ha bng E2 mi dn ti phn ng to thnh sn phm A c gi tr nng lng l E1. y E2>E1 nn E2-E1>0 v phn ng nghch l phn ng thu nhit. Cc nh nghin cu cho rng gia cc phn t tham gia vo phn ng (cht A) v cc phn t trng thi hot ha (A*) lun c s cn bng. Phn ng xy ra nh sau: A A* P (sn phm) K* hoc: A (A+E) P (sn phm) E l enzyme cn (A+E) l phc hp enzym vi c cht A.thi v c xc nh bng t s nng phn t trng thi hot ha trn nng phn t trng thi ban u.
K* =
[A ]
*
[A]
hoc
K* =
[A + E] . [A][E]
(2.51)
[A]: Nng cht A trc phn ng [A*]: Nng cht A trng thi hot ha
[A+E]: Nng phc hp enzym vi c cht A [E]: Nng enzyme i lng K* lin quan mt thit ti s thay i nhit , nng lng t do, entropi ca phn ng to phc cht hot ha nn K* c xem nh hng s cn bng ca phn ng v c xc nh theo phng trnh: R.T.lnK* = T.S - H (2.52) R: Hng s kh T: Nhit tuyt i (oK) S: S thay i ca entropi (ni ln s thay i trt t ca h khi b hot ha) H: S thay i ca entanpi (ni ln lng lin kt cn phi ph v to phc hot ha) Phng trnh (2.52) c th vit li nh sau:
S H K*= e R . e R .T (2.53) lnK = R R.T Trong c hc thng k xc lp c mi quan h gia hng s cn bng gia hai trng thi l K* v hng s tc phn ng l K theo phng trnh: R.T K=K* (2.54) N.h N: S Avogadro v h l hng s Planck Thay gi tr K* t phng trnh (2.53) vo phng trnh (2.54) s c:
R.T R R.T .e .e (2.55) N .h So snh vi phng trnh Arrenius cng thc (2.45) K= K = p.z.e
Eh R .T
R.T S Thnh phn (p.z) tng ng vi thnh phn .e R v s thay i N.h
entanpi (H) tng ng vi gi tr nng lng hot ha (Eh). Trong c th sng, phn ln cc phn ng ha sinh xy ra u c s tham gia ca enzyme, tc l u tri qua giai on to phc cht hot ha nn c th p dng phng php phc hot ha.
Thuyt phc hot ha c th gii thch c phn ng bin tnh hay phn hy ca Protein. V d nu un nng dung dch Protein hay cho vo dung dch Protein cc cht nh ur, cn, axit... s lm bin tnh hay phn hy phn t Protein. Khi biu hin r nht l ha tan ca Protein b gim xung cn nht li tng ln. Phn ng bin tnh hay phn hy Protein bnh thng khng xy ra v c nng lng hot ha rt cao. Mt nguyn t hay phn t mun tham gia vo phn ng th chng phi c hot ha. Nng lng hot ha phn ln c dng vo lm thay i trt t v cu trc phn t (tc lin quan ti yu t thay i entropi) cn phn nh c dng vo ph v cc lin kt trong phn t (tc lin quan ti yu t thay i entanpi). V d: Trong phn ng phn hy pepxin c 45.000KCal/M dng thay i trt t v cu trc phn t pepxin cn 18.300KCal/M c dng vo ph v cc lin kt ca phn t pepxin.
XV. S iu ha tc phn ng trong c th sng

Qu trnh ng ha thc n t mi trng l mt qu trnh phc tp, phi tri qua nhiu giai on bin i v mi giai on li c c trng bi cc phn ng sinh ha. Qu trnh bin i t cht A thnh cht K cho t bo c th ng ha c, gi l qu trnh chuyn ha. V d: Qu trnh chuyn ha t Protein thnh axit amin, t Lipit thnh axit bo v glixerin, t Gluxit thnh glucose v.v... Trong c th sng, qu trnh chuyn ha khng din ra mt cch n l m gia cc cht tham gia vo qu trnh chuyn ha chng cn c mi lin h vi nhau, th hin trn hai mt c bn l: nguyn liu v nng lng. Mi lin quan v mt nguyn liu l kh nng chuyn ha mt cht ny thnh cht kia thng qua mt s sn phm trung gian chung. V d saccharide d dng chuyn thnh Lipit (lipide) thng qua hp cht trung gian l axetil Coenzym A. Mi lin quan v mt nng lng th hin khi phn gii mt hp cht no , gii phng ra nng lng v c tch ly vo ATP. Ngun ATP c dng hot ha phn t (i vi phn ng thu nng lng), cc qu trnh sinh tng hp cc cht trong c th nh s hot ha cc axit amin trong tng hp Protein v thc hin cng nh qu trnh co c v.v... C ch iu ha gia qu trnh oxy ha (gii phng ra nng lng) v qu trnh photphorin ha (tch ly nng lng) c xc nh qua t s nng ADP v photpho v c trn nng ATP. [ADP] + [P] (2.56) K= [ATP]
Trong c th sng, khng c mt cht no l khng tham gia vo mng li ca cc phn ng ha sinh. C th sng l mt h m nn mng li cc phn ng ha sinh cng l mt h thng m, lun c s xm nhp ca cc cht t mi trng bn ngoi vo qua con ng thc n, tip n l qu trnh chuyn ha cht dinh dng v sn phm cui cng ca qu trnh trao i cht s c thi hi ra mi trng bn ngoi. Trong c th, s chuyn ha ca mt cht no c th c thc hin bng nhiu con ng khc nhau. Hinhenvut a ra nguyn l: "Trong cng mt iu kin, con ng chuyn ha no c tc phn ng ln hn th con ng c ngha quan trng hn". V d: glucose-6-photphat chuyn thnh 3 photpho-glyxerin aldehit c th c thc hin qua con ng glicoliz hoc qua chu trnh pentozo photphat, ty thuc vo hiu sut ca qu trnh chuyn ha. Con ng no t tiu tn nng lng v c tc phn ng din ra nhanh hn s quyt nh hng chuyn ha ca vt cht. i vi h kn, qu trnh din ra theo mt chui cc phn ng ni tip:
K1 A B K 2 C... K n P Phn ng no c tc chm nht s quyt nh tc ca ton b qu trnh chuyn ha. i vi h thng sng (h m), qu trnh chuyn ha s khng ph thuc vo tc phn ng ca mt giai on no m ph thuc vo s tng quan gia cc hng s tc phn ng. Mt c tnh quan trng ca c th sng l kh nng duy tr trng thi cn bng dng. Khi c th trng thi cn bng dng th tt c cc qu trnh thm thu, chuyn ha v thi hi s b chnh cho nhau duy tr nng cht khng i theo thi gian. Sau y l m hnh n gin c trng cho h m ni chung v h thng sng ni ring: Mi trng Mng T bo Mng Mi trng
S Ngun
K1 K2 A K
K 1
K 3
K3
Z Sn phm d ha
dinh dng Khi t bo trng thi cn bng dng th nng cht A (Ca) v cht K(Ck) s khng i. Do vy, nng cht A v cht K phi tha mn phng trnh: dC a dC k = 0 v =0 (2.57) dt dt
Di tc ng ca yu t mi trng (chng hn khi tng nhit ) s lm cho hng s tc phn ng K2 tng ln dn ti nng cht A gim xung cn nng cht K tng ln. duy tr nng cht A v cht K khng i th qu trnh vn chuyn cht A vo trong t bo phi tng ln (tc K1 tng) v qu trnh thi hi cht K ra khi t bo cng phi tng ln (tc K3 tng). Khi chm dt tng nhit th t bo khng chuyn ngay v nhit ban u m qu trnh h nhit din ra t t nn cc hng s tc phn ng K1, K2, K3 cng gim t t. Do vy, nng cht A vn cn cao v nng cht K vn cn thp hn so vi trng thi dng ban u. chuyn nhanh v trng thi cn bng dng, bt buc qu trnh vn chuyn cht A ra khi t bo v qu trnh vn chuyn cht K vo trong t bo phi tng ln (tc c K-1 v K-3 u phi tng). duy tr trng thi cn bng dng (tc duy tr Ca v Ck khng i) th qu trnh vn chuyn cht A vo trong t bo v vn chuyn cht A ra khi t bo cng nh qu trnh vn chuyn cht K vo trong t bo v vn chuyn cht K ra khi t bo phi b chnh cho nhau nh gii thch trn. M hnh trn thuc phm vi nguyn l Lostalie, c trng cho h m: "Khi mt h ang trng thi cn bng, nu ta thay i mt trong ba yu t (nng ; p sut; th tch) th cn bng s chuyn dch theo chiu c tc dng ngc li s thay i ". Trong v d trn, khi tng nhit tc l thay i yu t nng A v cht K. Nu nng cht A gim xung th s vn chuyn cht A vo trong t bo phi tng ln, nhm duy tr nng cht A n nh cn nng cht K tng ln th s thi hi cht K ra khi t bo phi tng ln cng nhm duy tr nng cht K n nh. Mt v d in hnh v s duy tr nng glucose trong mu c th ngi s minh chng cho nguyn l Lostalie. C th ngi khe mnh (tc duy tr trng thi cn bng dng) th hm lng glucose trong mu lun c mt gi tr n nh. Nu c th thc hin cng (nh chy in kinh) hay b i th hm lng glucose trong mu s gim xung. duy tr trng thi dng, lp tc glucogen tch ly gan s chuyn ha thnh glucoza-1-photphat enzym glucoza - 6 enzym photphat glucose v thm vo trong mu b vo lng glucose s dng. Hoc c th thc hin cng nhng khng s dng glucose trong mu m s dng glucogen. Nh enzym photphorilaza xc tc m glucogen chuyn thnh glucoza - 6 - photphat v tip tc b oxy ha iu kin ym kh gii phng ra nng lng, cung cp cho c th thc hin cng. Ngc li, nu c th t thc hin cng hay dng nhiu ng dn ti lm tng hm lng ng trong mu th lp tc xy ra cc phn ng: glucose glucoza - 6 - photphat glucoza - 1 - photphat, sau
trng hp thnh glucogen v c tch ly gan. y nng glucoza - 6 - photphat gi vai tr iu khin tc ca qu trnh chuyn ha thnh glucogen tch ly gan hay thnh glucose thm vo trong mu. Qu trnh chuyn t trng thi cn bng dng ny sang trng thi cn bng dng khc l mt qu trnh bt thun nghch. l s gi. Khng ai chng li c s gi. S pht trin ca c th ch din ra theo chiu thun t tr gi m khng din ra theo chiu nghch t gi tr. Mt s thng s trng thi ca c th nh nhit , pH ca mu, hm lng ng, mui khong trong mu th hu nh khng thay i trong sut thi gian sng. Ngi ta gi l tnh cht cn bng ni tnh ca c th sng (hay gi l cn bng dng bn).
Chng 3
TNH THM CA T BO V M
I. Cc phng php nghin cu tnh thm
Tnh thm ca t bo v m l kh nng cho cc cht i qua mng t bo khng c tnh chn lc (vn chuyn th ng) hoc c tnh chn lc (vn chuyn ch ng). nghin cu tnh thm ca t bo, cc nh khoa hc thng s dng cc phng php sau: - Phng php th tch: Theo di th tch ca t bo trng thi sinh l bnh thng v th tch ca t bo khi dung dch nhc trng. Ty mc ch v iu kin thit b m s dng cc phng php: * Ly tm huyn dch t bo sau xc nh th tch ca chng bng hng cu k. * Xc nh s thay i trong sut ca t bo bng phng php trc quang. *Xc nh s thay i chit xut ca t bo. Phng php th tch ch c th p dng nghin cu nhng i tng c kch thc ln v c bn cao trong dung dch nh to hay hng cu. V d nh theo di tnh thm ca m c ch i vi nc bng phng php trng lng (xem thc hnh L sinh). - Phng php s dng cht mu v cht ch th mu: phng php ny c tin hnh bng cch quan st trc tip di knh hin vi qu trnh tch lu cc cht c mu vo trong t bo. Cc cht ch th mu c s dng nghin cu tc thm ca cc loi axit v kim khc nhau vo trong t bo. Nhc im ca phng php ny l khi dng cc cht mu v ch th mu c nng thp th kh pht hin cn dng nng cao th chng tr thnh nhng c t i vi t bo. Khi s dng phng php ny cn lu cc cht c mu v ch th mu va ph thuc vo mc thm ca chng vo trong t bo li va ph thuc vo kh nng lin kt ca chng vi cc phn t Protein, axit amin v cc phn t khc trn mng hay trong t bo. V d nh s dng Xanhmetylen l mt cht c mu nghin cu tnh thm mt chiu ca da ch (xem thc hnh L sinh). - Phng php phn tch ho hc: y l phng php phn tch ho hc cc cht c nng rt nh trong t bo. Tuy nhin vi i tng nghin cu nh nh t bo li c thnh phn ho hc phc tp th vic nh lng mt cht no l rt kh khn. Nhng hin nay c nhng thit b phn tch hin i nh my quang ph hunh quang, my quang ph hp th nguyn t, my quang k ngn la... c th xc nh chnh xc nng cht cn nghin cu nng rt nh. Phng php phn tch ho hc c s dng rt rng ri trong nhiu lnh vc trong c nghin cu tnh thm ca t bo v m. V d nh xc nh nng natri (Na) bng my quang ph hp th nguyn t cho kt qu rt chnh xc. - Phng php dng cc cht ng v phng x nh du: Trong ho hc, ngoi nguyn t bnh thng nh Hydro (k hiu l 11H) cn c nguyn t phng x nh tritium (k hiu l 31H). Nguyn t tritium khc nguyn t Hydro ch trong ht nhn nguyn t c thm hai ht notron (n) khng mang in tch (gi l trung ho in). Cht phng x l n lun pht ra cc ht c mang in tch hoc cc ht khng mang in tch v chnh cc ht khi pht ra nh my o s xc nh c cht c gn cht phng x. Phng php s dng cht phng x nh du cho php nghin cu tnh thm ca t bo v m trong mt c th hon chnh (invivo) v cn dng nghin cu tin trin ca qu trnh chuyn ho
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com v o thi cht ra khi t bo. V d nh cc nh khoa hc s dng coban 58 phng x (27 Co) thay cho Coban bnh thng (5927Co) tng hp nn Vitamin B12, sau nghin cu qu trnh hp th B12 ca c th sng trong khong thi gian xc nh.
II. Mng t bo v vai tr ca mng t bo

Mng t bo ng mt vai tr v cng quan trng trong hot ng trao i cht ca t bo. Theo Frank: c th ni mt cch chc chn rng, cc chc nng quan trng nht ca t bo khng nhng b chi phi m cn ch c th thc hin c nh chc nng a dng ca cc loi mng. Mng t bo phi c cu trc p ng c yu cu ca qu trnh trao i vt cht v nng lng gia t bo vi mi trng, m bo thc hin cc chc nng sng ca t bo. c nhiu m hnh cu trc mng t bo c a ra nh m hnh DawsonDanielli (1935), m hnh Robertson (1959), m hnh Singer-Nicolson (1972). M hnh Singer-Nicolson vn k tha m hnh cu trc mng 3 lp: Protein - lp kp Lipit Protein do Rorbetson a ra nhng da trn cc kt qu nghin cu sau ny hai ng a ra m hnh mng khm lng (hnh 3.1) Theo cc tc gi, mng t bo l mt lp lipit kp trng thi lng cn cc phn t protein c nhng (khm) vo lp lipit vi mc nng hoc su khc nhau. Do trng thi lng nn cc phn t lipit v protein c th di chuyn theo c chiu dc ln chiu ngang. Tnh lng ca mng t bo ph thuc ch yu vo t l gia 3 thnh phn c bn tham gia vo cu trc mng l photpholipit, cholesteron, protein.
Hnh 3.1: M hnh cu trc mng t bo do Singer-Nicolson a ra nm 1972

Cc nh khoa hc cho rng duy tr trng thi bn lng ca lp lipit kp thuc mng t bo s c nhng ngha sinh hc nh: -Kh nng thch nghi ca t bo i vi nhit cao hay thp. -To s di ng d dng ca cc phc hp hocmn - th quan bn trong mng t bo trong vic tip nhn thng tin. -Tng cng s hot ng ca mt s enzyme vn chuyn cc cht qua mng t bo. -Gii thch c c ch vn chuyn cc ion qua mng t bo, c bit l Na+, K+ trong s hnh thnh in th tnh v in th hot ng.
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com -Gii thch c s bin mt ca mng nhn trung k v s xut hin mng nhn mt k trong phn bo nguyn nhim v gim nhim. Mng t bo c nhng chc nng c bn sau y: -Ngn cch gia cc t bo vi nhau, gia t bo vi mi trng bn ngoi v gi vai tr bo v t bo. -Thc hin qu trnh trao i vt cht v nng lng gia t bo vi mi trng, gp phn thc hin cc chc nng sng ca t bo. -Bi tit cc cht thi hoc cht c tng hp bn trong t bo ra mi trng bn ngoi. -Thc hin s nhn din t bo. Theo gi thit ca Ehrlich Tylerweiss th trn b mt mng t bo c nhng th quan nhn bit t bo theo kiu ging nh s nhn bit khng th vi khng nguyn. -Thc hin s kt dnh t bo. Cht kt dnh c bn cht glucoProtein c tng hp bn trong t bo v c tit ra mi trng bn ngoi di dng ho tan lm nhim v kt dnh t bo nh kiu trng cu gai tit ra cht ho tan trong nc bin c tc dng va dn d va to s kt dnh gia tinh trng vi trng. -Thc hin d dng s di chuyn ca t bo. V d nh Amip c th to chn gi di chuyn theo phng nm ngang c bn hng bt mi. -Thc hin s sinh sn ca t bo. Bnh thng t bo c ch s t l gia th tch nhn v th tch t bo cht (bng th tch ca t bo tr i th tch nhn) t mt gi tr hng nh th t bo khng phn chia. Khi ch s ny gim do th tch t bo cht tng ln th t bo phi thc hin qu trnh gin phn. -Mng t bo cn l ni c tr ca cc enzyme nht l cc enzyme tham gia vo qu trnh vn chuyn hay xc tc cho cc phn ng phn gii cht c lm tng tnh khng ca t bo. -Mng t bo c tnh bn thm nn c vai tr duy tr s bt i xng ion gia bn ngoi v bn trong t bo, hnh thnh nn in th tnh. -Mng t bo cn c tnh cht vt l nh tnh lng chit quang, sc cng mt ngoi nh, in tr ln v c cu trc khng ng nht...
III.Quy lut chung v s xm nhp ca vt cht vo trong t bo

Cc cht xm nhp vo trong t bo theo 2 con ng chnh sau y: -Cc cht xm nhp vo trong t bo qua siu l. Trn mng t bo c nhng l nh (siu l) c ng knh t 3,5 A0 - 8 A0. Bn trong siu l thng cha nc cn b mt c th cha mt s nhm phn cc. Con ng ny dnh cho cc cht ho tan trong nc nh ng, axit amin, ion... Cc cht ny mun xuyn qua mng qua siu l th phi vt qua c cc tr ngi nh phi tch ra khi cc lp v hydrat ho (nu l ion hay phn t c cha cc nhm phn cc), phi lch qua c lp phn t rt cht ch trn b mt mng t bo, phi thng c lc tng tc tnh in (lc ht nu l ion khc du v lc y tr li nu l ion cng du in tch vi cc nhm phn cc nm trn b mt ca siu l), phi vt qua c hng ro in th ( y l in th mng t bo). Do vy, cc cht mun xm nhp vo bn trong t bo th phi c mt gi tr nng lng nht nh thng c bn lc cn trn. Nu nng lng cn thit thng c bn lc cn trn cng ln th cng c t cht xm nhp c vo bn trong t bo.
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com -Cc cht xm nhp c vo bn trong t bo qua con ng ho tan trong lipit: Con ng ny dnh cho cc cht khng ho tan trong nc nhng ho tan tt trong lipit. Nm 1899, Overton khi nghin cu tnh thm ca t bo v m i vi cc cht gy m pht hin thy mi lin quan thun l tc thm ca cc cht ny cng cao khi kh nng ho tan ca chng trong lipit cng ln. Sau ny gi l hiu ng Overton. Jacobs gii thch hiu ng Overton theo cu trc in ca cc phn t. Theo Jacobs cc hp cht ho hc c phn thnh 2 nhm chnh theo tnh cht phn cc ca chng. Nhm th nht l nhng phn t c cha cc nhm phn cc nh NH2, COOH, OH... nn gi l cc phn t c cc. Nhm ny ho tan trong lipit km nn kh nng thm vo trong t bo t. Nhm th hai l nhng phn t cha cc nhm khng phn cc nh metyl, etyl, phenyl (CH3, C2H5, C6H5) nn gi l cc phn t khng phn cc. Nhm ny ho tan tt trong lipit nn kh nng thm vo bn trong t bo rt nhanh. Song cng c trng hp ngoi l nh ur v glucose ho tan km trong lipit v c kch thc ln hn siu l nhng li thm vo trong t bo rt nhanh. Ngc li, Citrat trimetyl ho tan tt trong Lipit nhng li rt kh thm vo bn trong t bo. iu ny c gii thch do ur v glucose l hai cht c mt trong qu trnh trao i cht ca t bo nn trn mng t bo c cht vn chuyn trung gian cn Citrat trimetyl th khng .
IV. Qu trnh vn chuyn th ng
Vn chuyn th ng l qu trnh xm nhp ca cc cht theo tng i s vct ca cc loi gradien v khng hao tn nng lng ca qu trnh trao i cht. Vn chuyn th ng din ra l do s tn ti ca cc loi gradien sau: -Gradien nng : L s chnh lch v nng gia bn trong v bn ngoi mng t bo. -Gradien p sut thm thu: L s chnh lch v p sut thm thu, c bit l p sut thm thu keo do cc phn t protein gy nn, gia bn trong v bn ngoi mng t bo. -Gradien mng: Xut hin do tnh bn thm ca mng. l do mng ch thm cc cht c kch thc nh nh ion, cc cht v c cn cc phn t c kch thc ln nh protein, lipit, gluxit th hon ton khng thm. Do vy gia hai pha ca mng c s chnh lch v nng to nn gradien mng. -Gradien ho tan: Xut hin trn ranh gii gia hai pha khng trn ln nh pha nc v pha lipit khi kh nng ho tan ca cc cht trong hai pha y l khc nhau, dn n s chnh lch v nng to nn gradien ho tan. -Gradien in th: Xut hin do s chnh lch v in th gia bn trong v bn ngoi mng t bo. Trong 5 loi gradien k trn th gradien no c gi tr tuyt i ln hn c s quyt nh hng vn chuyn ca dng vt cht. V d t bo hng cu, c, dy thn kinh, tn ti gradien mng c gi tr tuyt i ln hn c cho nn lng ion kali trong t bo lun cao gp t 30 n 50 ln so vi bn ngoi. c bit mt s loi to bin, nng it trong t bo cao gp hn hai triu ln so vi nc bin l do gradien mng. Cc gradien k trn u l hm s ca sinh l t bo v chng c lin quan vi nhau. Trong qu trnh hot ng sng ca t bo khng nhng ln ca cc gradien b thay i m c khi c hng ca chng cng b thay i. Cc gradien gi vai tr quan trng trong vic iu khin tc vn chuyn th ng cc cht vo trong t bo hoc i ra khi t bo.
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com Bn cnh vai tr ca 5 gradien k trn th hng vn chuyn ca dng vt cht cn ph thuc vo cng trao i cht ca t bo. Khi tng quan gia cc qu trnh tng hp v phn hu trong t bo thay i th hng vn chuyn ca dng vt cht cng b thay i. V d nhng t bo hng cu non thng xy ra qu trnh tch ly cc cht nn ion kali v photphat thng thm vo trong t bo vi cng ln. nhng t bo hng cu gi th nhu cu tch lu cc cht t cn qu trnh phn hy cc nucleotide din ra mnh nn ion kali v photphat li thi ra mi trng ngoi vi cng ln mc d vn tn ti gradien mng hng cu. Cui cng cn phi k n vai tr ca cc cht kch thch hoc gy thng tn i vi t bo. L thuyt hng phn ch ra rng ti nhng vng mng si trc noron, khi c sng hng phn truyn qua th tnh thm ca mng i vi ion tng ln. Khi dng thuc ph hy cc t bo ung th th gii phng gc photphat.
V. Qu trnh khuych tn v nh lut Fich

C ch vn chuyn ch yu ca cc cht ho tan trong nc qua mng l qu trnh khuych tn. Nu vt cht chuyn ng cng vi dng dung mi theo hng tng gradien, gi l qu trnh khuych tn thun. Trong trng hp ch tn ti gradien nng th vt cht vn chuyn theo hng gradien nng , gi l qu trnh khuych tn. Nm 1856, Fich tm ra nh lut khuych tn ca vt cht. Tc khuych tn ca vt cht trong mt n v thi gian t l thun vi gradien nng v din tch mng ni vt cht thm qua. Biu thc ton hc ca nh lut Fich: dm dC = D.S. (3.1) dt dx dm : tc khuych tn ca vt cht (gam/giy). dt D : h s khuych tn. i vi mi cht n l mt hng s. V d vi ng ma th D=0,384, vi ng Mantoza th D=0,373. n v ca h s khuych tn l Cm-1. s-1. S: Din tch b mt mng t bo ni vt cht thm qua (cm2). dC : gradien nng . dx Du tr v phi ca phng trnh (3.1) th hin s khuych tn ca vt cht theo chiu t ni c nng cao ti ni c nng thp lm cho s chnh lch v nng s gim dn. nh lut Fich c th c trnh by qua dng khuych tn ik, l lng vt cht thm qua mt n v din tch trong mt n v thi gian. dm ik = (3.2) S.dt So snh phng trnh (3.2) vi phng trnh (3.1) rt ra: dC i k = D (3.3) dx
dC khng thay i theo thi gian th dng khuych dx tn vt cht ik cng khng thay i theo thi gian, khi t bo trng thi cn bng dng. T phng trnh (3.1) cho thy vi mt cht xc nh th tc khuych tn trong mt dC n v thi gian ch cn ph thuc vo gradien nng . Trn thc t xc nh dx dy ca mng t bo l dx gp rt nhiu kh khn v dy mng thng thay i theo T (3.3) suy ra, nu gradien nng sinh l t bo nh hng cu dy ca mng thay i t 30 A n vi trm Angstron ( A ). Bi vy, hai tc gi l Berlien v Colender a ra phng trnh : dm = P.S(C1 C 2 ) (3.4) dt C1, C2 l nng cht hai pha ca mng t bo. P: Hng s thm ph thuc vo bn cht ca cc cht thm, c xc nh bng t s gia lng vt cht thm ra bn ngoi v lng vt cht c trong t bo. Vi ion Natri ( Na+ ) th P = [Na+] thm ra ngoi / [ Na+] trong t bo ([Na+] l nng ion Natri). Khuych tn l qu trnh vn chuyn th ng ca cc cht ho tan trong nuc nn nu xy ra c ch khuych tn lin hp tc l s vn chuyn ng thi hai cht cng mt lc nn hng s thm khng ch lin quan ti mt cht m cn lin quan ti cht khuych tn lin hp. V d s khuych tn ca Na+ c lin quan ti s khuych tn ca K+ nn hng s thm ph thuc c vo Na+ ln K+. Ngoi qu trnh khuych tn thng, s xm nhp ca vt cht vo trong t bo cn c thc hin theo c ch khuych tn trao i v khuych tn lin hp. -Khuych tn trao i : bng phng php ng v phng x nh du, cc nh khoa hc pht hin ra rng Na+ v cc ion khc hng cu thng c thay th bi chnh ion mi trng bn ngoi. Theo Uxing, qu trnh trao i Na+ bn trong v bn ngoi mng c s tham gia ca cht mang ( hay cht chuyn ). u tin cht mang lin kt vi Na+ trong ni bo sau vn chuyn ra bn ngoi mng. bn ngoi, Na+ trong t bo c gii phng cn Na+ c sn mi trng bn ngoi li lin kt vi cht mang v c vn chuyn vo trong ni bo. trong t bo, Na+ c ngun gc t mi trng li c gii phng cn cht mang lp li qu trnh trao i ion tip theo. Qu trnh khuych tn trao i Na+ vn m bo nng Na+ hai pha ca mng khng thay i. -Khuych tn lin hp : l qu trnh vn chuyn ca cht ny c lin quan ti qu trnh vn chuyn ca cht khc hoc l to phc cht vi mt cht khc. V d s vn chuyn ca Na+ ra bn ngoi t bo c lin quan ti s vn chuyn ca K+ vo trong t bo. Nu dng tc nhn ngoi c ch dng vn chuyn ra ca Na+ th dng vn chuyn vo ca K+ cng b c ch theo. Khuych tn lin hp theo kiu to phc cht c a ra gii thch s xm nhp ca glucose, glixerin, axit amin v mt s cht khc vo trong t bo. Glucose hu nh khng tan trong lipit v c kch thc ln hn kch thc siu l nhng glucose li thm qua mng rt nhanh. C ch thm ca glucose c gii thch do phn t glucose (k hiu l G) kt hp vi cht mang (k hiu l M) to thnh phc cht (k hiu l GM) v phc cht ny li ho tan tt trong lipit nn thm vo t bo rt nhanh. trong t bo
o o
glucose c gii phng cn cht mang trng thi t do s quay li nh v trn mng t bo, tham gia vo vn chuyn phn t glucose tip theo.
VI. Qu trnh vn chuyn tch cc

Vn chuyn tch cc l qu trnh vn chuyn cc cht ngc hng tng gradien v c tiu tn nng lng ca qu trnh trao i cht. trng thi sinh l bnh thng do mng t bo c tnh bn thm nn dn ti s phn b khng ng u ca mt s ion gia bn trong v bn ngoi mng. iu ny c th hin qua bng 3.1.
Bng 3.1: Nng ion trong t bo c v dch gian bo
Nng ion (n v l mili phn t gam) Dch gian bo Trong t bo
Na+ 145 12
K+ 4 115
Cl120 3.8
HCO327 8
Kt qu t bng 3.1 cho thy nng Na+ bn ngoi cao hn trong t bo trn 10 ln, nng K+ trong t bo li cao hn bn ngoi gn 30 ln cn nng Cl - bn ngoi cao hn trong t bo cng gn 30 ln. S phn b khng ng u ca mt s ion gia bn trong v bn ngoi mng t bo nhng i tng sinh vt khc nhau th khc nhau. Trong hot ng sng ca t bo, do nhu cu qu trnh trao i cht i hi phi vn chuyn ion v mt s cht ngc hng tng gradien nn phi cn ti qu trnh vn chuyn tch cc mi p ng c. V d : i vi nhng hng cu non do nhu cu tch lu cc cht cao nn K+ c thm vo trong t bo vi cng ln mc d phi vn chuyn ngc gradien nng . Song khi hng cu tr thnh mt t bo hon chnh th s vn chuyn tch cc ca K+ vo trong t bo s chm dt. Qu trnh vn chuyn tch cc i hi phi hao tn nng lng ca qu trnh trao i cht. Maidel v Haris chng minh rng nhit bnh thng ca c th khi trong mi trng c cht gluxit th cc ion dng c th vn chuyn ngc gradien nng . Nu dng thm axit monoit axtic hoc floritnatri th s vn chuyn tch cc cc ion dng s gim i r rt. iu ny c gii thch do qu trnh glicoliz phn gii gluxit thnh axit lactic v gii phng ra nng lng c tch lu vo 2 phn t ATP (phn gii ym kh) cn phn gii hiu kh s gii phng ra nng lng cao gp 20 ln phn gii ym kh. y chnh l ngun nng lng cung cp cho s vn chuyn tch cc cc ion dng. Khi thm axit monoit axtic hay florit natri, hai cht ny ph hu qu trnh glicoliz nn khng cn nng lng cung cp, v vy s vn chuyn tch cc cc ion dng b gim i. Qu trnh vn chuyn tch cc bao gi cng l s vn chuyn c tnh chn lc v ch din ra khi c nhu cu ca t bo. * Qu trnh vn chuyn ch ng cc ion dng Gi thuyt u tin a ra gii thch c ch vn chuyn cc ion dng l ca Hacxley v Convay, c a s cc nh nghin cu ng h . Hai ng cho rng trn mng phi c mt b my gi l bm Natri. Khi t bo trng thi tnh, bm Natri c kh nng bm Na+ t ni bo ra mi trng ngc gradien nng . Bm Natri hot ng thng qua h thng enzyme v s dng nng lng ca qu trnh trao i cht. Bm Natri thc cht l mt cht mang c sn trong mng hay c hnh thnh do s tng tc ca Na+ vi mt thnh phn no ca mng. Khi phc cht cht mang gn Na+" s xuyn qua mng ra mi trng ngoi v di tc dng ca enzyme th Na+ c gii phng cn cht mang trng thi t do li tr v nh v trn mng t bo thc hin
chu trnh vn chuyn Na+ tip theo. Sau ny Hodgkin, Katz v Scou (1954) u thng nht cho rng mng c mt b my gi l bm Natri - Kali. Bm ny c kh nng bm K+ t mi trng vo ni bo v bm Na+ t ni bo ra mi trng, khi t bo trng thi tnh. Ngun nng lng cung cp trc tip cho "bm Natri - Kali hot ng c ly t ATP. Cc tc gi cho rng bm Natri - Kali" thc cht l mt cht chuyn trung gian c sn trong mng hay c to thnh do nhu cu ca s vn chuyn tch cc ca ion. Cht chuyn trung gian c kh nng lin kt vi Na+ mt trong ca mng , sau chuyn Na+ ra mt ngoi ca mng t bo. mi trng ngoi, Na+ c tch ra cn K+ li lin kt vi cht chuyn v c a vo mt trong ca mng t bo. trong t bo, K+ c gii phng cn cht chuyn trng thi t do li thc hin chu trnh vn chuyn ion Na+ v K+ tip theo. Hodgkin tnh ton l thuyt cho rng nng lng thy phn 1mol ATP vn chuyn 1mol ion dng (cation) qua mng ngc gradien in th c gi tr khong 400mV. Thc nghim mi ch xc nh tnh ton ca Hodgkin ph hp vi s vn chuyn H+ qua mng d dy c gradien in th ln khong 400mV. noron thn kinh c gradien in th gn bng 120 mV nn nng lng thu phn 1ATP vn chuyn t 2 n 3Na+, ngc gradien in th. Thc nghim xc nh bm Natri Kali chnh l cht chuyn trung gian, nh v trn mng t bo (xem hnh 3.2). thc hin s vn chuyn ch ng Na+ v K+ th chnh hai ion ny hot ho enzyme ATPase xc tc cho phn ng thu phn ATP, gii phng ra nng lng cung cp cho qu trnh vn chuyn Na+ ra bn ngoi ng thi vn chuyn K+ vo trong t bo qua cht chuyn trung gian.
Hnh 3.2: S c ch hot ng ca bm Natri Kali (theo gi thuyt Hodgkin, Katz v Scou)
Kt qu nghin cu trn noron thn kinh v t bo thn xc nh do nhu cu cn c s vn chuyn ion ch ng, trong mng xut hin (hoc c trc) cht chuyn trung gian c bn cht l lipoProtein hoc photphoProtein. Qu trnh vn chuyn ch ng Na+ v K+ tri qua 3 giai on sau: - Giai on 1: Xy ra phn ng photphoril ha (tc chuyn gc photphat cho cht chuyn trung gian). Phn ng ch c th xy ra khi enzyme ATPase c hot ha bi Na+ xc tc cho phn ng thy phn ATP gii phng nng lng v chuyn gc photphat. Kt qu l Na+ v gc photphat c gn vo cht chuyn trung gian v phn ng xy ra bn trong t bo:
ATPase ATP + photphoprotein + Na+ Na+ - photphoprotein - P+ADP.
- Giai on 2: Phc cht Na+ - photphoprotein - P xuyn qua mng t bo ra mi trng ngoi. bn ngoi, xy ra phn ng trao i ion: Na+ - photphoprotein - P + K+ K+ - photphoprotein - P + Na+ - Giai on 3: Phc cht K+ - photphoprotein - P li xuyn qua mng vo trong ni bo. trong t bo, xy ra phn ng dephotphat (loi b gc photphat) v gii phng K+. K+ - photphoprotein - P K+ + photphoprotein + P Phn t photphoprotein trng thi t do li tip tc tham gia vo qu trnh vn chuyn ch ng Na+ v K+ khc. gii thch v sao cht chuyn trung gian li c th khi th gn vi Na+, khi li gn vi K+, cc nh khoa hc da vo thuyt Eidenman. Thuyt ny cho rng cht chuyn trung gian c in tch m, khi n thay i cc nhm mang in tch m s thay i lc ht tnh in. Do vy, cht chuyn trung gian c kh nng khi th "ht" Na+, khi li "ht" K+. - Gi thuyt Opit v Trernoc: Hai ng cho rng "bm Natri - Kali" thc cht l mt protein xuyn mng, l thnh phn cu trc ca mng t bo.
Hnh 3.3. S hot ng ca "bm Natri - Kali"

(Theo Bruce Alberts et all, 1994)
Protein xuyn mng c hai min A v B, trong min A c i lc (lc ht tnh in) i vi Na+ cn min B li c i lc i vi K+. Khi t bo c nhu cu vn chuyn ch ng ion th ATP tng tc vi protein xuyn mng dn ti c s gn Na+ vo min A ng thi khi chuyn gc photphat t ATP sang protein xuyn mng lm cho phn t protein xuyn mng thay i hnh th (hoc quay 180o). By gi c min A v min B quay ra pha ngoi mng v Na+ c gii phng ra mi trng ngoi bo cn K+ li c gn vo min B ca protein xuyn mng. Tip xy ra phn ng dephotphat (loi b gc photphat) v phn t protein xuyn mng tr v v tr lc ban u (hoc quay 180o). By gi min B li quay v pha trong mng v K+ li c gii phng ra t bo cht cn phn t protein xuyn mng li tip tc vng vn chuyn ch ng ion Na+ v K+ khc.
Cn theo Bruce Alberts et all (xem hnh 3.3) th protein xuyn mng l ATPase cng c min nhn Na+ v min nhn K+. Nh phn ng thy phn ATP m gc photphat t ATP c chuyn sang protein xuyn mng, lm cho phn t protein xuyn mng thay i hnh th (tc mt trong ca n m ra) cho Na+ gn vo min A. Sau mt trong ng li cn mt ngoi li m ra gii phng Na+ ng thi K+ li c gn vo min B. Tip theo ATPase loi b gc photphat tr v hnh th ban u (tc mt ngoi ng li cn mt trong m ra) gii phng K+ vo trong t bo. Phn t ATPase trng thi t do li tham gia vo qu trnh vn chuyn ion tip theo.
VII. Qu trnh vn chuyn cc cht hu c

Qu trnh vn chuyn ch ng cc cht hu c (c bit l ng v axit amin) n ph thuc vo cu trc mng t bo v cu trc phn t cht hu c. V d t bo c ch ch thm cc loi ng glucose, galactose, dng L-azabinose cn khng thm i vi dng D-azabinose (c nhm OH nm pha bn phi nguyn t cacbon ng lin sau nhm COOH). T bo gan li thm ng hecxose, pentose rt nhanh cn ng mantose li thm rt t. Qu trnh vn chuyn ch ng cc axit amin cng xy ra theo c ch tng t nh i vi ng. V d mng th riboxom thm glixin, dng D-alanin (nhm NH2 nhm pha bn phi nguyn t cacbon ng lin sau nhm COOH) v dng L-valin d dng, trong khi li hon ton khng thm i vi dng L-alanin. * C ch vn chuyn ch ng i vi ng: Cc nh khoa hc cng cho rng vn chuyn ng ngc gradien nng cng cn c s tham gia ca cht chuyn c trng, enzyme v ATP. iu ny c Heliman v Ianoy chng minh bng thc nghim vo nm 1971. Hai ng thy rng mng t bo bta ca tuyn ty ng vt c v c kh nng thm dng D-glucose ngc gradien nng cn hon ton khng thm dng L-glucose. Nhng nu thm vo mi trng nui cy t cht floritzin (l mt c t) th kh nng thm ca t bo bta tuyn ty i vi dng D-glucose cng b mt i. iu ny ch c th gii thch do floritzin ngn cn phn ng thy phn ATP cung cp nng lng hay c ch enzyme xc tc cho phn ng lin kt gia cht chuyn c trng vi D-glucose hoc ngn cn c hai phn ng trn. Bng chng tip theo l kt qu nghin cu ca Udas theo di s vn chuyn glucose qua mng nhau thai cu. Udas thy rng nu nng glucose mi trng ngoi nh hn gi tr nng gii hn, nu tng nng glucose ln th qu trnh vn chuyn glucose qua mng nhau thai cu cng tng theo. Song nu nng glucose mi trng ngoi tip tc tng khi ln hn gi tr nng gii hn th s vn chuyn glucose qua mng nhau thai cu khng tng thm na. T Udas cho rng phi c mt cht chuyn c trng i vi ng glucose. Khi nng glucose mi trng nh hn nng gii hn nu tng ln th vn cht chuyn c trng (vi quan nim mt cht chuyn gn vi mt phn t ng) nn s vn chuyn glucose qua mng nhau thai cu cng tng theo. Khi nng glucose mi trng ngoi tng cao vt qu nng gii hn do nng cht chuyn c trng khng tng thm na nn s vn chuyn glucose qua mng nhau thai cu s khng tng. Qu trnh vn chuyn ch ng ca ng cn theo c ch khuych tn lin hp. ng ha tan tt trong nc nhng khng ha tan trong lipit v c kch thc ln hn ng knh siu l nn khng th thm trc tip qua mng t bo. Khi phn t ng s to phc vi cht chuyn c trng (phn ln to phc vi enzyme). Phc cht ny khi li ha tan tt trong lipit nn khuych tn qua mng t bo mt cch d dng. trong t bo, phc cht b enzyme phn gii thnh cht chuyn c trng v phn t ng. Nh vy, phn t ng c vn chuyn vo trong t bo cn cht chuyn c trng li quay
tr li mng t bo tip tc qu trnh vn chuyn mi. ng cng c th c vn chuyn ch ng nh vo h thng enzyme vi s tham gia ca ATP. V d ng glucose bnh thng khng th thm ngc gradien nng . Nh enzyme hexokinase xc tc dn ti phn ng:
Glucose + ATP hexokinase glucozo - 6 - P + ADP Khi nng glucozo - 6 - P bn ngoi li cao hn bn trong t bo v ha tan trong Lipit nn d dng thm vo trong t bo. trong t bo, vi s xc tc ca enzyme glucozo - 6 -photphatase dn ti phn ng:
Glucozo - 6 - P G 6 e glucose photphatas Nh vy, nh hai enzyme vi s tham gia ca ATP m glucose c vn chuyn ngc gradien nng vo trong t bo. Qu trnh vn chuyn ng cn ph thuc vo pH v nhit mi trng. Theo cc tc gi Arnost, Ausiello, Denis (1970), s hp th ng xy ra mnh pH t 6,2 n 8,2. Tc thm cc loi ng qua mng lin quan ti nhit c gii thch bng phng trnh do Michalis v Menthen a ra dng gii thch s ph thuc ca tc phn ng men vo nhit . l: v .[ S ] v = max (3.5) [S ] + Km v: Tc phn ng vmax: tc phn ng cc i nhit thch ng [S]: Nng c cht Km: Hng s Michalis c gi tr t 10-1 n 10-6 p dng i vi ng: v .C v = max n (3.6) Cn + K v: Tc thm ca ng qua mng vmax: Tc thm cc i ca ng nhit thch ng Cn: Nng ng mi trng ngoi K: Hng s lin quan ti kh nng lin kt gia phn t ng vi cht chuyn c trng. mi i tng nghin cu, hng s K c mt gi tr xc nh. V d chut ng, vi ng glucose th K=9mM cn vi ng galactose th K=35mM. * C ch vn chuyn ch ng i vi axit amin: Cng ging nh con ng vn chuyn ch ng i vi ng, s vn chuyn ch ng cc axit amin cng c s tham gia ca cht chuyn c trng, ATP v enzyme. V d qu trnh ti hp thu axit amin ca t bo biu m ng ln ca thn, nh xc tc bi enzyme gamaglutamyltransferase m xy ra phn ng chuyn nhm glutamyl ca glutathion cho axit amin din ra nh sau: transferase axit amin + glutathion gamaglutamyltransferase - a.amin + xystenylglixin Sau gamaglutamyl - a.amin v xystenylglixxin u thm vo bo tng. Trong bo tng axit amin c tch ra cn gamaglutamyl li kt hp vi xystenylglixin to thnh
glutathion. Cc phn ng chuyn nhm, tch nhm v kt hp c s cung cp nng lng t 3ATP. Glutathion sau khi c ti lp li tip tc vng vn chuyn axit amin khc. Qu trnh thm cc axit amin ph thuc vo pH. Theo Lidonald (1971), tr glixin v alanin cn phn ln cc axit amin s hp thu pH=7 din ra mnh hn nhiu so vi pH=5 hoc pH=6. Qu trnh vn chuyn cc cht hu c c lin quan ti mt s ion, chng hn nh Na+. Thc nghim chng minh xtrophantin l cht c ch s vn chuyn Na+ ng thi xtrophantin cng c ch s vn chuyn cc loi ng, nucleotide, peptide, axit amin.
VIII. Tnh thm ca t bo i vi nc

Qu trnh thm nc ca t bo c thc hin nh cc yu t sau: - Do s chnh lch v p sut thm thu, c bit l p sut thm thu keo do cc i phn t sinh hc gy nn (protein, lipit...) - Do s chnh lch v p sut thy tnh. - Do s chnh lch v in th. - Do s thay i cu hnh phn t protein. C ch ny c trng cho cc loi sinh vt tin ha thp c c quan iu ha nc l nhng khng bo co bp do s thay i cu hnh ca cc phn t protein. Qu trnh thm nc vo trong t bo c thc hin theo nhng con ng ch yu sau: - S thm thu: Mng t bo c tnh cht bn thm, ch cho mt s cht i qua nh ion, nc, mui khong, ng, axit amin... cn cc cao phn t sinh hc th hon ton khng thm qua. Cc cht ha tan trong nc to nn p sut thm thu. Theo nh lut VantHoff, p sut thm thu P ca dung dch t l thun vi nng cht tan C (mol) v nhit tuyt i T (oK) theo cng thc: (3.7) P=.C.R.T : H s phn ly (i vi cht khng phn ly th =1) R: Hng s kh bng 0,082lt.atmtphe /M. Khi nng cht tan cng cao th p sut thm thu cng cng ln nhng do mng t bo c tnh bn thm nn cc phn t cht tan khng th khuych tn t ni c nng cht tan cao n ni c nng cht tan thp. Do vy, h bt nng cht tan ch c cc phn t nc s thm thu t ni c p sut thm thu thp (tc c nng cht tan thp) sang ni c p sut thm thu cao (tc c nng cht tan cao). Nh vy, s thm thu ca nc thc cht l qu trnh khuych tn nn c th p dng nh lut Fich xc nh tc khuych tn ca nc qua mt n v din tch mng, trong mt n v thi gian. T phng trnh (3.1) ca nh lut Fich cho bit tc khuych tn ca nc t l thun vi tit din S, tc l t l thun vi bnh phng bn knh siu l. - Qu trnh siu lc ca nc: Ngoi c ch khuych tn, s vn chuyn ca nc cn c thc hin theo c ch siu lc, do s chnh lch ca p sut thy tnh. Tc vn chuyn ca nc theo c ch siu lc tun theo nh lut Puazeli ging nh chuyn ng ca cht lng qua mao qun thy tinh do p lc t s chnh lch ca p sut thy tnh.
dv r 4 .( P P2 ) 1 = dt 8.l
(3.8)
dv : Tc chuyn ng ca nc qua mao qun (cm3/giy) dt r: Bn knh mao qun (cm) P: p sut thy tnh (P1>P2) : nht ca nc (Centipuaz: Cp) l: Chiu di mao qun (cm) Kch thc siu l c nh hng n s vn chuyn ca nc theo c c ch khuych tn ln siu lc. Ty thuc vo kch thc siu l m c ch no chim u th hn. Theo
Renkin v Papenhanner (1957), nu bn knh siu l bng 5 A th c ch khuyn tn v siu lc l tng ng nhau. Khi bn knh siu l ln hn 5 A , qu trnh siu lc s ln hn qu trnh khuych tn cn khi bn knh siu l nh hn 5 A th qu trnh siu lc li nh hn qu trnh khuych tn. V d in hnh v mi tng quan gia tnh thm khuych tn v siu lc l th nghim v s vn chuyn nc qua da ch. Trc th nghim, nc khuych tn qua da ch chim gn 356ml/cm2/gi cn thm siu lc chim 9,2ml/cm2/gi. Sau khi phun hoc mn tuyn yn l pituitrin ln da ch th nc khuych tn qua da ch ch tng ln t 12% n 13% trong khi c ch thm siu lc tng ln hn hai ln. Kt qu cho thy hoc mn tuyn yn lm tng ng knh ca siu l trn da ch. S thm thu v siu lc c ngha quan trng i vi qu trnh trao i nc gia mu v m. p sut thm thu ca mu ngi v ng vt c v do cc cht tan l v c v hu c to nn, c gi tr t 7,8 n 8 atmotphe (at). Song ch c p sut thm thu keo do cc phn t Protein to nn l gi vai tr i vi s trao i nc gia mu v m. p sut thm thu keo ca mu ngi bng 30mmHg cn p sut thm thu keo ca dch m v bch huyt bng 10mmHg. Nh nc s thm thu t dch m v bch huyt (ni c p sut thm thu keo thp) vo mu (ni c p sut thm thu keo cao). S trao i nc gia mu v bch huyt cn lin quan ti p sut thy tnh. p sut thy tnh c to ra t s co bp ca tim. cui ng mch, p sut thy tnh ln hn p sut thm thu keo, do nc c siu lc t mu (ni c p sut thy tnh cao) vo bch huyt v m lin kt (ni c p sut thy tnh thp). Ngc li, cui tnh mch, p sut thy tnh gim nn nh hn p sut thm thu keo, do nc li c thm thu t m lin kt v bch huyt (ni c p sut thm thu keo nh) vo mu (ni c p sut thm thu keo ln). phn gia ca ng mch v tnh mch c p sut thm thu keo v p sut thy tnh bng nhau. Khi c th khe mnh, s trao i nc din ra bnh thng th nc c vn chuyn t bch huyt vo mu cui tnh mch v t mu vo bch huyt cui ng mch s cn bng nhau nn m bo lng nc trong c th n nh. Theo quan im nhit ng hc th c th khe mnh s lun duy tr trng thi cn bng dng. Khi c th b bnh cao huyt p th cn bng gia p sut thm thu keo v p sut thy tnh s b ph v. bnh nhn cao huyt p s c s nhp p ca tim tng ln, do vy p sut thy tnh tng cao. bnh nhn h huyt p do mt mu nhiu, b nhim phng x, b i ko di... s lm p sut thm thu keo ca mu gim mnh. Do vy, bnh nhn cao huyt p v h huyt p u dn ti s trao i nc ca c th b ri lon.
o o o
Trong h sinh vt, do iu kin mi trng sng m c th c c quan iu ha p sut thm thu. sinh vt bc thp nh cc loi tim mao nc ngt c khng bo thng co bp tit ra dch nhc trng duy tr p sut thm thu cho t bo. Do vy, nc bn ngoi s khng thm thu vo trong t bo lm cho t bo khng b trng ln. cc loi tm nc ngt c tuyn di ru l c quan iu ha p sut thm thu v cng tit ra dch nhc trng (c nng mui rt nh hoc bng khng) cn bng vi p sut thm thu mi trng ngoi. c nc ngt cng nh c bin, c quan iu ha p sut thm thu l mang. Mang c nc ngt th tit ra dch nhc trng (ging tm nc ngt) cn mang c nc mn li tit ra dch u trng (c nng mui cao) cn bng vi p sut thm thu ca nc bin, gi cho c khi b mt nc. ngi v ng vt c v th c quan iu ha p sut thm thu l thn. Thn thi ra nc tiu nhc trng so vi mu khi c th n thc n lng (c th tha nc) cn thn thi ra nc tiu u trng khi c th n thc n kh (c th thiu nc).
IX. Tnh thm ca t bo v m i vi axit v kim

* i vi axit mnh v kim mnh: Trong dung dch axit mnh c c trng bi nng H+ cao cn kim mnh c trng bi nng OH- cao. Mc d kch thc ca ion nh hn kch thc siu l nhng cc ion khng th i qua siu l. iu ny c gii thch do trong dung dch cc ion bao gi cng b hydrat ha hay b cc ion khc du bao bc xung quanh. Ion c kch thc cng nh th lp v hydrat cng dy. V d K+ c
ng knh 2,6 A ch c 4 phn t nc bao quanh cn Na+ c ng knh 1,9 A li c ti 8 phn t nc bao quanh. Do vy, kch thc thc t ca ion trong dung dch ln hn rt nhiu so vi kch thc ca siu l. Mt khc, cc ion do b lp v hydrat bao bc nn hn ch kh nng tng tc tnh in vi cc ion trn thnh siu l. iu c bit l cc ion H+ v OH- u c kh nng tham gia vo cc phn ng ho hc rt cao cho nn chng d b cc phn t trn b mt mng hp ph. Chnh v vy t bo v m khng b tn thng bi axit hay kim th chng cng hon ton khng thm axit mnh v kim mnh. Axit mnh v kim mnh ch thm vo trong t bo khi t bo b tn thng do tc nhn vt l hay ho hc. Trakhochin tin hnh th nghim trn trng cu gai. ng tim vital khng mu (mt cht ch th gp axit s cho mu ) vo trng cu gai v sau th trng vo nc bin c nng H2SO4 thp. V H2SO4 l axit mnh nn trng cu gai hon ton khng thm nn trng vn c mu nh trng bnh thng. Song nu dng tia t ngoi chiu ln trng cu gai th trng s b tn thng nn H2SO4 d dng i vo t bo lm trng cu gai s c mu (do vital phn ng vi H2SO4 cho hp cht c mu ). * i vi axit yu v kim yu: Cc axit yu v kim yu c th thm qua mng t bo mt cch d dng v chng ha tan tt trong Lipit. Khi trong t bo cc axit yu v kim yu c th b phn ly thnh cc ion t khuych tn v li b hydrat ha nn chng khng thot ra ngoi mi trng c. l nguyn nhn dn n lm cho cc cht in phn yu thng c nng cao trong t bo. Hin tng trn gi l tnh thm mt chiu. Tnh thm ca t bo i vi axit yu v kim yu ph thuc vo pH v thnh phn ha hc ca mi trng. Nu pH ca mi trng dch chuyn v pha axit (tc giu H+) th phn ly ca axit yu s gim xung nn d xm nhp vo trong t bo. Nu pH ca mi trng dch chuyn v pha kim (tc giu OH-) th phn ly ca cc baz yu s gim xung nn cng d xm nhp vo trong t bo. Vn ny c Overton th nghim trn nng nc ch. Nng nc sng trong nc ngt, nu thm vo
nc mt t cht nitratstricnin nng 0,01% th nng nc vn sng bnh thng mc d ion stricnin rt c nhng n hon ton khng thm qua da nng nc c. Nu ta tip tc thm vo mt t cht cacbonat natri th nng nc s cht ngay. Nguyn nhn do ion stricnin kt hp vi ion cacbonat to thnh axit yu l stricnin cacbonat v d dng thm qua da nng nc nn lm cho nng nc b cht nhanh. Axit yu v kim yu c tnh thm mt chiu theo hng t mi trng vo trong t bo. Bn cht ca hin tng tnh thm mt chiu theo Rubinstein l do s khc nhau v gi tr ca cc tham s ha l mi trng v trong ni bo, do s gy nn s thay i trng thi ha l ca cc cht thm qua mng t bo. Th d mi trng bn ngoi phn t dng khng phn ly, khuych tn mnh (v d nh H2CO3) nhng khi thm vo trong t bo, phn t li phn ly (H2CO3 H+ + HCO3-) v khuych tn yu. Khi dng cc ion, chng s b lp v hidrat bao bc nn khng th thm ra mi trng ngoi c.
X. Thc bo v ung bo
Ngoi hai c ch vn chuyn th ng v vn chuyn ch ng, s xm nhp ca vt cht vo trong t bo cn theo c ch thc bo v ung bo. phn hy cc ht cht rn hay git cht lng khi chng b thc bo hay ung bo, trong t bo c bo quan ng vai tr nh mt bo quan tiu ha. chnh l lizoxom, mt bo quan bn trong c cha nhiu enzyme c th phn gii nhiu cht khi c t bo hp thu. * Thc bo: a s t bo trng thi t do nh sinh vt n bo (to) hay nm trong thnh phn ca m nh sinh vt a bo (m c, m gan, m rut non...) u c th thc bo. Thc bo l hin tng t bo c kh nng hp th cc ht nh vi khun, virus... Qu trnh thc bo thng xy ra hai giai on. Giai on th nht l t bo hp ph ht v gi cht ht trn b mt mng t bo. Giai on th hai l mng t bo un lm vo pha trong t bo cht bc ly ht cn a vo ni bo. Giai on hp ph ht ch yu lin quan ti cc yu t ha l nh in tch b mt hoc do tng tc ho hc. Giai on hai ch yu lin quan n mng t bo. C ch thc bo hp ph cht dinh dng hay gp nhng loi sinh vt tin ha thp nh vi sinh vt, to, ng vt nguyn sinh (amip). sinh vt tin ha cao nh ng vt c v v ngi, s thc bo c ngha chnh l bo v c th. ng vt c v th bch cu ht, t bo Kuffer c hot ng thc bo thng xuyn nh tiu dit vi khun, virus khi c th b nhim cng nh tiu dit t bo hng cu gi, mnh xc t bo b thng tn... * Ung bo: L hin tng t bo c kh nng ht nhng git cht lng nh git m vo trong ni bo. Hin tng ung bo c Maxt v Doil tin hnh th nghim v quan st trn i tng amip vo nm 1934. Hai ng s dng phng php nh du phn t protein bng cht pht quang nh vo mi trng gn con amip. Ch sau thi gian ngn quan st c c mt lng ln protein c con amip hp th theo c ch ung bo. Qu trnh ung bo cng din ra theo hai giai on. Giai on u tin l giai on hp ph git cht lng ln b mt mng t bo. Giai on th hai l mng t bo un lm vo pha trong t bo cht bc ly git cht lng, to khng bo a vo ni bo. Th d in hnh v ung bo l hot ng ca cc t bo ty xng. Trong vng ty to hng cu, c loi t bo ngoi bin cha y cc phn t pheritin. Quan st di knh hin vi thy r cc phn t pheritin thot ra khi t bo ngoi vi sau c hp ph ln b mt ca nguyn hng cu. Tip theo b mt ca nguyn hng cu un lm vo pha trong t bo cht to nn mt git nh bc ly phn t pheritin v a vo trong
t bo. trong t bo, mng bao bc phn t pheritin b ha tan cn phn t pheritin c s dng tng hp nn phn t hemoglobin. Kt qu nghin cu v s thm ht m t bo biu m rut non qua knh hin vi in t cho thy thc cht ca qu trnh ny l hin tng siu ung bo. Nu cho ng vt n m ng th ch sau t pht quan st thy nhng git m ng cc nh nm gia cc siu nhung mao ca t bo biu m rut non v mt s git m (lipit) nm gn trong vng un cong vo ca mng siu nhung mao. Thi gian cng v sau th s lng cc git m b hp th bi t bo biu m rut non cng nhiu. Qu trnh thc bo v ung bo cng l qu trnh hp th cht mt cch tch cc, i hi phi s dng nng lng song c tnh chn lc khng cao. Do vy, trong mt s trng hp t bo hp th ln b mt c nhng cht c, c hi i vi n.
Chng 4
IN TR CA T BO V M
I. in tr ca t bo v m.
Khi nghin cu tnh cht in ca h thng sng, cc qu trnh in sinh hc ca c th, ta cn phi nm vng cc kin thc vt l ng dung vo cc h thng sng. Ngy nay, vic ng dng cc hin tng in sinh vt kho st, thm d v iu tr trong Y hc l vic lm tng i ph bin. Xc nh cc thng s in ca t bo, m, si c... lm c s cho vic chn on v iu tr bnh. Vic tr liu bng phng php ny nhm gii thch s hot ng ca c th hoc h thng sng, trng thi trao i cht bnh thng cng nh phn tch trong cc tnh trng bnh l khc nhau. Nh vy bng phng php l sinh da trn hin tng in hc, khoa hc a ra c nhiu i lng c th, cc thng s trng thi c trng cho c th. Hn na, ngy nay nh vo cc thit b vi k thut hin i cung cp cho ta nhiu hng nghin cu mi trn t chc sng. Thc nghim cho thy rng, in tr hoc dn in ca cc i tng sinh vt c gi tr hng nh trong cng mt iu kin xc nh. l cc thng s c trng cn thit trong xt nghim cn lm sng, cc ch s ht sc c bn trong nghin cu y-sinh hc. V vy nghin cu tnh cht in nh o in dn, in th... ca cc t bo v m c mt ngha tht s quan trng. Vic nghin cu thng s in nhm mc ch tm hiu mt s c tnh, cu trc cng nh s nh hng ca cc tc nhn vt l ln cc c quan, t chc sinh vt. Vic xc nh gi tr in tr xut, in dn xut, tnh dn in hay ni chung l cc thng s in ca t bo v m cho php ta nh gi c trng thi sinh l v chc nng hot ng ca cc i tng kho st. Ngoi ra, t cc thng s ghi o c, ngi nghin cu cng c th suy on ra c cu trc ca cc t chc sng mt cch d dng. Bng nhiu phng php khc nhau ca l sinh hc, ta c th nhn bit v bn cht ca t chc sinh vt, m khng cn phi ph hu cu trc ca i tng kho st. Ngay t th u k th XIX, cc nh khoa hc bit cch chn phng php nghin cu ng theo hnh thc o in. Bng cch d dng

nht l o in tr cng nh xc nh dn in ca mu, t bo, m sng, c, si thn kinh. ng thi cng c th a ra c nhiu thng s lin quan m nhiu cng trnh nghin cu sau ny cp n. l cc ti nghin cu v in tr, in dn, tnh dn in trn cc ng vt, hoc ngay c trn cc t bo thc vt ... Nhiu kt qu thc nghim trc y chng minh cho ta thy rng: Vi cc t bo ng vt hoc thc vt cng nh i vi cc m sng, di tc ng ca dng mt chiu th in tr sut c gi tr trung bnh vo khong 106 107 /cm. Nn di tc dng ca dng in mt chiu cho thy, dn in ca cc i tng sinh vt gn ging nh tnh dn ca mt cht bn dn in. Cn di tc dng ca dng in xoay chiu th gi tr in dn o c c gi tr nh hn rt nhiu. xc nh c thng s v in tr thun, in tr khng ca cc h thng sng l mt vic lm khng n gin. Thng thng, ta gp phi nhng kh khn v phc tp trong khi o v: - i tng sng l mt h a pha v t chc khng ng nht v cu trc. - Th tch t bo khng c nh m c th bin i tu theo trng thi sinh l ca i tng khi kho st. - B mt t bo c mt lp v protin bao bc, lp mng bo v t bo c in dn rt ln. - Ngoi ra, dng in i vo m ch yu chy qua lp gian bo c dn in tt v bn thn n cha nhiu loi ion vi nng rt cao. - Cc vi in cc lm tn thng mng. Nhng s liu ng tin cy l o in tr ca cc t bo c kch thc nh vi hnh dng d tnh ton v th huyn ph. ph hp vi kt qu nghin cu t thc nghim, khi s dng cng thc Maxwell o thng s in tr. Phng trnh ny c dng nh sau:
r1 1 r = r1 +2 r r1 1 r2 r1 +2 r2
(4.1)
Trong , r : in tr sut ca dung dch. r1 : in tr sut ca mi trng.

r2 : in tr sut ca t bo. : l t s gia th tch ca t bo vi ton b th tch ca dung dch. Vi phng php xc nh nh trn, thc nghim cho ta thy rng in tr sut ca hng cu c gi tr khong 1012 /cm. Nh vy hng cu l mt vt th c c trng ca mt cht in mi. Ngy nay t cc kt qu nghin cu c, ngi ta a ra nhiu phng php khc nhau xc nh cc thng s in ca i tng kho st tu theo yu cu. Vic ng dng trong lm sng cng c mt ngha ht sc to ln. Ngoi ra, c s l thuyt cng c xy dng mt cch hon chnh, cng nh gii thch c cho nhiu c ch hnh thnh mt cch hp l vic ng dng hin tng in trong nghin cu y-sinh hc.
II. in tr ca t bo v m di nh hng dng in mt chiu.

xc nh gi tr v in tr thun trn c th ngi v ng vt, ta c th tin hnh theo mt s phng php n gin. o in tr bng cch dng in cc t ti hai v tr khc nhau trn i tng nghin cu, c m phng nh (hnh 4.1) di y:
Hnh 4.1: o in tr ti hai v tr xc nh ca c th.

Kt qu thc nghim cho ta thy rng: Vi cc t bo, m sng khc nhau th gi tr ghi nhn c cng khc nhau. Tht vy, khi cho dng in mt chiu i qua cc i tng nghin cu trn th cng dng in o c khng phi l mt hng s. Gi tr o c gim mt cch lin tc theo thi gian n mt gi tr xc nh no , mc d hiu in th ca ngun cung cp l khng i. Theo nh lut Ohm, cng dng in (I) s bin i mt cch tuyn tnh theo hiu s in th p t vo mch o. Vi ngun in mt chiu th cng dng (I) l khng i, nhng y kt qu ghi nhn c th hon ton c s khc bit. Tht vy, kt qu thc nghim cho ta thy rng dng in ghi o c trn i tng sng khng phi l hng s m gi tr ca n b thay i nhiu theo thi gian. Cng dng in l mt hm s bin i theo thi gian nh (hnh 4.2): I I = Const.
I = f (t) t
Hnh 4.2: S ph thuc cng dng in (I) theo thi gian (t).
Nguyn nhn c s gim dng in nh nu trn l do hin tng phn cc xy ra trong i tng nghin cu. Khi cho dng in mt chiu i qua, th ngay trn bn thn h thng sng xut hin mt sut in ng (E). Hin tng phn cc P(t) tng n mt gi tr xc nh no v pht trin theo chiu hng ngc li so vi s phn cc ban u. S phn cc ton phn ny lm hiu in th ton mch gim, do dng in ngoi cng gim theo. p dng nh lut OHM cho mt mch kn, trong trng hp ny ta c:
I=
U P(t ) R
(4.2)

Trong : U: l hiu in th p t vo i tng kho st I : l cng dng in qua on mch R : l in tr ca t bo, m sng m ta nghin cu. Tng t nh vy, trong h thng sng v c s phn cc khi cho dng in chy qua, nn ta cng c th xc nh c gi tr in dung C tng ng theo hng s in mi ca cc i tng nghin cu bng biu thc: C= Vi: : hng s in mi S : din tch b mt bn cc d : khong cch gia 2 bn cc. Trong t bo gm in dung tnh v in dung phn cc (Cp) ca cc loi t bo, m sng, c ...di tc dng ca dng in:
S 4d
(4.3)
Cp =
Trong :
Idt
0
(4.4)
R(I 0 I t )
I0, It : Cng dng in ti im t = 0 v ti thi im t bt k I : Cng dng in ca on mch. h thng sng, trong trng thi bnh thng th in dung phn cc c gi tr ln, cn khi i tng ny b tn thng hay hoi t th kt qu o c v in dung li c gi tr nh hn nhiu. in dung phn cc ca cc i tng sinh vt ni chung c gi tr trung bnh vo khong t 0,1 F/cm2 n 10 F/cm2 , cn i vi cc si c th in dung ca n c th ln n khong 40 F/cm2.

III. S bin i in tr theo tn s dng xoay chiu. Di tc dng ca dng in xoay chiu th tr khng ca t bo, m sng ph thuc nhiu vo tn s () ca ngun pht. Do , thnh phn in khng ca i tng sinh vt nh cm khng, dung khng v tng tr mch s ph thuc nhiu vo tn s ca ngun. Khi mt trng thi sinh l no , ng vi mt tn s nht nh ca ngun xoay chiu th in tr ca t bo v m sng c gi tr khng i. T nm 1920, nhiu cng trnh nghin cu kho st s bin i ca in tr di tc dng ca ngun xoay chiu cho thy: in tr ca hu ht cc loi m, c, t bo sng s b thay i di cc tn s khc nhau. c tuyn bin i ca in tr theo tn s () c dng nh (hnh 4.3) di y:
Hnh 4.3: S bin i in tr ca t bo m theo tn s.
T th trn ta thy: Vi < 1 v > 2 : in tr tng i n nh. 1 < < 2 : in tr gim khi tn s tng. Quy lut ny hon ton xy ra v bin i l nh nhau vi mi loi t bo, m v nhiu i tng khc. c bit, kt qu xy ra cng tng t i vi cc si c mc d gi tr tr khng tuyt i ca c l khc nhau. Cc gi tr 1 2 c gi tr thay i trong khong t 102 108 Hz.

a s cc m sng, in tr cc tiu hay tr s dn in cao ng vi tn s khong 106 Hz, cn i vi si thn kinh th gi tr ny tng ng vi tn s vo khong 109 Hz. T thc nghim cho thy, dn in ca t bo v m thay i ch yu tn s thp. tn s cao tnh cht ny t thay i hn v tn s thp hin tng phn cc din ra chm chp v r rt hn nhiu. S ph thuc dn in vo tn s l i lng c trng biu din cho i tng l t bo hoc m sng. i vi cc t bo, m b tn thng tu theo mc , tnh cht ny cng gim dn theo, n khi t bo cht th tnh cht cng khng cn na. Ngha l in tr ca t bo khi b tn thng khng cn ph thuc nhiu vo tn s. hiu r iu , ta kho st in tr ca mt s m thc vt trng thi sinh l bnh thng v b tn thng vi cc mc khc nhau di tc dng nhit, nh (h.4.4):
R (1 )
(2 )
(3
(1)
(4 )
Hnh 4.4: Bin i in tr m thc vt theo tn s trng thi sinh l bnh thng v khi b tn thng.
(1): Trng thi bnh thng. (2): un 500C trong 2 pht (3): un 500C trong 4 pht (4): un 1000C trong 20 pht.
T c trng bin i ca tng tr theo tn s ca dng in xoay chiu trn, ta thy ti nhit cao nu thi gian tc dng cng di th mc tn thng cng ln, ng thi s ph thuc ca in tr theo tn s t hn. Da trn c s nghin cu v cc thng s in c th d dng xc nh, nh gi cng nh m t i tng theo cc ch s sinh hc ca

mt t chc sng trng thi sinh l bnh thng hay b bnh l. Ngy nay ngi ta thng biu din ch s sinh hc qua c trng bin i ca in tr theo tn s R() cho cc loi t bo, m sng cng nh xc nh cho nhiu i tng nghin cu sinh vt khc.
IV. Tng tr ca t bo v m.
c trng nh lng cho mi lin h k trn, ng thi tin vic kho st, Tarutxop a ra h s K cho vic nh gi i tng nghin cu s nhanh v c ngha hn, thay v dng th biu din s bin i ca in tr nh trnh by trn. H s ny c xc nh nh sau:
k= R1 R 2
(2.5)
Trong 1 , 2 c chn tu theo i tng nghin cu. V du: Khi kho st trn mt dy thn kinh, thc nghim cho thy rng in tr c gi tr cc tiu tn s 2 = 109 Hz v 1 = 104 Hz nn:
K= R10 4 R109
(2.6)
Gi tr K cng ln s ph thuc ca in tr vo tn s ny cng mnh v ngc li gi tr ny nh khi t bo, m b cht. Trong cc t bo v m bnh thng, h s K ph thuc vo v tr ca i tng theo bc thang tin ho ca s sng. V d: H s K ca gan ng vt c v khong 9 10 H s K ca gan ng vt mu lnh khong 2 3 Cn trong c th ngi hoc ng vt, h s K ph thuc vo cng trao i cht ca tng loi m. - cc m sng c cng trao i cht mnh m nh gan, lch th h s K c gi tr ln. - Cn mt s c quan c cng trao i cht thp hn (c) th h s K li c gi tr nh. Cc gi tr R1, R2 c tin hnh trong cng nhit , iu kin th nghim nh nhau, v thi gian rt gn nhau. Nn t s o c tng i n nh khi m v t bo trng thi sinh l bnh thng ( tn s cao in tr ca c t thay i trong mt thi gian di, trong khi in tr vng tn s ngun in thp b thay i nhiu hn)

S thay i in tr lm cho tnh dn in ca t bo v m b thay i theo, ngun gc chnh l do kh nng xy ra hin tng phn cc ca chng. Do h s K cn c gi l h s phn cc
V. p dng phng php o in trong chn on v iu tr bnh.

1. Trong chn on v iu tr bnh. Ngy nay, phng php o in tr bng dng in th ng c s dng rng ri trong vic nghin cu qu trnh din bin xy ra t bo, m sng v cc t chc sinh hc di tc nhn ca cc yu t vt l, ho hc cng nh cc qu trnh bin i trong trng thi bnh l. V s dng vi ngun in p c gi tr nh, do tc nhn ny khng gy nn nhng bin i nhiu v tnh cht l ho trong t chc sng cng nh khng lm tn thng n i tng nghin cu l cc t chc sinh vt. Thc nghim cho ta thy rng, tnh trng bnh l bin i c lin quan nhiu n s thay i v gi tr in tr, in dn v cc thng s in in ca t bo. Khi mt t chc sinh hc b vim nhim th ngay trong giai on u, th tch t bo b trng ln, khong khng gian gia cc t bo b thu hp li, in tr ca m tng ln rt nhanh. Du hiu d dng thy nht l di tc dng ca dng in c tn s thp, in tr ca m lc ny c xc nh ch yu bi in tr ca cc thnh phn cht dch khong gian bo.
giai on u trong qu trnh vim, cc thnh phn v nht l cu trc t bo cha c s thay i ng k. Th tch t bo bin i khng ng k, nn gi tr in dung ca n cha c s thay i nhiu. Hin nhin rng: Khi tng gi tr in tr m in dung vn gi nguyn khng thay i th l du hiu ca s trng t bo, ngc li nu gim in tr m vn gi nguyn gi tr in dung th l du hiu ca s gim th tch ca t bo. giai on sau trong qa trnh vim nhim, ta thy c s thay i v cu trc t bo mt cch r rt, hin tng vim lm tng thm qua mng t bo. V vy lm thay i tnh dn in ca t bo v m sng, dn n s gim gi tr in dung v in tr. Ngy nay hin tng in sinh hc c ng dng nhiu cho vic chn on, iu tr ng dng trong y hc da vo cc tham s trn. Chng hn, p dng phng php o in tr xc nh nhn p, o chnh

lch in tr gia na mt bn phi v bn tri, o in c tai, kho st phn ng c bng cch o sut Galvanic. Xc nh mc tn thng t bo di cc tc nhn thng qua cc gi tr o in dn, o gi tr in tr thun v tr khng ca da...
2. Phn cc trong h thng sng.

Nhiu nh khoa hc cho rng, s phn cc cc ion ch yu xy ra trn b mt ca mng t bo. Quan nim phn cc mng xc nh l t bo sng ging nh mt vt th, m bn thn n c cha cc cht in phn trong cu trc thnh phn ca h. Cc cht in ly thng nm trng thi t do. Tht vy, khi trng thi sinh l bnh thng, t bo khng cho mt s cht i qua chng hn nh i vi ion Na+ m ch thm mt chiu i vi ion K+. Khi c in trng bn ngoi, cc ion trong t bo s di chuyn v cc cc in tng ng, tu thuc vo du ca in tch. Cui cng cc ion tri du s tp trung hai pha u cc ca mng t bo. Tng ng vi s tch t in tch ion pha bn trong mng, th mi trng bn ngoi mng t bo pha i din cng c s tch t cc in tch ion mang du ngc li. Nh vy, hai pha cc ca mng s xut hin hai lp in tch b mt. Hin tng to thnh trn t bo c m t ging nh hai bn cc ca mt t in v cng gi vai tr tch in trong cht in mi. y l gi thuyt a ra gii thch v s xut hin gi tr in dung ca mt t bo. in dung ph thuc vo hng s in mi ca t bo, cn gi tr in tr xut hin c xc nh thng qua in tr b mt ca n. Ngy nay bng phng php ng v phng x kim chng c iu . Mng t bo khng ch thm i vi ion K+ m cn thm ng thi i vi c nhiu loi ion khc nhau. Trong trng thi sinh l bnh thng hay qu trnh trao i cht bnh thng, ta thy lun lun xy ra hin tng vn chuyn cc loi ion Na+ - K+ qua li hai phi ca mng. Bng nhiu kt qu thc nghim khc nhau, ta d dng thy rng s phn cc trong t bo khng ch xut hin trn b mt ca mng m cn xy ra trong ton b th tch ca t bo. Tht vy, do hin tng in hng cc vt dn di tc dng ca in trng ngoi, lm cho cc in tch t do dch chuyn. Cc phn t tch in tng ng c s phn b in tch tr li trn pha i din trong cc t bo, nn cc lp bn trong t bo cng c s tch in vi du in tch ph hp.
P (t)
Hnh 4.5: S hnh thnh sc in ng phn cc.
T bo sinh vt c xem nh l mt h a pha, mi pha c tnh cht ho l khc nhau, do in tr v in dn ca n cng khc nhau. ng thi cc cht khi thm nhp vo t bo sng cng tn ti cc pha ho tan hon ton khc nhau. S phn cc xy ra trong ton b th tch ca t bo, ngay trn b mt v k c cc thnh phn bn trong t chc sng. c bit s phn cc trong ni bo ng mt vai tr ht sc quan trng. a s cc dch sinh vt trong h thng sng l cc dung dch in ly, cha cc ion v cc i phn t tch in. Do , c ch phn cc trong h thng sng c lin quan vi tnh cht ca mt dch a in phn trong dung dch in ly ca i a s cc phn t sinh vt. Trong c th, a s cc t chc u c cc mng ngn cch gia cc c phn vi nhau hoc cc t chc ring bit. Theo thc t c th c cu trc phc tp, ging nh bn thn chng c chia ra thnh nhiu ngn c lp. S dch chuyn ca cc ion qua cc vch ngn v th cng khng cn t do di tc dng ca in trng ngoi. Cc ion tch t trn t bo, m sng, trong cc t chc sinh vt to ra mt qu trnh phn cc in di tc dng ca in trng ngoi nh (hnh 4.6) di y:
Hnh 4.6: S cc tnh ho t chc sng. Ngy nay cc s liu thc nghim cho thy rng, qu trnh phn cc h thng sng c xc nh khng nhng bi s phn cc ca cc ion m cn c xc nh bi s phn cc (c tnh lng cc in) cc cht hu c. Cc phn t protein, nucleotid u c gi tr moment lng cc kh ln, hng s in mi ca dung dch in ly cng b thay i nhiu theo tn s dng in bn ngoi. i a s cc phn t nm trong thnh phn nguyn sinh cht u cha cc nhm phn cc, di tc dng ca cc yu t bn ngoi gy tn thng trong nguyn sinh cht. Do , trng thi thay i thng hnh thnh hng lot cu trc mi c s nh hng khc nhau. S phn cc in mi trong h thng sng cng c th do s dch chuyn in tch trong thnh phn cu trc, m bao gm s phn ln l cc phn t hu c. Qu trnh ny thng c gi l s phn cc in trong cu trc v m.
VI. Mt s phng php in di

in di l phng php phn tch da trn s dch chuyn cc in tch, phn t nhim in di tc dng ca trng lc in khng i. Cc thnh phn khc nhau trong dch sinh vt khi nhim in s mang mt in lng c ln v du in tch khc nhau. Tu theo loi phn t m cc ht nhim in s dch chuyn vi tc nhanh hay chm di tc dng ca in trng ngoi. Cc phn t sinh vt d dng nhim in v tch in, nn da vo tnh cht ny xy dng mt s phng php in di. Ngy nay, phng php in di c p dng rng ri v ng dng nhiu trong cc nghin cu y - sinh hc. Thng thng, ta s dng phng php in di tch chit cc thnh phn albumin, globulin trong huyt thanh; in di protein (phc cht lipide vi protein); in di glucoprotein (xc nh cc thnh phn ca protein v glucide trong huyt thanh).

Nm 1937, Tiselius l ngi u tin a ra phng php in di trong mi trng t do kho st s hin din ca protein trong huyt thanh. Sau Tiselius, Cremer ta thy c nhiu nh khoa hc khc nh Durrum xy dng l thuyt hon chnh v hin tng ny mt cch tng minh v pht hin thm nhiu ch tiu o t khc trong cc thit b mi ca mnh. Nm 1950, cc nh sn xut a ra mt loi thit b mi hn, l s phn tch da trn phng php in di hin th v kho st thng tin bnh l bng cch ghi li cc c tuyn bin i ca n trn bng giy ch th. Ngy nay c nhiu cng trnh nghin cu y-sinh hc p dng phng php in di, da trn nguyn tc dch chuyn ion tch chit v phn tch nhiu thnh phn khc nhau ca dch sinh vt. Cc hng nghin cu bng phng php in di thng tp trung trong ba loi ch yu l: - Pht hin s dch chuyn ca cc phn t bng cch kho st di knh hin vi (in di t bo). - in di trong dung dch t do (xc nh theo linh ng ca cc ht nhim in trong dung dch). - in di trn cc cht gi (phng php in di kho st trn bng giy).
Chng 5
IN SINH HC
I. M u:
T lu chu u, ngi ta tin hnh nhng th nghim l th khm ph v cc kh nng lm xut hin dng in trn c th ng vt. T khi nim v in ng vt mi xut hin v c chng minh s tn ti ca n. Mt s loi c sinh sng sng v bin c b phn bo v c bit pht in nh c tr in, c ui in, chnh in...Ngc dng lch s v s pht hin ra cc dng in t sinh vt trn cho thy, t rt lu ngi Ai Cp gp phi v lm quen vi nhng hin tng in ny. Mt tnh cht c trng ca t bo ng vt l gia chng v mi trng bn ngoi lun lun tn ti mt s chnh lch in th. o hiu in th trn cc loi t bo khc nhau th s chnh lch ny vo khong 0,1 V. c bit c mt s loi c in c th sinh ra cc xung in rt cao n khong 600V, vi dng in c hng trm mA. Tnh cht in sinh hc c Dr. Louis De Galvanie khm ph. Sau , ti ny thu ht nhiu nh khoa hc khc quan tm v u t vo vic nghin cu mt cch l th. Tuy nhin sau hn 100 nm, k t nhng pht hin u tin di s ghi nhn ca cc nh khoa hc, con ngi vn cha gii thch c c ch hnh thnh hin tng in sinh vt mt cch r rng. Cc kt qu thc nghim vn cn ng khung trong vic m t hin tng. Trong vi thp k gn y, nh cc phng tin ghi o c nhy cao, chnh xc, cng nh cc thit b in t hin i...ngi ta mi khm ph c nhiu qui lut hnh thnh dng in ca t bo. T kt qu thc nghim o c bng cc phng php khc nhau nh ng v phng x, ng hc phn t, hin vi in t, ho t bo..., cc nh khoa hc cho thy bn cht ca dng in sinh hc. Vic xy dng c s l thuyt v gii thch c ch ca vic hnh thnh dng in sinh hc cn c nhiu hn ch. S d nh vy l v khi nghin cu hin tng in sinh vt thng gp phi mt s gii hn sau: - Tc bin i tn hiu trn i tng nghin cu thay i qu nhanh, trong khi cc gi tr o c thng rt nh, nn yu cu v thit b nghin cu phi l cc dng c ghi o tht nhy v c chnh xc tht cao.

i tng nghin cu thng c kch thc ht sc nh (vo c kch thc t bo). iu kin nghin cu, phi c tin hnh vi phng php nh th no khng lm nh hng n trng thi sinh l ca i tng kho st.
Trc khi tm hiu v cc loi in th sinh vt, ta lu rng cc dch th hai pha trong v ngoi mng t bo l cc dung dch in phn (electrolytic solutions). Nng trung bnh ca cc anion c gi tr khong 155 mEq/l, ng thi c xut hin mt nng tng ng ca cc loi cation pht trin theo pha ngc li. Theo c ch vn chuyn vt cht qua mng sinh hc ta thy c s phn b tr li ca cc anion v cation hai pha mng. ng thi vi qu trnh vn chuyn tch cc, th c c s khuych tn ca cc ion vi cc thm khc nhau. Kt qu cui cng l trong ton b qu trnh h c s chnh lch nng ion hai pha mng, do lm xut hin mt hiu s in th mng (membranne potential). Hai yu t c bn c lin quan n s hnh thnh hiu th mng sinh hc c ngha quyt nh l: - S khuych tn nhng ion qua mng do s chnh lch nng ca cc loi ion hai pha mng. - S vn chuyn tch cc ca nhng ion qua mng khi chuyn dch t pha (phase) ny sang pha khc, to thnh mt cn bng mi l s cn bng c bit ca cc ion. Vi mt s c im nu trn th mc ch v yu cu khi nghin cu hin tng in sinh vt l: Hiu c bn cht ca cc loi in th sinh vt c bn nh loi in th ngh, in th tn thng, in th hot ng...Ngoi ra cn nm vng v cch ghi o, iu kin th nghim, cc giai on xut hin. Xy dng l thuyt ph hp gii thch s hnh thnh cc loi in th trn. Gii thch v cc kt qu ghi nhn c, k c cc mi quan h gia chng. Tm hiu mt s ng dng in sinh hc ca cc cng trnh nghin cu trong Y-Sinh hc. a ra mt s ng dng hin tng in trong cng tc chn on, thm d chc nng, cng nh cc ng dng iu tr bnh trong Y hc. Vic nghin cu cc hin tng in sinh vt v k thut ghi o cc thng s lin quan c mt ngha ht sc quan trng. c bit, ngy nay vi cc thit b khoa hc hin i, vic ng dng hin tng in

trong Y hc, xt nghim trn cn lm sng c s dng kh ph bin. Do ta cn phi nm k phng php ghi o, hiu r bn cht ca cc loi in th sinh vt c bn.
II. in th mng v in th pha. 1. in th pha.

* Gradient in th (the electric gradient) trng thi bnh thng ta thy c s phn b khng u ca cc ion hai pha ca mng sinh vt. Do c s chnh lch nng , cc ion ny s khuych tn qua mng t ni c nng cao n ni c nng thp hn. Di nh hng ca gradien nng , cc ion c khuynh hng tin ti trng thi cn bng mi, ng thi hnh thnh gia n mt lp in tch kp ngay trong dch sinh vt. nh hng ca in trng ngoi s lm xut hin gradien in th do s phn cc ca mng. Cc ion dng c xu hng ri mi trng c n pha m, ngc li ion m dch chuyn v mi trng c th dng hn. S chuyn di ca cc ht mang in s dng li khi lc tc dng ln cc ion di nh hng ca gradien in th cn bng vi gradien nng . Trong gradien nng pht trin theo hng ngc li so vi gradien in th nh (hnh 5.1) di y:
Hnh 5.1: nh hng ca gradien in th v gradien nng
: Dng chuyn dch di tc dng gradient in th : Dng chuyn dch di tc dng gradient nng

* in th in cc (electrode) in th in cc l hiu th c hnh thnh gia in cc v dung dch in phn. Hay ni cch khc l in th xut hin lp in kp khi in cc nhng vo dung dch in phn. V d: C th ly v d bng cch kho st trng hp hiu th ny khi s dng in cc bc (Ag) trong dung dch nitrat bc (AgNO3). Nu gi: ic l th ho hc ion kim loi trong in cc idd l th ho hc ca ion trong dung dch. Ta thy: - Nu ic < idd , di nh hng ca gradient th ho hc lm cc ion bc (Ag+) chuyn dch vo kim loi lm thanh kim loi tch in dng. Chuyn ng ca cc ion ny dng li khi h thng trng thi cn bng in ho. Thanh kim loi tch in dng v xung quanh s c mt lp ion NO3- bao bc to thnh lp in kp. trng thi cn bng in ho, s chnh lch in th ho hc gia in cc (c) v dung dch (dd) s c gi tr bng hiu s in th in ho ca lp in tch kp ny. Ta c: ZF = idd - ic (5.1) Trong : : in th ca in cc i vi dung dch Z : Ho tr ca cc ion t do F : Hng s Faraday - Nu ic > idd in cc s tch in m v xung quanh c lp ion Ag+ v cc ion Ag+ s ri khi thanh kim loi i vo dung dch. in cc b ho tan dn dn trong dung dch, hin tng chuyn dch ion ch dng li khi t ti trng thi cn bng in ho. - Nu ic = idd Trong trng hp ny s chuyn dch ca ion theo hai hng l cn bng nhau, nn th in cc bng khng. Da vo biu thc tnh cng thm thu phi thc hin khi tng nng dung dch ln 1gam/mol, ngha l lm thay i p sut thm thu ion t P1 n P2 tnh hiu th in cc. Ta c: P A = RT ln 2 (5.2) P1

Trong A: l cng thm thu R: Hng s kh l tng. T: Nhit tuyt i ca mi trng. Ngoi ra, cng ca lc in trong qu trnh chuyn ho lm thay i nng ion trong dung dch. Cng ny c xc nh trong trng th nng t nh in l: A = UF (5.3) Cng ca qu trnh in ho v cng thm thu trong mi qu trnh cn bng nhau, nn t (5.2) v (5.3) ta c: P2 (5.4) = UF P 1 RT P2 (5.5) U = ln F P 1 Theo phng trnh Vant Hoff th p sut thm thu t l thun vi nng nn: RT ln
U= RT C 2 ln F C1
(5.6)
Trong th nghim trn th C1, C2 chnh l nng ion trong dung dch (Cdd) v trong in cc (Cc). Vi Z l ho tr ca ion kim loi, nn tng qut ta c th vit li hiu in th ion nh sau:
U
i
C RT ln c ZF C dd
(5.7)
Vi Ui l hiu in th xut hin do s chnh lch in th ca cc ion to thnh. y chnh l phng trnh Nernst xc nh gi tr hiu s in th in ho (electrochemical potential different) ca cc dung dch in ly.
* Hiu in th pha. Nu nhng hai in cc cng kim loi vo hai bnh ng dung dch cha cng mt cht in ly vi ho tan C1 v C2 khc nhau. mi in cc, sau khi t n trng thi cn bng s xut hin mt in th m

ln ca n s t l vi t s nng cc ion kim loi trong in cc v trong dung dch . V nng ion kim loi trong hai dung dch l khc nhau nn gi tr in th xut hin trn mi in cc cng khc nhau, gia chng tn ti mt hiu in th Uc gi l hiu in th nng .
U c = U c1 U c2
RT C c RT C c Uc = ln ln ZF C1 ZF C 2
(5.8) (5.9) (5.10)
Uc =
RT C2 ln ZF C1
Vi C1, C2 l cc nng ion kim loi trong dung dch. Do , khi nghin cu trn thn nhit ngi bnh thng, nhanh chng cho vic tnh ton ngi ta thay vo cc gi tr hng nh, cng thc c xc nh li l:
UC =
C 61 log 2 Z C1
(5.11)
Nh trn ta thy di nh hng ca gradient nng v gradient in th m cc ion khuych tn qua mng vi tc khc nhau. Ni cch khc mi loi ion c d? linh ng khc nhau. Do s khuych tn ca mi loi ion khc nhau, nn gia hai lp dung dch c nng khc nhau th lp in tch kp xut hin cng c gi tr khc nhau. Hiu in th hnh thnh nn do s khych tn ca tng loi ion, to thnh cc pha ho tan trong mi dung dch. Hiu in th ny c gi l hiu in th pha khuych tn (UKT) c xc nh :
U KT =
RT + C 2 ln ZF + + C1
(5.12)
vi + , - l linh ng ca cc ion dng v ion m Hiu in th pha (khuych tn) tn ti trong mt thi gian rt ngn cho n khi c s phn b ng u tr li cc ion m v ion dng

trong ton b th tch dung dch. i vi dch sinh vt c dng l mt h in ly rt phc tp, nn vic ng dng cng thc ny vo h thng sng cng gp phi rt nhiu kh khn v tn ti nhiu i phn t ion ho khc nhau.
2. in th mng
* Cn bng Gibbs - Donnan Trong chng trc trnh by v trng thi cn bng GibbsDonnan, nn phn ny ta ch nhc n mt s tnh cht c lin quan v hin tng in m thi, l: -Trong c th ngi v ng vt c cc protein (R+) dng mui, n l cc i phn t khng lt c qua mng. Mc du cc phn t ny khng qua c mng nhng n ng mt vai tr ht sc quan trng, l lm nh hng nhiu n tc dng ca p sut thm thu ln mng. -Do s phn phi tr li cc ion khi trng thi cn bng ng c hnh thnh, nn hai pha mng c s chnh lch nng cc ion (c kh nng khuych tn c) qua mng. - Mt s ion khc cn li m khng c kh nng chuyn dch t pha ny n pha kia c, th s to thnh mt s cn bng c bit. chnh l cn bng Donnan. Thc nghim cho thy cn bng Gibbs-Donnan khng nhng ph thuc vo bn cht dung dch, tnh thm chn lc ion, kch thc ca mng m cn ph thuc nhiu vo loi in tch ca cc ion trong h sinh vt.
hiu r bn cht s phn b cc loi ion trong cn bng trn, ta kho st th nghim di y: Dung dch Protein cho vo bnh th nht RCl, c ion R+ l cc proteine mang in tch dng c kch thc ln khng qua c mng ngn cch gia hai bnh. Bnh th 2 cha dung dch mui NaCl, cc ion Na+ v Cl- c th d dng qua mng. Bnh th nht c nng C1, bnh th hai c nng C2. trng thi ban u cc ion c phn b trong mi bnh nh sau: [R+]1 = [Cl-]1 = C1 [Na+]2 = [Cl-]2 = C2 (5.13) C th biu din s phn b cc loi ion hai bnh (1) v (2) cha dung dch RCl v NaCl c nng ban u l (C1) v (C2) . Hai bnh (1) v (2) ngn cch nhau bng mt mng gia nh m hnh di y:
(RCl) (NaCl)
R+ Na+ ClCl(C2) (C1) (C2) (C1)
(1)
Hnh 5.2 : S phn b ion hai pha mng trong trng thi ban u.
Gi s sau mt thi gian ngn, s c mt lng ion Na+ v Cl- i qua mng vi nng X. Nh vy c s di chuyn ca Na+ v Cl- t bnh (2) sang bnh (1). By gi ta thy nng ca mi loi ion hai pha mng c phn b tr li trong qu trnh cn bng mi trng thi tip theo, c m t nh s di y:
(RCl) (NaCl)
R+ Na+ Na+ Cl(C2-X) (C1) (X) (C2-X) (C1+X) Cl-
(1)
Hnh 5.3 : S phn b ion hai pha mng trng thi cn bng mi trnh tnh trng tch t in tch dng v mt pha, nn s di chuyn thng xy ra mt cch ng thi vi c hai loi ion Na+ v Cl- . S dch chuyn ca cc ion ch tm dng li khi tin n trng thi cn bng : [Na+]1 [Cl-]1 = [Na+]2 [Cl-]2 (5.14)

Cn bng mi ny c gi l trng thi cn bng Donnan (Donnan equilibrium). c im ca trng thi ny l khi cn bng din ra, trong mi ngn u c s trung ho v in:
[R+]1 + [Na+]1 = [Cl-]1 [Na+]2 = [Cl-]2 (5.15)
trng thi sau l trng thi m h tin ti s cn bng ng. Khi trng thi cn bng xy ra, nu thay cc gi tr (5.15) vo phng trnh (5.14) ta c: (C2 - X )2 = (C1 + X ) X
2 C2 X = C1 + 2C 2
(5.16)
Kho st trong cc trng hp c bit, t phng trnh trn ta thy: *Nu nng ion R+ hay Cl- lc u rt b (C1 << C2 ), th c th xem nh nng C1 rt long so vi C2 (C1 0), nn c cht vn chuyn qua mng trong trng hp ny l: X = C2 / 2 (5.17)
*Nu ion R+ hay Cl- mi trng bn trong rt ln (C1 >>C2 ) so vi mi trng bn ngoi, th c cht khuych tn qua mng xem nh khng ng k: X 0 (5.18) *Nu nng dung dch ho tan gia hai mi trng cn bng nhau (C1 = C2 ) th : X = C2 / 3 (5.19) T cc trng hp trn ta c nhn xt l: Khi t bo tip xc vi dung dch cht in ly c cng loi v vi gc proteine l i phn t ion chnh ca t bo, th trong tt c cc trng hp u c mt lng cht nht nh i vo t bo. Di nh hng ca qu trnh vn chuyn lm cho p sut thm thu pha bn trong ca t bo lun lun c gi tr ln hn so vi mi trng xung quanh.
* Hiu in th trng thi cn bng Gibbs - Donnan th gia hai pha mng lun lun tn ti mt s chnh lch in th. Hiu in th xut hin l do c s

phn b khng ng u cc ion trng thi cn bng Donnan. Hiu in th c gi l hiu in th mng (Um). Theo th d trn ta thy, khi cn bng din ra th phng trnh cn bng Gibbs -Donnan cho ta: [Na+]1 [Cl-]1 = [Na+]2 [Cl-]2 C th vit li theo t s nng ion nh sau: [Cl ]1 [ Na + ] 2 = [Cl ] 2 [ Na + ]1
(5.20)
Lc , phng trnh hiu s in th in ho ca Nernst c vit li cho hiu in th mng l:
RT [ Na + ] 2 RT [Cl ]1 Um = = ln ln ZF [ Na + ]1 ZF [Cl ] 2
(5.21)
III. in th tnh.
Trong c th ng vt, trn cc t bo, m sng thng xut hin v tn ti nhiu loi in th khc nhau. Cc loi in th ny c cng ngun gc nh nhau nhng tu theo nguyn nhn xut hin, phng php o c v iu kin th nghim m ta c th phn chia ra thnh nhiu loi c tn gi khc nhau. l cc loi in th c bn nh in th ngh, in th tn thng, in th hot ng, in th ti ch. in th tnh hay cn gi l in th ngh. l in th c trng cho trng thi sinh l bnh thng ca i tng sinh vt. Ni cch khc, in th ny cng c trng cho tnh cht in ca h thng sng trng thi trao i cht bnh thng. in th tnh chnh l hiu in th bnh thng tn ti hai pha mng, c xc nh bng cch ghi o s chnh lch hiu th gia t bo cht v dch ngoi bo. C th tin hnh th nghim nh di y.
1. Th nghim. kho st s bin i dng in v o hiu in th mng ca mt t bo (m sng hay mt si thn kinh...) no , thng thng ta hay s dng phng php ghi o vi in cc ni bo. Th nghim c tin hnh nh hnh 5.4 (a,b,c) di y:
( ( b c Hnh 5.4: Ghi o in th ngh. a) t hai in cc pha ngoi mng sinh hc. b) t mt in cc bn ngoi v mt vi in cc xuyn qua mng. c) Cm hai vi in cc xuyn qua mng. Ghi o bng cch t hai in cc trn b mt si thn kinh, ta thy kim in k ng h o dng in khng lch khi im khng. iu chng t khng c s chnh lch in th gia chng (hnh 5.4a). Nu t mt in cc pha bn ngoi mng v mt vi in cc cm xuyn qua mng, ta thy gia hai in cc ny c xut hin mt hiu in th (hnh 5.4b). Cn khi chc c hai vi in cc xuyn qua mng th ta cng thy kim in k vn ch gi tr khng. iu chng t gia hai in cc khng c mt s chnh lch in th no. (Hnh 5.4c). Kt qu th nghim trn cho thy: Gia mt ngoi t bo khng b tn thng v mi trng bn ngoi khng c s chnh lch in th. Ngc li gia phn bn trong t bo v mi trng bn ngoi lun lun tn ti mt hiu in th no . S chnh lch in th ny c gi l in th ngh hay in th tnh ca mng (Resting membrane potential). 2. c im. in th ngh c hai c im nh sau: - Mt trong t bo sng lun lun c gi tr in th m so vi mt bn ngoi. Ni cch khc chiu in th ngh l khng i. - Bnh thng in th ngh c gi tr in th bin i rt chm theo thi gian. Bng cc phng php v k thut ghi o tt, ta c th duy tr dng in ny trong mt thi gian di. ln in th gim chm theo thi gian. Gi tr ny ch gim i khi chc nng ca t bo, hay ca si c bt u xut hin.
( a

3.Cc yu t nh hng n in th ngh. in th ngh c trng cho trng thi sinh l bnh thng ca h thng sng. Nu thay i trng thi sinh l s lin quan n trng thi chc nng ca h. Do bt k yu t no lm nh hng n qu trnh trao i cht bnh thng ca n cng u nh hng n in th ngh ca h, chng hn nh: - Di tc dng ca dng in bn ngoi. - Gi tr in th b thay i khi lm thay i thnh phn ion ca mi trng. - S tc ng ca mt s c t ln h thng sng cng lm bin i nhanh in th mng. - Khi thay i lng oxy trong mi trng cng s lin quan n qu trnh h hp ca m, c..., do s lm nh hng n in th ngh. - ...
cc loi t bo khc nhau th in th ngh cng c gi tr khc nhau. Gi tr ny thay i trong khong t -10mV n -100mV. S chnh lch in th tn ti gia cc phn khc nhau trong mt h sinh vt cng l mt trong nhng yu t c trng cho c th sng.
IV. in th tn thng.
in th tn thng l hiu in th xut hin do s chnh lch in th gia vng b tn thng v vng khng b tn thng. S tn thng xy ra c th do nhiu nguyn nhn khc nhau (nh di tc ng c hc, nhit, in, hoc ho hc ...) u lm xut hin s chnh lch in th. Loi in th ny c cng dng nh nhau trn cc i tng sinh vt. c trng c bn ca in th tn thng l: - Gi tr ca hiu in th gim dn v bin i chm theo thi gian. - in th tn thng ph thuc nhiu vo iu kin kho st v phng php ghi o. - ln in th b nh hng nhiu tu thuc vo iu kin sinh l ca cc i tng nghin cu.
1. i tng ng vt.
Thc nghim cho thy rng, trng thi sinh l bnh thng th cc thnh phn ion mt trong mng t bo (m, c...) v pha bn ngoi c s phn b n nh. Cn gia cc v tr khc nhau mi trng bn ngoi khng b tn thng so vi mi trng xung quanh s khng c s

chnh lch no v in th. Ni cch khc, trng thi sinh l bnh thng ta thy c s phn b in tch ban u hai pha mng sinh hc. Nu khi cc t bo (m) b tn thng, s lm nh hng n qu trnh vn chuyn cht, m c th l s trao i cc cht qua mng t bo. Ni tm li, s tn thng i tng sng m c th nh t bo (m, c,..) lm thay i trng thi chc nng ca t bo hay s lm thay i trng thi sinh l bnh thng ca cc i tng nghin cu.
2. i tng thc vt.

Kho st tnh cht in trn i tng thc vt cng cho thy c nhiu im tng t nh ng vt, l: - C s chnh lch in th gia vng b tn thng v vng khng b tn thng. - in th tn thng c gi tr m. - in th ny tn ti trong mt khong thi gian ngn. - Gi tr in th gim nhanh theo thi gian v tu thuc vo iu kin th nghim, ph thuc vo khong cch gia cc vng kho st. - Kh nng xut hin in th ny ch khu tr ti vng b thng tn.
3. Cc yu t nh hng.
Thc nghim chng t rng, cc yu t no lm nh hng n qu trnh trao i cht bnh thng ca t bo v m u lm thay i gi tr in th tn thng nh: - nh hng ca nhit mi trng. - Thay i thnh phn mi trng, nht l i vi Oxy lin quan nhiu trong qu trnh trao i cht. - S tc ng ca cc trng lc in bn ngoi (in trng, t trng..) lin quan n s chuyn dch ca cc ion qua mng. - S tc ng ca cc c t vo mi trng c lin quan n s thay i iu kin sinh l bnh thng.
V. in th hot ng.
in th hot ng l s dao ng nhanh ca in th mng. Dao ng in mng xut hin trong cc t bo thn kinh, c, v mt s t bo khc khi c sng hng phn truyn qua. Do dng in lm xut hin in th ny cn c gi l dng in hng phn. Tt c t bo sng u c c tnh d b kch thch, tc l c kh nng chuyn t iu kin sinh l bnh thng trng thi tnh sang trng thi hot ho. Di nh hng

ca tc nhn kch thch no , t bo s d dng b thay i tnh cht ho l ca mng. Khi c sng hng phn truyn n, du hiu in tch hai pha mng t bo b o ngc hn li so vi gi tr in th ngh lc ban u. Hiu in th ny xut hin l do c s chnh lch v gi tr in th gia hai pha ca mng. Lc ny gi tr ca in th mt bn ngoi s m hn so vi in th mt bn trong ca n. xc nh in th hot ng, thng thng ta s dng cc k thut ghi o vi in cc ni bo. C th tin hnh kho st s xut hin in th hot ng bng hai phng php nh di y:
1. Phng php 2 pha.

C th tin hnh kho st trn si thn kinh c kch thch ti v tr (1), hai in cc t ti hai v tr (2) v (3) trn mt si thn kinh. Theo di s bin i gi tr in th ca chng qua mt in k G nhy ni gia hai in cc nh hnh (5.5):
U = 0 (a) (b) U = 60V
(1) (2)
(3) U = 0 (d)
(1) (2)
(3) U = 60V
(c)
(1) (2)
(3)
(1) (2)
(3)
Hnh 5.5: Ghi o in th hot ng hai pha a) Kch thch ti v tr (1). b) Sng hng phn truyn n v tr (2). c) Sng hng phn nm gia v tr (2) v (3).
d) Sng hng phn truyn n v tr (3).

Nu dng mt tc nhn no kch thch si thn kinh ti v tr (1); th theo quan nim c in s c mt sng hng phn mang in tch m truyn dc theo si thn kinh. - Khi sng kch thch lan truyn n v tr (2) th gia hai in cc t ti v tr (2) v (3) s xut hin mt gi tr hiu in th U no , khong 60 mV (hnh 5.5b) - Sng kch thch lan dn v v tr (3) th hiu in th ny gim dn v tin gn n gi tr in p bng khng (U = O mV) khi sng hng phn trong vng gia v tr (2) v (3). Khi sng kch thch tin ti v tr (3) th hiu in th gia hai cc bin i v pha in th m (hnh 5.5c). - Khi sng kch thch truyn n v tr (3) th in th m ny t gi tr in p ti hn (Uth) (Uth = -60 mV) nh hnh 5.5d. - Khi sng ri khi v tr (3) th hiu in th gia hai in cc tr v li gi tr U bng khng nh ban u. Theo di c tuyn bin i theo thi gian ta c dng in th hot ng nh (hnh 5.6):
U Ub
Ud
Hnh 5.6: c tuyn bin i ca in th hot ng hai pha theo thi gian
2. Phng php mt pha.

* Phng php ghi o. Phng php mt pha l phng php ghi o in th hot ng bng cch dng mt in cc t ti v tr (2) v mt vi in cc khc cm xuyn qua mng t v tr (3). Sau kch thch ti v tr (1) v kho st sng hng phn kch thch truyn dc theo i tng nghin cu (t bo, si c, ...) nh (hnh 5.7):

U = 60V (a) (b)
U = 0
(1) (2)
(3) U = 60V
(1) (2)
(3)
(c)
(1) (2)
(3)
Hnh 5.7: S ghi o in th hot ng mt pha trn si thn kinh. a) Kch thch ti v tr (1). b) Sng kch thch truyn n v tr (2). c) Sng kch thch truyn n v tr (3).
- Khi cha kch thch, gia in cc (2) v vi in cc (3), c xut hin mt s chnh lch in th, l in th ngh ca si thn kinh. in th ny c gi tr khong - 60mV n - 100mV. - Khi kch thch ti v tr (1), sng hng phn lan truyn n v tr (2) th hiu in th ny tng dn ln t gi tr in th m n gi tr khng. Hiu th ny tng nhanh v t ti gi tr cao nht ti in th khng (U = 0) khi sng hng phn n v tr (2) (hnh 5.7b). - Khi sng hng phn truyn t v tr (2) n (3) th hiu in th hot ng mt pha gim tr li v in th ngh nh lc u (-80mV) Vy in th hot ng mt pha chnh l s bin i nhanh chng ca in th ngh di tc dng ca mt tc nhn kch thch no . Dng in th hot ng mt pha bin i theo thi gian trong th nghim trn mt si thn kinh, c biu din nh (hnh 5.8):
U(mV ) 0 40 80
a c b Hnh 5.8: c tuyn bin i ca in th hot ng mt pha theo thi gian. * Cc giai on hnh thnh. Khong vi thp nin tr li y, nh cc thit b ghi o hin i, in th hot ng mt pha c biu din mt cch t m, chnh xc hn. S hnh thnh in th hot ng c chia ra lm nhiu giai on nh (hnh 5.9). o trn si thn trc khng l ca thn kinh c mc, ta thy in th ngh c gi tr khong -60mV phn nh ca in th hot ng c gi tr khong 50mV. U Kch (mV) thch 60 B 40 20 Cm vi
in cc
O 20
40 C A 60 80 Hnh 5.9: Cc giai on bin i ca in th hot ng

in th hot ng c cc giai on bin i l: + Giai on kh cc (Depolarization), on AA. Lc ny hiu in th hai pha mng bin i t gi tr in th ngh (U ngh) n im c in th bng khng (U = 0 mV) + Giai on qu kh cc, on ABB. Trong giai on ny hiu in th hai pha mng vt qu gi tr in th khng, tip tc bin i v pha c in th dng. + Giai on phn cc li (Repolarization), on BC. l giai on m hiu in th hai pha mng gim tr li v gi tr in th ngh. + Giai on qu phn cc, on CD. Giai on ny ng vi lc hiu in th hai pha mng c gi tr m hn in th ngh. Nu kch thch c cng ln ta nhn thy rng: - Trong thi gian xut hin pha ln (nhnh ln) in th mng vt qu gi tr in th khng, ta thy c s o cc ca in th mng. - Trong pha xung (nhnh xung), mng c s phn cc li. in th hot ng pha ny ph thuc vo khong cch gia hai in cc v ph thuc nhiu vo tc dn truyn hng phn. Cc nghin cu ca Erlange v Gatse chng minh rng: -in th hot ng ghi c t mt thn kinh l tng cc in th lan truyn trn cc si t c cu to nn si trc thn kinh (hnh 5.10)
U (mV) 10 0 50
0, 4
1, 2
2, 4
t (ms)
Hnh 5.10: in th hot ng ca t c v si thn kinh.
T c tuyn trn, tc gi gii thch cho ta thy rng:-C s tng ng gia gi tr in th hot ng ghi o c trn si ca thn kinh ca

mo (nt t) v in th hot ng xut hin ghi o c trn tng si t thn kinh tng hp nn si trc thn kinh (nt lin).
VI. Bn cht ca in th tnh v in th tn thng.

gii thch v c ch, bn cht v ngun gc ca cc loi in th sinh vt, ta da vo mt s gi thuyt, cc l thuyt ion cng nh mt s cch l gii khc ca cc nh khoa hc: C nhiu quan im khc nhau gii thch v s hnh thnh in th sinh hc. Tuy nhin l thuyt m ang c nhiu nh khoa hc chp nhn v c c s vng chc hn c, l L thuyt ion mng. Theo thuyt ny, trong qu trnh hnh thnh in th sinh vt th cc ion (c bit l cc ion Na+, K+, Cl-) trong dch ni bo v bn ngoi t bo ng vai tr quyt nh. Cho n nay l thuyt ny vn chim nhiu u th trong vic gii thch hin tng in sinh vt. Da vo l thuyt trn, ta c th gii thch v s hnh thnh cc loi in th sinh vt c bn. Trc khi gii thch c ch hnh thnh in th ngh v in th tn thng, ta kho st s phn b cc loi ion chnh nh hng n hiu th mng. trng thi bnh thng, c th xc nh c gi tr in th tnh tng ng vi s phn b nng ion hai pha mng. Chng hn nh s phn b ion trong t bo c Mamalian nh (bng 5.1):
Bng 5.1: Nng mt s loi ion trong t bo c Mamalian
Nng ion dch ngoi bo [ion]0 (M/cm3) Cations: Na+ 145 K+ 4 + -5 H 3,8.10 .. .. .. ion khc 5
Nng ion dch ni bo [ion]i (M/cm3) Cations Na+ 12 K+ 155 H+ 13.10-5 .. .. ..
[ion]0 / [ion]i
m (mV)
12,1 1/39 1/3,4
66 -97 -32

Anion ClHCO3.. .. .. Ion khc Anion ClHCO3.. .. .. A-
120 27 7
4 8 155
30 3,7
-90 -32
in th
-90
1/30
-90
1. Nhn xt: Kho st cc thnh phn tng t nh trn nhiu i tng nghin cu khc nhau, nh thn kinh ch, tim chut cng, c xng ch .. .., ta thy c s phn b khng ng u ca cc loi ion hai pha mng. c bit, i vi cc loi ion Na+, K+, Cl- cho thy t l gia cc ion ny thng l: - Ion K+ trong t bo ln hn bn ngoi khong vi chc ln. - Ion Na+ bn ngoi ln hn bn trong rt nhiu. - Ion Cl- bn ngoi ln hn bn trong khong 30 ln. 2. L thuyt ion mng Bernstein l ngi u tin cho rng, in th tnh l kt qu ca s phn b khng u cc ion hai pha mng t bo. trng thi tnh, mng khng thm ion Na+ v Cl- m ch cho cc ion K+ lt qua. Hin tng vn chuyn cc cht xy ra, do c s phn b khng u c ba loi ion ny hai pha mng t bo. Ngoi ra mng c tnh bn thm v tnh thm ca mng i vi tng loi ion l khc nhau, l yu t c bn to nn in th tnh. in th tnh c cc gi tr khc nhau tu thuc vo i tng nghin cu.
Boyler v Conwey pht trin thm quan im ca Bernstein, bng cch chng minh cho ta thy rng mng thm ng thi i vi c ion K+ v Cl-. trng thi tnh, cc ion Na+, K+, Cl- c phn b tr li ti hai pha mng. Qu trnh vn chuyn v c ch hot ng ging nh s phn b ca cc ion trong trng thi cn bng Donnan. Do , in th tnh (US) cng c xc nh bng t s nng ca cc loi ion khuych tn qua mng. Hn na do tnh bn thm ca mng i vi tng loi ion m c s phn b li cc in tch ca chng hai pha mng. Thc nghim cho thy pha trong mng tch in m cn pha ngoi mng tch in dng.

Khi cn bng Donnan in th tnh ca mng t bo ng vt c th xc nh bi cng thc:
US =
Trong : [K+]0 v [Cl-]0 l cc nng ion mi trng bn ngoi t bo. [K+]i v [Cl-]i l cc nng ion pha bn trong t bo. Goldmann a ra gi thuyt l: - Mng t bo c tnh ng nht v cu trc v in trng tc dng ln mng ti mi v tr l khng i. - Dung dch in ly ca cc dch sinh vt c coi nh l dung dch l tng. - Mng c tnh cht bn thm nhng khng phi hon ton tuyt i, ngha l c th cho ion ny qua cn ion khc th khng th qua c. c trng cho kh nng dch chuyn ca cc ion qua c mng nhiu hay t, ta dng i lng h s thm (P) cho tng loi ion. - Cc ion Natri cng c tham gia vo qu trnh hnh thnh nn in th tnh ny. Do cng thc v in th tnh c Goldmann xc nh li nh sau: RT PK [ K + ] 0 + PNa [ Na + ] 0 + PCl [Cl ] i US = ln (5.23) F PK [ K + ] i + PNa [ Na + ] i + PCl [Cl ] 0 Trong : PK, PNa, PCl, l h s thm i vi cc ion Kali, Natri v Clo. Nhng nghin cu gn y xc nh v s hin din ca ion Natri tham gia vo qu trnh hnh thnh in th mng. Bng phng php ng v phng x nh du cho ta kt qu nh sau: -Mng t bo thm tt i vi ion K+ , Cl- v t thm i vi ion Na+ (tc dng ion Natri vo khong 14. 10-12 mol/ cm2 /sec). T cc kt qu thc nghim, Hodgkin v Keynes khng nh mt cch chc chn rng: -Mng t bo thm i vi ion Natri, mc du t nhng khng th b qua c. Tm li thuyt ion mng chim nhiu u th v vic gii thch bn cht s hnh thnh in th tnh. Kt qu o t thc nghim hon ton ph hp vi l thuyt. Thc vy, gi tr tnh ton t l thuyt gn ng vi kt qu o c t thc t. Tuy nhin l thuyt cng rc ri v cng khng
RT [ K + ]0 RT [Cl ]i = ln ln ZF [ K + ]i ZF [Cl ]0
(5.22)

th d dng gii thch trn nhiu i tng khi m iu kin th nghim kh phc tp. Tht vy l thuyt ion mng cho rng, cc ion u khuych tn qua mng di nh hng ca Gradient. Tuy nhin khi nghin cu trn mt s ng vt th cc ion Kali khng phi hon ton trng thi t do m c th cn lin kt vi cc cht bn trong t bo. Kt qu l thuyt v thc nghim khi kho st trn c ch theo ba phng php c xc nh nh (hnh 5.13).
UM (mV) 0 -20 -40 -60 -80 100 120 Kt qua thc nghim Tnh theo Goldmann Tnh theo Nernst
mM/ l 0, 1 5 5 10 50 100
[K+]O
Hnh 5.13: Gi tr in th tnh ca c ch theo cc php o
Nu lm thay i nng ion Kali mi trng bn ngoi c ch th in th tnh o c c s sai khc nhau t nhiu so vi cc kt qu tnh ton t l thuyt.
VII. Bn cht ca in th hot ng.

Tt c t bo sng u c c tnh d b kch thch, ngha l c nhiu kh nng chuyn t trng thi tnh sang trng thi hot ho di nh hng ca cc tc nhn. S bin i cc thng s c trng cho trng thi, thc ra l do b thay i tnh thm ca mng. Tuy nhin thut ng t bo d hng phn thng thng hay c s dng i vi cc loi t bo thn kinh, c.. ..ngha l cc i tng ny c kh nng p ng ngay di tc dng ca ngun kch thch. p ng thay i do kch thch thng c biu hin bng s xut hin

mt in th hot ng. V bn cht v c ch hnh thnh in th kh phc tp, da vo l thuyt ion mng ta mi c th gii thch mt cch hp l nht.
1. S kh cc v ti phn cc. - Ta bit rng trng thi ngh, c s phn b cc loi ion hai pha mng lm cho bn trong mng tch in m v pha bn ngoi mng tch in dng. in th chnh l gi tr cu in th ngh ca t bo trng thi bnh thng (hnh 5.14a). - Khi mng t bo c kch thch th t bo trng thi hng phn. Theo Bernstein v mt s tc gi khc cho rng mng t bo thm vi mt s loi ion no . Khi tnh thm ca mng i vi nhng ion Na+ t nhin tng, th nhiu ion Na+ thm t ngoi vo pha trong mng, mang lng in tch dng vo pha trong. Trng thi ngh bnh thng bin mt, pha trong mng c gi tr in th dng hn so vi gi tr in th m lc bnh thng. S phn cc tr li trong lc ny c gi l in th bin i (reversal potention) v giai on ny c gi l giai on kh cc (hnh 5.14b).
a) Trng thi ngh
b)Trng thi kh cc
c)Trng thi phn cc li
Hnh 5.14: S phn b in tch cc giai on ca in th hot ng
- Ngay lp tc sau khi c s kh cc khong mt phn trm giy (milisecond), mng hu nh thm hon ton i vi cc ion Na+. Do mt cn bng ion th bm Na+ v K+ xut hin a ion Na+ quay tr li. V vy to thnh to s cn bng mi ca cc ion gia hai pha mng. S phn cc lc ca mng ging nh s phn b ion lc ban u, nn giai on ny c gi l giai on phn cc li (hnh 5.14c).
2. S thay i tnh thm ca mng. Theo Hodgkin hay Huxley th gia in th mng v nhng ion i qua mng c mi lin quan vi nhau. Cc ion i qua mng tu thuc vo tnh thm ca mng, nn da vo tnh cht ny i vi cc loi ion hoc ni mt cch khc l c th da vo s thay i v dn in bi cc ion. in dn thay i lm in th mng (Um) cng thay i theo khi i tng sinh vt trong trng thi hot ng.
* Kho st th biu din dn in ca ion Na+ v ion K+ , tng ng nh khi kho st c trng tnh thm ca mng, ta c kt qu nh (hnh 5.15). Ta thy mi s kh cc mng u lm tng tnh thm ca mng i vi ion Na+ . Khi s kh cc t ti mt gi tr no (ngng kh cc) th tnh thm ca mng i vi ion Na+ t nhin tng vt ln. Tng ng khi dn in ca mng i vi cc ion Na+ cng tng ln hng ngn ln. S gia tng ny ch tm thi trong sut thi gian rt ngn, khong mt phn nh ca mili giy (ms). Cn i vi ion K+ ta thy lc mng trng thi ngh, dn in ca ion K+ ln gp khong 100 ln i vi ion Na+. Nhng trong giai on u hnh thnh in th hot ng, dn ion K+ ch tng ln khong 30 ln n 40 ln trong khi dn i vi ion Na+ li tng ln hng ngn ln. V tnh thm ion K+ xy ra tr hn v ko di trong mt thi gian lu hn so vi s gia tng ca ion Na+. Cc giai on bin i trong in th hot ng thng khng ng b nhau, nn ta phi kho st in th hot ng da trn s thay i v t s thm gia chng, ngha l da vo gi tr: PNa / PK Hoc tng ng vi s thay i v tnh thm, ta cng c th kho st s bin i ph thuc theo t s dn in: Na / K

60 40 in th 10 0 10 in th hoat ng 20 0 20 40 60 80 100
0, 1 0,0 1 0,00 1 10 0 10
Ty s dn
PNa PK
0, 1 0,0 1 0,00 1
ms 0, 1 1, 5 5 Hnh 5.15: Bin i dn Na+, K+ mng t bo tng ng vi s

0
hnh thnh in th hot ng (theo Hodgkim v Huxley).

+ S pht trin ca giai on kh cc: Trong giai on ny, th dn ca ion Na+ tng ln hng ngn ln, ng thi khi dn ca ion K+ thay i khng ng k. Kt qu o c cho thy c s phn cc ngc tr li so vi ban u. Lc ny dn ca ion Na+ ln hn dn ca ion K+ khong 30 ln. Ni cch khc tnh thm ca mng i vi ion Na+ by gi ln hn nhiu so vi ion K+. V vy in th mng trong giai on ny c xc nh gn nh hon ton bi s khuych tn ca ion Na+ hn l do bi cc ion K+. Da vo cng thc tnh in th ion, ta c.
RT [ Na + ] 0 (3.24) U Na = ln ZF [ Na + ]i Vy khi t bo trng thi hng phn, mng t bo b kh i dn n lm in th ngh gim. S gim in th ngh lm cho cc ion Na+ chuyn ng theo hng gradient nng vo t bo mt cch mnh m hn trc. Dng in do cc ion ny to ra cng b kh cc mnh, chnh l giai on qu kh cc ca mng (hnh 5.14b).
+ Giai on phn cc li: dn in ca ion Na+ ln hn dn in ca ion K+ ch trong khong thi gian vi mili giy (milisecond), nn giai on tip theo sau ta thy mng ging nh tr nn khng hot ng na. Tnh thm ca mng i vi ion Na+ li b c ch, cn tnh thm ca mng i vi ion K+ li tng ln. in th mng lc ny chu s nh hng nhiu bi ion K+. Tnh thm K+ gia tng tr nhng ko di lu hn, lng ion K+ khuych tn t trong ra ngoi t bo qua mng theo hng gradient nng mt cch mnh m lm cho mt trong t bo c gi tr m hn mt bn ngoi. Qu trnh pht trin theo khuynh hng tin ti cn bng, v in th lc ny c xc nh ch yu bi s tham gia ca ion K+, mng b tng nhanh qu trnh phn cc tr li hai pha mng. Hu qu ca giai on trn kt hp cng vi s hot ng ca bm + ion Na - K+ trong giai on phn cc li a mng tr v in th ngh ban u. Mng cng c gi tr in th m hn nhiu. ng thi vi s pht trin ca ion K+ lc ny khuych tn qua mng mt cch hon ton, lm cho mng c s phn cc nhiu hn. Do in th pha trong mng lc ny c gi tr m hn in th ngh bnh thng. Giai on hnh thnh ca in th hot ng ny chnh l giai on qu phn cc ca mng t bo.

Da vo cng thc Nernst xc nh gi tr in th hnh thnh trong giai on phn cc li (ch yu do K+ to nn), ta c:
UK =
RT [ K + ] 0 ln ZF [ K + ] i
(3.25)
VIII. p dng phng php o in th trong chn on v iu tr bnh.

Hin tng in ng, ngy nay c s dng mt cch ph bin trong Y-Sinh hc. Da vo cc quy lut t nhin, cc phng php vt l, l thuyt l sinh nghin cu h thng sng mt cch khoa hc. Tu theo ngun tc nhn c bn cht l ho khc nhau m lm thay i qu trnh phn b in tch hai pha mng t bo, thay i tnh thm, thay i p sut thm thu ...ca i tng sinh vt. Ni chung, cc yu t tc ng trn u lm thay i trng thi nhit ng, do lm nh hng qu trnh trao i cht, dn n s thay i trng thi chc nng ca cc c quan trong h thng sng. Tuy nhin di nh hng ca tc nhn l cc loi dng in ngoi c th xy ra nhiu tc dng khc nhau v nhiu mt ln c th ngi v ng vt. Chng hn nh: - Di tc dng ca dng in mt chiu, ta thy cc in tch-ion chu tc ng ca lc in trng. S phn b tr li cc in tch trn i tng kho st lm nh hng n qu trnh trao i cht cng nh trao i ion qua mng. Ni mt cch khi qut l c s thay i tnh cht ho l, cc ch tiu sinh hc ca nhng i tng sinh vt. Dng in mt chiu c ng dng nhiu trong Y hc lm ngun tc nhn kch thch c quan th cm thn kinh, o sut Galvanotonus, hoc s dng iu tr trong ion liu php (th nghim LeDue) - Di tc dng ca dao ng in t : Dng in cao tn c s dng rng ri trong y hc iu tr bnh. Da trn cc hiu ng nhit, hiu ng in - c khng mang bn cht nhit hoc hiu ng kch thch. Ngy nay trong vt l tr liu ng dng loi dng in ny iu tr cc bnh v xng, khp, kch thch c, thn kinh v phc hi chc nng... - Di tc dng ca ngun kch thch dng xung dng in hay xung in p vi cng nh v bin i trong khong thi gian ngn, tn s thch hp to cc kch thch in. Trong Vt l tr liu, ngi ta thng dng xung kch thch iu tr cc

bnh thn kinh, mch mu, ri lon chuyn ho...Trong lm sng, ngi ta cn s dng ngun tc nhn kch thch to cc kch thch in, sc in (electric shock). thm d chc nng hot ng ca c, thn kinh, trong nghin cu y hc ngi ta thng a ra cc thng s nh thi tr (Chronaxy), thi gian p ng hoc xc nh v ngng kch thch (Rhobase) ca cc i tng nghin cu. - Da trn hin tng in ng lm thay i tnh cht ho l ca cc t bo, thay i qu trnh trao i ion cng nh trao i cht trong h thng sng. Tm li, ngy nay vic ng dng v hin tng in ng trong Y-Sinh c s dng kh ph bin v ng vai tr thit yu khng th thiu trong vic chn on, thm d v iu tr bnh.
Chng 6
IN NG HC
I. Cc hin tng in ng hc.
Cc t bo, cc t chc sng, cc c quan ca h thng sng l mt h keo d th phc tp bao gm nhiu pha khc nhau. Do tc ng ca in trng ngoi khng i lm xut hin s chuyn ng tng i gia cc pha trong h, ngc li nu cc pha c thnh phn cht ho tan khc nhau th di chuyn ng c hc ca cc ion cng s to nn trong h mt hiu in th no . Cc hin tng in xut hin trong qu trnh ny c gi l cc hin tng in ng v chng c phn thnh cc loi sau y nh: in di, in thm, in th chy, in th lng... Nm 1809 Reis l ngi u tin pht hin thy cc hin tng in ng hc khi nghin cu chuyn ng ca cc ht t st di tc dng ca dng in mt chiu. Qua th nghim ca mnh, Reis thy rng cc ht keo mang in tch cng c kh nng vn chuyn c trong in trng, ng thi cng vi qu trnh bin i th mi trng phn tn cng s chuyn ng theo. Nh vy trong th nghim trn, Reis pht hin ra hai hin tng c bit quan trng, l khi cc ht tch in s dch chuyn di tc dng ca in trng ngoi, l hin tng in di hay in thm.
1. in thm.
L s chuyn ng ca cc mi trng phn tn ti pha in cc cng du vi din tch b mt ca pha phn tn. Da vo s dch chuyn ca cc ion trong in trng ta d dng phn tch mt hn hp polime sinh vt bng hin tng in thm hay in chuyn trn bng ghi. Dch sinh vt l cc dung dch in ly c nhiu thnh phn, vi ho tan khc nhau. Do di tc dng ca in trng ngoi, cc dung dch long v cc i phn t ion ho c tc dch chuyn khc nhau. Hin tng in thm d dng phn bit c l s chuyn ng ca dng cht lng, trong khi dng chuyn ng ca cc ht (phn t v cc i phn t ion ho) l do hin tng in di to nn. Qa trnh in thm c th xy ra trong nhiu trng hp v qua cc t chc sinh hc khc nhau, chng hn nh da ch, mng t bo, thnh ng mch, mao qun, vch mao mch...

Da vo gi tr hiu s in th in ho ca lp in tch kp ( ), Smolukhovski a ra cng thc xc nh tc chuyn ng tng i ca cc ht gia hai lp l:
v=
Trong :
E 4
(6.1)
l hng s in mi ca mi trng l h s ni ma st ca dung dch, E l cng in trng tnh ( v E c tnh theo n v mV )
2. in th chy.
in th chy xut hin khi cht lng chuyn ng di tc dng ca p sut thu tnh qua cc mao qun, hoc cc l nh ca mng, m thnh l c mang in tch. Hin tng dch chuyn ca cc cht lm xut hin in th chy theo chiu hng ngc li so vi hin tng in thm. y s chuyn ng ca mi trng phn tn s to nn mt hiu in th trong bn thn h. thy r iu ny, ta tin hnh th nghim nh sau: -Dng mt bnh thu tinh hai ngn cha dung dch sinh l, ngn cch nhau bng mt mng da ch. y dung dch sinh vt l cc mi trng phn tn, cn mng da ch ng vai tr to ra cc pha phn tn. Hin tng xy ra c biu din nh (hnh 6.1) di y:
Hnh 6.1: in th chy xut hin qua cc l mng. Nu tng p sut na bnh pha bn tri th cht lng s chuyn ng v bn phi bnh, ng thi gia hai pha ca bnh s xut hin mt hiu in th. in th mi hnh thnh chnh l gi tr ca in th chy. S chnh lch in th hai bnh l hiu in

th o c hai pha ca mng ngn cch hai dung dch. Nguyn nhn xut hin hiu in th trn l do trng thi cn bng tnh in b ph v. in th chy cng d dng kho st c khi tin hnh th nghim vi cc dch sinh vt v cho chuyn ng qua cc mng xp hay cc mng bn thm. Do , da vo hin tng trnh by trn, thng thng ngi ta hay s dng cc mng lc o xp ca cc i tng nghin cu l cc t chc sinh vt.
3. in th lng.
in th lng l hiu s in th xut hin gia lp trn v lp di ca dung dch a pha trong qu trnh lng cc ht mang in ca cc pha phn tn di tc dng ca trng lc. Bn cht ca hin tng lm xut hin loi in th ny khc hn so vi hin tng lm xut hin hiu in th trong qu trnh in di. C th kho st hin tng xy ra trong qu trnh lng ca mu chng hn (hnh 6.2). Mu l mt dung dch keo, cc thnh phn hu hnh ca mu (hng cu, bch cu) c trng lng ring ln hn huyt thanh nn s lng xung y bnh. Trong qu trnh lng mu lm xut hin mt s chnh lch in th gia lp trn v lp di ca dch sinh vt. in th xut hin trong trng hp ny chnh l in th lng.
Hnh 6.2: in th lng trong qu trnh lng mu
Qu trnh chuyn ng lm xut hin in th lng ca mu c th gii thch nh sau: Hin tng lng mu lm cc ion dng tch ra khi s chuyn ng ca cc thnh phn hu hnh. Kt qu thc nghim cho ta thy r iu , cc lp di ca dung dch c in th m v mang in lng m, cn cc lp trn mang in th dng v c nhiu in tch dng.

Tt c cc hin tng in ng u lin quan n s xut hin hiu in th gia pha phn tn v mi trng phn tn. in th ny cn c gi l in th in ng hoc Zeta in th ( - in th). Th in ng ch xut hin do qu trnh chuyn ng ca cc pha trong h d th. Hiu in th ny c hnh thnh trn ranh gii gia mng dung mi cc mng (gi l lp hp ph) trn b mt ca ht v ton b phn cht lng cn li ca dung dch.
4. in di. in di l phng php phn tch da trn s dch chuyn cc in tch, phn t nhim in di tc dng ca trng lc in khng i. Thng thng, ta s dng phng php in di tch chit cc thnh phn albumin, globulin trong huyt thanh; in di protin (phc cht lipid vi protin); in di glucoprotin (xc nh cc thnh phn ca protin v glucid trong huyt thanh). Nm 1937, Tiselius l ngi u tin a ra phng php in di trong mi trng t do kho st s hin din ca protin trong huyt thanh. Sau Tiselius, Cremer ta thy c nhiu nh khoa hc khc nh Durrum xy dng l thuyt v hin tng ny mt cch tng minh. Trong h thng sng, cc dch sinh vt l cc dung dch in ly, thng thng l s phn ly khng tuyt i hon ton. Nu gi l h s phn ly, vi qu trnh sinh vt thun nghch th phn ng s din ra nh sau: Th d: Kho st s ho tan ca Axit Acetic trong nc. Nu dung dch c nng C mol Axit Axetic khi ho tan trong mt lt nc, ta c: CH3 - COOH CH3COO- + H+ V nng phn t gam l: C(1- ) C C Vi l h s phn ly.
Theo nh lut Guldberg v Waage th: C .C = Const. C (1 ) (6.2)
2C =K 1
Vi K l hng s ion.
(6.3)

Tc ca cc loi ion l khc nhau, nn nu gi + v - l linh ng ca cc ion dng v m th mt dng in l:
i = ne ( + + ) S
Vi i l dng in bn ngoi, S l tit din ca vt.
(6.4)
Khi ht mang in c in lng q l cc i phn t Protin, cc ht keo hay ht mixen th di tc dng ca in trng E, chng s dch chuyn vi vn tc v. Cn khi lc in trng (fe) m cn bng vi lc ma st (fms) th ht s chuyn ng u:
qE = k v (6.5) v = qE/ k (6.6) Trong k l h s ma st ph thuc vo hnh dng, kch thc phn t ho tan v nht ca dung dch. Theo nh lut Stock, ta c lc ni ma st c xc nh l: Fms = 6rv (6.7)
Vi: r l bn knh ht nhim in l nht ca mi trng T (6.5) v (6.7) ta c:

v = qE/ 6r (6.8)
Do hin tng ni ma st xy ra, khi qu cu c bn knh r, chuyn ng tnh tin trong khi cht lu, th qu cu s ko lp cht lu gn mt tip xc vi n chuyn ng theo. Vn tc bin i theo hng chuyn ng, c biu din nh (hnh 6.3) di y:
2/3 r v
Hnh 6.3: Lc tc dng ln qu cu bn knh r chuyn ng trong cht lu

Nu t U l linh ng in di, th theo nh ngha: U =/E =q/k Khi in trng E c gi tr bng mt n v cng in trng th: U = . Vy: linh ng in di chnh l tc di chuyn in di ca cc ht nhim in di tc dng in trng ngoi c cng in trng l 1Volt/mt (V/m). V lc in trng l trng lc th, ta c th p dng cng thc th nng trng lc in khi lm dch chuyn mt n v in tch t im kho st xa ra v cng, nn in th Zeta ( ) c xc nh l: = q / r (6.9) = q / r Th (6.10) vo (6.8), ta c: = E / 6 Hay: (6.11) (6.12) (6.10)
6 v E
y l cng thc Smolukhovski xc nh Zeta ( - in th ca cc phn t, i phn t ion, cc ht hnh cu mang in ca lp in tch kp trong hin tng in di.
II. Ngun gc in tch b mt.

1. Phn b in tch mt. Dch sinh vt l cc cht c cu to t cc dung dch ho tan, c bit trong c cha nhiu i phn t (nh protin, polisacarit, axit nuclid...) cht ho tan, trng thi keo. i phn t sinh vt lun lun l dng cc polime cao phn t c phn b mt cch ri rc cha cc nhm phn cc. Cc nhm c cc ny gi cht vi cc phn t nc, do chng khng cn ht ln nhau na hoc l b ion ho v tt c cc phn t ny u mang nhng in tch cng du nn chng thng y nhau.

Cc t bo ng vt, cc t chc sinh hc trong h thng sng l cc i phn t phc tp, l cc h keo. Trong sinh vt ta thy c cc polime cao phn t nh: - Protin to thnh t cc mch peptid ca cc axit amin - Polisacarit l cc polime ca glucopirano - Axit nucleid l cc polime ca nuclotid. Thng thng, trn ranh gii gia hai pha ca h trong dch sinh vt c th xut hin mt hiu in th do cc lp in tch b mt. Cc in tch t do xut hin di s phn ly ion trong cc nhm chc ca cc i phn t. Chng hn nh i phn t protin dng lng tnh:
NH2 R
NH3+ R
Trong mi trng axit, cc phn t protin ng vai tr ca mt ion dng:

NH3+ R + HCl NH3+ R + Cl-
Trong dung dch kim, th i phn t Protin li ng vai tr ca mt dng ion m:
NH2 R + NaOH
NH2 R +Na+ +H2O
Cl- v Na+ l cc ion nghch Kt qu s to thnh hai loi ion y l do qu trnh ion ho cc nhm NH2 v COOH.

Tm li, cc in tch t do tn ti trn b mt cc ht c lin quan n pha phn tn. S xut hin in tch trn b mt ca cc i tng sinh vt c th do hai c ch: - C ch ion ho cc nhm phn ly. - C ch hp ph cc ion ca mi trng phn tn ln b mt ca cc i phn t. Trong mt s cc loi ion, ta thy c s xut hin ca ion H+ hay OH-. Do c ch hp ph khc nhau m in tch b mt ca cc i phn t sinh hc c in lng khc nhau nhiu. Ni cch khc, in tch mt ngoi c th xc nh c qua s bin i pH ca dung dch. Trong th d trn ta thy, i vi mi trng toan (axit mnh) th i phn t sinh hc mang in tch dng, ngc li trong mi trng kim ( pH cao) th i phn t sinh hc li mang in tch m. Nh vy do hin tng in chuyn m du ca in tch ca cc i phn t ph thuc vo pH ca mi trng. Do s phn b li cc in tch hai pha phn tn trong dch sinh vt lm xut hin mt th in ng do lp in tch kp to thnh. Do qu trnh ion ho cc nhm chc trong phn t m mt s ion s i vo mi trng phn tn, nhng ion ny gi l nhng ion nghch. Mt s ion cn li trong mi trng trn s c nh trn cc ht pha phn tn, chng s xc nh du ca in tch b mt gi l ion to th.
III. in th Zetta v phng php xc nh.

Lp in tch kp xut hin trong cu trc sinh vt c th din ra theo nhiu c ch khc nhau. Vi quan im tnh in hc, ta thy s xut hin lp in tch kp l do hin tng phn b cc loi in tch hai pha mng sinh hc. Ni mt cch tng qut hn l s sp xp ca hai loi ion tri du nhau trn ranh gii gia hai pha phn tn v mi trng phn tn nh (hnh 6.4) di y:
+ + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + ai phn t+ SH + +
+ +
+ +
+ +
+ +
(a)
(b)
Hnh 6.4: Lp in tch kp trn b mt i phn t sinh hc (a) v trn mng t bo (b).
Hai lp in tch kp trn cch nhau mt khong d. Theo Debye v Huxkey, ta c th xc nh c b dy d ca lp in kp ti nhit 250C l:
d =
3 ,. 5
(A0)
(6.13)
Vi l lc ion. Theo l thuyt ny, lc ion c gi tr c xc nh bng na tng s cc nng (gam/lit) ion c mt, cn nng cc thnh phn th c tnh bng cch nhn vi bnh phng ho tr nguyn t tng ng. Th d: Dung dch NaCl (0,01N) c: = (0,01 . 12 + 0,01 . 12 ) / 2 = 0,01 Hiu in th gia hai lp in kp gi l in th Zeta (). C th xc nh in th da vo hin tng in chuyn di tc dng ca lc in trng nh sau: Nu gi () l hng s in mi ca mi trng, (d) l chiu dy lp in tch kp, () l hiu in th lp kp, th in tch mt ngoi (q) ca cc i phn t sinh vt c xc nh l:
q =
4 d
(6.14)
Di tc dng ca in trng ngoi E th cc i phn t chu tc dng ca lc tnh in l:

F = qE
(6.15)
Tc dng ca lc in trng lm cho cc i phn t dch chuyn vi vn tc v. Gi s khi ion mi trng (lp ngoi) khng chuyn ng v ( l h s nht ca mi trng th ln ca lc in trng c xc nh l: Eq = T (6.14) v (6.16), ta c: qd = v d (6.16)
v
E
(6.17)
Cng thc (6.17) chnh l phng trnh Smolukhovski. Trong c tnh bng milivolt (mV), E l V/cm, v l (m/s th i vi nc ( = 80), ta c: = 0,0778 E Cu to lp in tch kp cng c th biu din theo s nh (hnh 6.5). Trong dung dch in ly, cc ion to th nm trn b mt ht keo, cn cc ion tri du th c phn thnh hai trng hp sau: - Trng hp th nht l cc ht mang in nm gn b mt ht keo (c kch thc phn t) c gi cht cnh b mt ht nh vo lc hp ph v c gi l lp hp ph. - Trng hp th hai l cc ht mang in chuyn ng t do di tc dng nhit trong mi trng phn tn to thnh lp khuych tn. AB: l b dy lp phn t, lp ny c b dy d cn c gi l lp hp ph BC: l mt phng ring khi cc phn t, i phn t sp xp trong in trng.

A B C
(a )
+ + -
- + + + + - + + - - + - + + + - + + - + + - + + -
+ + + + +
+ + + + + +
(b ) d
Hnh 6.5: Th nhit ng hc v th in ng. (a): S cu to lp in kp (b): c trng in th theo khong cch cc lp
Trong s trn ta c: E : th nhit ng hc. : th in ng
IV. Cc yu t nh hng n in th Zetta.

Khi cc phn t trong h chuyn ng, nu tt c cc ion tri du u b tch ra khi b mt ht keo th gia hai lp in tch ny s xut hin mt hiu in th E gi l th nhit ng. Thc t cho thy, do lc lin kt cc ion tri du trong lp hp ph bao gi cng chuyn ng cng vi ht keo. Do dng in to ra ch bi cc ht mang in t do cn hiu in th by gi c xc nh ch yu xy

ra gia lp hp ph v lp khuych tn. Hiu in th gia hai lp ny c gi l th in ng hay - in th. Do vy trong sinh vt, th in ng bao gi cng c gi tr nh hn th nhit ng. Tnh cht mi trng nh hng nhiu n gi tr ca - in th. Chng hn nh khi thay i nng ion trong mi trng phn tn, nhit mi trng lm thay i kh nng hp ph ion. Do thay i lp in tch kp lm cho gi tr - in th thay i theo. i lng - in th l mt ch s vt l c trng cho h thng sng. Da vo h s ny c th nh gi trng thi bnh l ca cc i tng sinh vt. Chng hn nh trng thi bnh thng, t bo hng cu mu ngi c gi tr khong 16,3 mv cn nu c s sai khc th l du hiu ca bnh l hoc ri lon v trng thi chc nng.
IV. ngha sinh hc ca Zetta in th.

Da vo bng phn tch v thnh phn cc cht cu trc bn trong v bn ngoi mng t bo hay s liu v - in th ca cc i tng sinh vt, ta thy chng ph thuc nhiu vo mi trng phn tn. Trong nhm axit phn ly mnh (nhm photphat nh hng ca phn t xephalin) cho thy th thay i ph thuc vo pH ca mi trng. Tht vy i vi hng cu, ta thy in th ph thuc vo in tch t do trn b mt t bo. Cc in tch b mt hng cu li b chi phi bi mc ion ha ca cc nhm phn ly trong phn t photpholipit. Nghin cu trn mt s ng vt, cc s liu ghi nhn c cho thy rng: Cc ng vt khc nhau s c th in ng khc nhau nh bng (6.1).
Bng 6.1: Th in ng ca hng cu ng vt trong dung dch m photphat pH=7.4 (ng trng). (Theo: Nguyn Th Kim Ngn-L sinh hc-HQG H ni-2001)
i tng Th Ln Chut nht Ngi Kh
(-in th (mV) 7 12.5 14.2 16.3 17.0

Mo Chut bch Ch 17.8 18.6 21.1
Cc loi vi sinh vt thng c kh nng hp ph cao i vi phn t protin cng nh cc cht hu c. Tht vy trong dch sinh vt c cc loi albumin, gelatin hoc cc sn phm phn hu t t bo, s hp ph trn b mt mng cng khc nhau, do th in ng xut hin cng c gi tr tng ng. Ngoi ra s hp ph b mt cn ph thuc vo i tng, thnh phn cng nh c im cu trc b mt ca chng.
Chng 7
C S HA L CA S HNG PHN
I. Khi nim hng phn v ngng hng phn
* Khi nim hng phn Hng phn l s chuyn t trng thi ngh ngi sang trng thi hot ng. Hng phn bao gm hai c ch: C ch tip nhn kch thch bi cc th quan v c ch chuyn tn hiu kch thch thnh tn hiu in, truyn v no x l thng tin v pht tn hiu thc hin phn ng tr li. Tn hiu kch thch rt a dng nhng ch yu l tn hiu vt l (nhit, nh sng, p sut...) v tn hiu ha hc (hocmn, mi, v...). Chc nng chuyn tn hiu kch thch thnh tn hiu in (tc sng hng phn) v dn truyn sng hng phn do noron thc hin. Thc hin phn ng tr li c th l c quan, m, t bo v c mc phn t. Trong h sinh vt, t sinh vt n bo ti sinh vt a bo tuy c mc tin ha khc xa nhau nhng u tn ti tnh hng phn thch nghi vi s thay i ca mi trng sng. * Khi nim ngng hng phn Cng
E D B C
2 Reobaz G 1 Reobaz A O Thi tr
Thi gian
Thi gian c ch
Hnh 7.1: Tng quan gia cng v thi gian kch thch
Ngng hng phn c xc nh bng cng nh nht v thi gian kch thch ngn nht c th to nn s hng phn. Cng nh nht kch thch to ra c phn ng tr li gi l 1 reobaz. Thi gian ngn nht khi kch thch 1 reobaz to ra c phn ng tr li l thi gian c ch (xem hnh 7.1). Trong thc nghim xc nh thi gian c ch rt kh nn Lapicque ly thi gian ng vi 2 reobaz o ngng thi gian kch thch, gi l thi tr. ng biu din tng quan gia cng v thi gian kch thch l ng hipecbol, ng vi phng trnh do Weiss a ra nm 1901:
i=
a +b t
(7.1)
i: Cng ngng t: Thi gian ngng
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com a: Hng s ng vi ng thng thi gian chy song song vi trc tung b: Hng s ng vi ng thng cng chy song song vi trc honh Nu cng i = 2b, ngha l bng 2 reobaz th phng trnh (7.1) s c dng: 2b =
a +b t
(7.2)
b=
a t
t=
a b
Thi tr thay i ty theo m. V d ngi thi tr ca c dui di gp t 1,5 n 2 ln so vi c gp.
II. L thuyt hng phn ca Heinbrun (1928)

Trn c s nhng s liu thc nghim v qu trnh hng phn c lin quan ti s thay i cu trc ha l ca nguyn sinh cht, nh l thay i tnh cht keo th hin t bo thc vt bc cao v ng vt nguyn sinh (amip) m Heinbrun a ra thuyt ng t vo nm 1928. Heinbrun cho rng: Tt c cc yu t kch thch u gy nn qu trnh ng t nguyn sinh cht km theo s tng t ngt nht cu trc ca n. Quan nim ny c cng c qua s liu v cc cht c ch s hng phn nh thuc ng, thuc m u lm gim nht ca nguyn sinh cht. Thuyt ng t ca Heinbrun gii thch cc yu t kch thch c bn cht khc nhau khi tc dng ln cc t bo ca th quan, trc tin gii phng Ca++ m trc trong nguyn sinh cht Ca++ li trng thi lin kt. Chnh do Ca++ c gii phng dn ti lm ng t nguyn sinh cht. Thuyt ng t ca Heinbrun mi ch gii thch c hin tng s kch thch dn ti lm ng t nguyn sinh cht cn nhiu hin tng khc lin quan ti s hng phn, thuyt ng t khng gii thch c.
III. Thuyt ph hy cu trc ca Naxonov v Alecxandrov (1940-1943)

Naxonov v Alecxandrov xem qu trnh hng phn nh l mt qu trnh ph hy cu trc. Hai ng quan nim nguyn sinh cht l pha khng ha tan trong nc v s phn b khng ng u ca cc cht trong ni bo v ngoi mi trng l do kh nng ha tan ca cc cht pha nc v nguyn sinh cht khc nhau v do kh nng lin kt ca cc cht vi phn t protein. Trong t bo cc cht in phn phn ln lin kt vi cc phn t protein, ch s t trng thi t do. Khi b kch thch hay b tn thng nguyn sinh cht c nhng thay i sau: - phn tn ca cc ht keo gim xung r rt v c ca ht nhn v nguyn sinh cht xut hin rt sm. - S tng nht ca nguyn sinh cht gm 2 pha: Khi yu t kch thch yu th nht nguyn sinh cht gim v khi yu t kch thch tng ln th nht nguyn sinh cht tng ln rt nhanh. - Khi kch thch, ban u qu trnh to ht trong nguyn sinh cht tng ln v sau qu trnh ny b c ch. - Khi kch thch pH ca nguyn sinh cht dch chuyn v pha axit. - Khi kch thch K+, PO4 , creatin c gii phng ra mi trng cn Na+, Cl- li xm nhp vo trong t bo rt nhanh.
3
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com Cng nh thuyt ng t, thuyt ph hy cu trc a ra ch l gii thch v s thay i tnh cht ha l ca nguyn sinh cht, mi gii thch hin tng ca qu trnh hng phn m cha gii thch c bn cht ca qu trnh hng phn.
IV. L thuyt hng phn ca Nernst (1899)

Nm 1887, Arenius cng b cc cht khi ha tan trong nc s phn ly thnh cc ion dng v ion m, di tc dng ca dng in ngoi, cc ion s chy v in cc mang in tch tri du vi in tch ion. Cc ion c kch thc v bn cht in tch khc nhau nn c vn tc chuyn ng v hai cc cng khc nhau. Sau khong thi gian nht nh s to nn nhng lp c mt ion khc nhau, tc s xut hin mt hiu in th. Nu ngt ngun in bn ngoi, thay vo cm 2 in cc vo dung dch in phn, ni vi mt bng in th n s sng. l dng in xut hin trong dung dch in phn. Nm 1899, Nernst da trn kt qu nghin cu ca Arenius cng xem t bo nh mt dung dch cht in phn c bao bc bi mng t bo. Di tc dng ca dng in kch thch, cc ion m v dng trong t bo cht s chy v hng in cc kch thch c in tch tri du vi in tch ion. Sau mt thi gian cc ion m v dng chuyn ng theo hai hng khc nhau s tp trung hai pha ca mng t bo. ngoi bo cng c cc ion dng v m, do lc ht tnh in, nu mt pha t bo, mt trong tch in m th mt ngoi mng tch in dng cn pha kia ca t bo mt trong s tch in dng v mt ngoi mng tch in m. Kt qu l gia bn trong v bn ngoi mng t bo hnh thnh nn mt hiu in th v khi in th ny t gi tr ngng th to ra s hng phn. S chnh lch v nng ion lin quan n cng v thi gian kch thch v to ra hng phn phi tha mn cng thc:
C Co = .i .
t k
(7.3)
C: Nng ion t do khi t bo hng phn Co: Nng ion t do khi t bo trng thi ngh ngi : S lng ion c dch chuyn do mt n v cng dng in i: Cng dng in; t: Thi gian kch thch k: H s khuych tn ca ion Nu kch thch bng dng in mt chiu th mi lin quan gia cng dng in v thi gian kch thch phi tha mn cng thc:
i t = hng s
(7.4)
Trong gii hn v cng dng in kch thch th nu cng dng in tng th thi gian kch thch gim v ngc li, duy tr tch s ca cng dng in vi thi gian lun l mt hng s. Nu kch thch bng dng in xoay chiu th mi lin quan gia cng v tn s dng in phi tha mn cng thc: i = hng s (7.5) Trong gii hn v cng dng in kch thch, nu tn s cao th cng dng in phi ln cn tn s thp th cng dng in nh, duy tr t s gia cng v
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com tn s lun l mt hng s. Cc cng thc trn do Nernst a ra ch ng trong phm vi tn s 100 Hz n 3 kHz. Kt qu thc nghim cho thy nu kch thch cng di ngng v kch thch lu hoc kch thch cng ln vi thi gian kch thch ngn th u khng gy ra hng phn. mi i tng nghin cu khi kch thch cng dng in l 1 reobaz th c mt thi gian kch thch cn thit (thi gian hu ch).
Bng 7.1: Thi gian hu ch ca mt s i tng nghin cu
TT 1 2 3 4 5
i tng nghin cu C trn d dy ch C trn ca nhuyn th C chn ca nhuyn th Thn kinh ch Thn kinh ng vt mu nng
Thi gian hu ch 1 giy 10-1 giy 10-2 giy 10-3 giy 10-4 giy
L thuyt hng phn ca Nernst khng nu ra c th s thay i nng ca nhng ion no v gii hn ngng nng l bao nhiu c th to ra hng phn.
V. L thuyt hng phn ca Bernstein (1906)

Nm 1906, Bernstein a ra l thuyt hng phn gii thch c ch hnh thnh in th tnh khi t bo trng thi ngh ngi v c ch hnh thnh in th hot ng khi t bo trng thi hng phn. Bernstein cho rng t bo khi trng thi ngh ngi do mng c tnh bn thm tc l thm K+ d dng cn Na+ thm t, trong khi hon ton khng thm cc phn t hu c mang in tch m (gi l cc amion hu c). Do vy, t bo trng thi tnh, bn trong c in tch m cn ngoi mng c in tch dng nn tn ti in th tnh v chiu in trng hng t ngoi vo trong t bo. Bernstein n thun ch xt s chnh lch nng K+ bn trong v bn ngoi t bo, p dng cng thc ca Nernst tnh c gi tr in th tnh ph hp vi gi tr o trc tip bng phng php vi in cc. Sau ny c nhiu s liu thc nghim lm sng t thm l thuyt ca Bernstein. Xegan chng minh cc ion ha tr hai (nh Ca++, Mg++,...) ht phn t nc (H2O) mnh hn so vi ion ha tr mt (nh Na+, K+, Cl-...). V th lp v hidrat ha ca ion ha tr hai dy hn so vi ion ha tr mt, nn ion ha tr hai kh lt qua siu l trn mng so vi ion ha tr mt. Mt khc, cng mt loi ion, nu ion no c kch thc nh (hay nguyn t lng b) s c in trng ln so vi ion c kch thc ln (hay nguyn t lng ln). V th ion c kch thc b ht nc mnh hn so vi ion c kch thc ln nn xy ra hin tng, trong dung dch ion c kch nh li kh thm qua siu l trn mng so vi ion c kch thc ln hn. V d Na+ c ng knh 1,9A0 b 8 phn t H2O bao quanh cn K+ c ng knh 2,6A0 ch c 4 phn t H2O bao quanh nn K+ thm qua siu l trn mng d hn so vi Na+. Nm 1955, Hodgkin v Katz bng phng php ng v phng x nh du xc nh c khi t bo trng thi tnh, dng Na+ i ra bng dng Na+ i vo cn dng K+ i ra ln gp xp x 1,63 ln so vi dng K+ i vo. Hn na khi t bo trng thi tnh, nng Na+ bn ngoi ln hn so vi bn trong nn chiu gradien nng Na+ hng t ngoi vo trong cn nng K+ bn trong li cao hn bn ngoi nn chiu gradien nng K+ li hng t trong ra ngoi . Song do tnh bn thm ca mng, theo tnh ton ca Goldman cho thy K+ i ra theo gradien nng K+ nhanh gp 76 ln so vi Na+ i vo cng chiu gradien nng Na+. Kt qu
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com thc nghim khng nh tnh ng n ca gi thuyt Bernstein, tnh bn thm ca mng l nguyn nhn dn n s phn b khng ng u ca cc ion gia bn trong v bn ngoi mng t bo nn to thnh in th tnh. Sau ny, nm 1942, Cuatite v Cole li chng minh rng K+ ng vai tr chnh trong vic duy tr in th tnh. Hai ng tng nng K+ ngoi bo ln (tc lm gim s chnh lch nng K+ gia bn trong v bn ngoi mng t bo) th gi tr in th tnh cng gim dn. Khi nng K+ bn ngoi bng nng K+ bn trong t bo th in th tnh s bng khng. Khi gim dn nng K+ bn ngoi v gi tr nh lc ban u th in th tnh li c khi phc. Song nu tng nng Na+ trong t bo ln th in th tnh vn khng thay i. Khi b kch thch, t bo chuyn t trng thi tnh sang trng thi hng phn v t bo b i cc, tc l bn trong t bo c in tch dng cn bn ngoi c in tch m. Khi t bo hng phn, Bernstein cho rng mng t bo mt tnh bn thm nn khng cn phn bit tnh thm gia K+ vi Na+ v cho c cc anion hu c thm ra bn ngoi. Chnh do cc anion hu c thm ra ngoi v cc Na+ thm vo trong d dng hn so vi K+ thm ra ngoi lm cho t bo b i cc. Sau ny thc nghim o c gi tr tuyt i ca in th hot ng ln hn gi tr tuyt i ca in th tnh nhng theo gi thuyt ca Bernstein th khng gii thch c bn cht ca hin tng ny. V d noron thn kinh lc ngh ngi c in th tnh l -90mV cn khi hng phn c in th hot ng t 120 n 130mV. Nm 1949, Hodgkin v Katz lm th nghim chng minh rng khi noron hng phn tnh thm ca mng i vi Na+ vo trong t bo tng ln 500 ln so vi khi noron ngh ngi. Ngc li, khi noron ngh ngi mng t bo cho K+ thm ra ngoi li nhanh gp 76 ln so vi Na+ thm vo trong t bo. Sau ny Goldman a ra cng thc tnh in th tnh v in th hot ng. Mi ion c mt h s thm c trng. Khi noron ngh ngi, nu ly tnh thm ca mng i vi K+ lm n v so snh v qui c PK+ =1 th tnh thm ca mng i vi Na+ l PNa+ = 0,013 cn tnh thm ca mng i vi Cl- l PCl-_= 0,045. Cng vi vic xc nh nng K+, Na+, Cl- bn ngoi v bn trong noron lc ngh ngi, thay cc gi tr bit vo cng thc Gondman s tnh c in th tnh bng -89mV. Khi noron hng phn nu vn ly PK+ = 1 lm n v so snh th PNa+ = 20, PCl- = 0,045. Cng xc nh nng K+, Na+, Cl- bn trong v bn ngoi noron lc hng phn, thay cc gi tr bit vo cng thc Goldman s tnh c in th hot ng bng 38mV. Khi noron hng phn, in mng chuyn t -89mV ln 38mV, do vy gi tr tuyt i ca in th hot ng phi l |-89mV|+38mV=127mV. Nh vy, bn cht ca hin tng in th hot ng c gi tr tuyt i ln hn gi tr tuyt i ca in th tnh. Khi ngng kch thch, mng noron s khi phc li tnh bn thm duy tr s chnh lch v nng ion ging nh lc ban u, tc l duy tr in th tnh. L thuyt mng ca Bernstein cha cp n vai tr ca cc ion ha tr hai nh Ca++, Mg++... Nhiu kt qu thc nghim xc nh vai tr ca Ca++ trong qu trnh hnh thnh in th sinh vt. Mt s nh khoa hc gi thit rng, trn mng t bo c cc knh dn "nhanh" v "chm" i vi cc ion. Khi t bo hng phn, cc knh dn "nhanh" cho dng Na+ vo trong t bo kh cc t bo (tc i du in tch m sang dng). Sau cc knh dn "chm" tip tc cho Na+ v Ca++ vo trong t bo kt thc qu trnh kh cc (tc l xut hin in th hot ng). L thuyt mng ca Bernstein, mc d c cc nh khoa hc b sung nhng vn cha y . Do vy, vn ny vn cn phi tip tc nghin cu v hon thin. VI. L thuyt hng phn ca Laxarev
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com Nm 1887, Ringer l ngi u tin pht hin dung dch ng trng NaCl c cho thm KCl v CaCl2 theo mt t l nht nh, gi cho c ch c phn x co c ging nh trong c th cn nguyn vn trong thi gian lu hn nhiu so vi c ch ch ngm trong dung dch ng trng NaCl. Sau ny Zac v Lob tin hnh th nghim trn trng c Fundulus thy rng: Trng khng n trong dung dch ch c NaCl. Nu dng dung dch CaSO4 c nng xc nh thm vo dung dch NaCl vi mt t l thch hp, to ra s tng quan ti u gia cc ion th trng c Fundulus s t t l to thnh phi cao nht l 75%. Sau ny tip tc c nhiu s liu thc nghim chng minh rng noron thn kinh ch c th hng phn khi trong bo tng ca si trc c c ion ha tr mt v ion ha tr hai. T nhng kt qu nghin cu trn, Laxarev a ra l thuyt ion v s tng thch hoc i khng gia mt s ion. ng cho rng, khi t bo trng thi ngh ngi s duy tr t l gia ion ha tr mt v ion ha tr hai mt gi tr xc nh v khng thay i: C1 = hng s (7.6) C2 C1: Nng ion ha tr 1 C2: Nng ion ha tr 2 Khi kch thch, t s ny b thay i dn n s hng phn nu t s ny tng hoc dn ti s c ch hng phn nu t s ny gim. L thuyt hng phn ca Laxarev gii thch c kt qu th nghim ca Flygn. l hin tng kch thch bng dng in mt chiu, khi ng mch th hng phn xut hin cc m cn cc dng th b c ch. Ngc li khi ngt mch, hng phn li xut hin cc dng cn cc m li b c ch. Laxarev gii thch nh sau: Khi ng mch, cc ion dng s ri khi cc dng v pha cc m theo hng ca in trng. Ion dng ha tr mt linh ng hn so vi ion dng ha tr hai cho nn tp trung cc m nhiu hn. Do vy, cc m t l gia ion ha tr mt trn ion ha tr hai tng ln, dn n hng phn xut hin cc m. Ngc li cc dng, cc ion dng ha tr hai ri chm nn c nng cao hn so vi ion dng ha tr mt ri nhanh lm cho t l ion ha tr mt trn ion ha tr hai gim xung gy ra s c ch hng phn cc dng. Khi ngt mch, khng cn dng in kch thch, cc ion s tr v trng thi phn b nh lc ban u (lc cha kch thch). Cc ion dng li di chuyn t cc m v pha cc dng. Ion dng ha tr mt b t phn t nc bao quanh so vi ion dng ha tr hai nn ion dng ha tr mt d dng thot ra khi cc phn t nc bao quanh hn ion ha tr hai. Ti cc dng, t l ion dng ha tr mt trn ion dng ha tr hai tng ln, dn n hng phn li xut hin cc dng. Ngc li, cc m, cc ion dng ha tr hai ri chm nn li c nng cao hn so vi ion dng ha tr mt ri nhanh lm cho t l ion ha tr mt trn ion ha tr hai gim xung, gy ra s c ch hng phn ti cc m. L thuyt hng phn ca Laxarev cha a ra c th ngng v t l gia ion ha tr mt v ion ha tr hai khi t bo trng thi ngh ngi bng bao nhiu? t bit c khi c kch thch dn n t l ion ha tr mt trn ion ha tr hai t gi tr vt ngng s gy ra hng phn cn bng hoc nh hn ngng s khng gy ra s hng phn. Vn ny cc nh khoa hc ang tip tc nghin cu nht l v vai tr c th ca ion ha tr hai b sung cho quan im ca Laxarev.
VII. C ch dn truyn sng hng phn trong dy thn kinh
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com Th nghim ca Hodgkin v Katz chng minh dng in hng phn xut hin trong dy thn kinh khi b kch thch c bn cht ion. Hodgkin v Katz cng ch r K+ c vai tr chnh trong vic duy tr in th tnh cn Na+ li c vai tr chnh trong vic hnh thnh nn in th hot ng (tc sng hng phn). Ty thuc vo bn cht ca dy thn kinh nh c mielin bao bc hay khng, ng knh si trc, chc nng ca noron m c tc dn truyn sng hng phn khc nhau (xem bng 7.2).
Bng 7. 2: Kiu si thn kinh v tc dn truyn sng hng phn trong dy thn kinh
ng Tc truyn (m/s) Kiu si knh si Bin nhit ng nhit (20oC) (37oC) () Anpha 10-20 20-40 60-120 Beta 7 - 15 15-30 40 - 90 Gamma 4-8 8 - 15 30 - 45 Denta 2,5-5 5-9 15 - 25 B 1-3 2-6 3-5 C 0,3-1,5 0,3 - 0,8 0,5 - 2
Chc nng Si vn ng c Si th cm (s m) Si hng tm t c Si th cm da (nng, lnh) Si tin hch dinh dng Si hu hch giao cm
Kt qu bng 7.2 cho thy ng vt ng nhit (chim, th, ngi) c tc dn truyn sng hng phn trong dy thn kinh nhanh hn so vi ng vt bin nhit (ch, c, lng th). Cc si thn kinh dn truyn cm gic au n c tc dn truyn chm nht (0,71,3m/s), cc si hng tm dn truyn cm gic s m, c tc cao hn t 50m/s cn cc si vn ng c tc dn truyn nhanh nht t ti 160m/s. Si trc thn kinh cng l mt dy dn in v nu l si trn (khng c mielin bao bc) th dch bo tng bn trong si trc c in tr l Rt cn mng noron c in tr l Rm. i vi dy thn kinh c mielin bao bc v do mielin l mt cht cch in rt tt nn noron ch tip xc vi mi trng ngoi qua eo Ranvie. Khi noron ch tip nhn kch thch qua eo Ranvie v dng in hng phn cng ch b suy gim do truyn in ra bn ngoi qua eo Ranvie. Khi b kch thch s xut hin xung in th hot ng ti in cc kch thch (cc m) v c k hiu l Vo. Do b tiu hao mt phn nng lng in thng in tr trong ca bo tng si trc v b r in qua mng noron nn gi tr ca in th hot ng b gim dn. in tr trong ca bo tng cng nh th in th hot ng b gim cng t v in tr mng noron cng ln th in th hot ng cng b gim cng t. Ngc li in tr trong ca bo tng ln th in th hot ng b gim nhiu v in tr mng noron nh th in th hot ng b gim cng nhiu. Cc nh khoa hc xc nh c gi tr in th hot ng sau khi pht sinh l Vo, truyn theo si trc thn kinh qung ng l x c gi tr l Vx c tnh theo cng thc: Vx=Vo. e (7.7) Rm: in tr mng noron t l thun vi in tr ring ca 1cm2 mng (k hiu l rm) v t l nghch vi bn knh si trc thn kinh (k hiu l r).
x Rm Rt
Rm =
rm 2r
(7.8)
Rt : in tr trong ca bo tng cng t l thun vi in tr ring ca 1cm3 bo tng (k hiu l rt) v t l nghch vi bnh phng bn knh si trc (r).

rt (7.9) r 2 Cc nh khoa hc tnh c ng vt thuc lp th, si trc dy thn kinh c mielin bao bc c bn knh r =15m, rm = 5000/cm2 v rt =50/cm3, in th hot ng Vo truyn c 1mm (l khong cch gia 2 eo Ranvie) cn li gi tr Vx c tnh theo cng thc: Vx = Vo.0,5 (7.10) Nu in cc kch thch t eo Ranvie th nht (gi l Ranvie 1) lm pht sinh in th hot ng l Vo=100mV khi truyn n eo Ranvie th hai (gi l Ranvie 2) theo cng thc (7.10) s cn 50mV. Thc nghim xc nh eo Ranvie c ngng kch thch in l 20mV. Do , dng in hng phn, tc in th hot ng pht sinh eo Ranvie 1 c gi tr l 100mV khi truyn n eo Ranvie 2 cn 50mV kch thch eo Ranvie 2 pht sinh in th hot ng cng c gi tr 100mV. C lp li nh vy, dng in hng phn hay cc xung in th hot ng c ln 100mV c truyn i theo noron cm gic v h thn kinh trung ng pht tn hiu truyn theo noron vn ng n m hay c quan thc hin phn ng tr li. i vi dy thn kinh khng c mielin bao bc, khi kch thch mt vng no th ti vng mng mt phn cc ri o cc nn c in tch tri du vi vng xung quanh cn ang trng thi tnh (xem hnh 7.1). Ti vng hng phn xut hin dng in hng phn n li kch thch vng ln cn v li to ra dng in hng phn mi ging nh dng in hng phn pht sinh ti vng b kch thch. S xut hin ca dng in hng phn sau khi b kch thch c lan truyn nh vy trn sut chiu di ca dy thn kinh mt cch lin tc. V vy, tc dn truyn ca dng in hng phn trong dy thn kinh khng c mielin bao bc thng chm v tiu hao nhiu nng lng. Rt =
A B C + + + -
Hnh 7.1: Dn truyn hng phn trong dy thn kinh khng c mielin bao bc
A v C: Vng noron trng thi tnh (trong m, ngoi dng) B: Vng noron trng thi hng phn (trong dng, ngoi m) : Si trc noron 2 : Dng in hng phn 3 : Hng truyn ca dng in hng phn v h thn kinh trung ng
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com i vi dy thn kinh c mielin bao bc, do mielin l mt cht cch in tt nn mng noron ch tip nhn kch thch eo Ranvie v mng noron cng ch mt phn cc v o cc (tc pht sinh in th hot ng) ti eo Ranvie (xem hnh 7.2). R1 R2 +2 1 + + + tnh + 2 hng phn + tnh
Hnh 7.2: Dn truyn hng phn trong dy thn kinh c mielin bao bc
R3 4
3 3
+ -
R1 v R3: Eo Ranvie 1 v eo Ranvie 3 trng thi tnh R2: Eo Ranvie 2 trng thi hng phn khi b kch thch : Si trc noron; 2: Bao mielin 3 : Dng in hng phn 4 : Hng truyn ca dng in hng phn v h thn kinh trung ng Theo hnh 7.2, khi kch thch eo Ranvie 2 th mng noron hng phn dn ti b o cc (trong c in tch dng (+), ngoi c in tch m (-)), c in tch tri du vi eo Ranvie 1 v eo Ranvie 3 ang trng thi tnh (trong c in tch m (-), ngoi c in tch dng (+)). Ti eo Ranvie 2 s xut hin in th hot ng (tc dng in hng phn) v dng in hng phn ny khi truyn n eo Ranvie 3 tuy gim i khong mt na nhng vn ln hn ngng gy kch thch nn to ra hng phn eo Ranvie 3, tc l li to ra in th hot ng mi c ln ging nh in th hot ng pht sinh lc ban u eo Ranvie 2. Dng in hng phn c lan truyn theo kiu "nhy" t eo Ranvie ny n eo Ranvie ln cn vi khong cch bc nhy l 1 milimt, theo hng v h thn kinh trung ng nn c tc truyn nhanh hn v t tiu hao nng lng hn so vi dy thn kinh khng c mielin bao bc. Nh hnh (7.1) dng in hng phn truyn theo hng t vng B n vng C cn hnh 7.2, dng in hng phn "nhy" t eo Ranvie 2 sang eo Ranvie 3 theo hng v h thn kinh trung ng (hoc ty sng) i vi dy thn kinh hng tm cn theo hng t ty sng hay h thn kinh trung ng ti m hay c quan thc hin phn ng tr li vi dy thn kinh ly tm. Mc d vy, vi dy thn kinh trn, pha ngoi mng noron c dng in truyn t vng C v vng B (hnh 7.1), t eo Ranvie 3 v eo Ranvie 2 (hnh 7.2) nhng u khng gy ra hng phn v khi mng noron tr tuyt i nu ang pha mt phn cc v o cc (khong 1 miligiy (ms)) hoc tr tng i nu ang pha ti phn cc (khong 3 ms) nn khng tip nhn kch thch. Nh vy, khi kch thch vng B xut hin dng in hng phn truyn n kch thch vng pha trc l C th hng phn li xut hin d dng cn nu truyn n vng C li quay v vng B th mng noron khng tip nhn s kch thch. i vi dy thn kinh ng vt mu nng, thi gian tr tuyt i ko di khong 0,002 giy -
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com 0,0004 giy. T bng 7.1, nu ta ly vn tc dn truyn trung bnh ca dy thn kinh nhm A l 60m/s, khi truyn t eo Ranvie 2 sang eo Ranvie 3 ri quay tr v eo Ranvie 2, qung ng l 2mm, tnh ra thi gian ch mt c 0,3ms, nh hn 1ms nn eo Ranvie 2 ang tr tuyt i nn khng tip nhn bt k kch thch no. Do vy, dng in hng phn truyn trong dy thn kinh ch theo mt chiu xc nh. Do mng noron c tnh tr nn mng noron khng th pht sinh cc xung in th hot ng mt cch lin tc c. Thi gian tr cng di th s lng ti a cc xung in th hot ng c mng noron pht sinh trong mt n v thi gian cng t v ngc li. Vedenski a ra khi nim tnh linh hot chc nng biu th kh nng hng phn ca cc t chc sng. Noron c tnh linh hot chc nng cng cao khi c kh nng truyn c s lng ti a cc xung in th hot ng trong mt n v thi gian cng nhiu. Ngc li, s lng ti a cc xung in th hot ng c truyn i trong mt n v thi gian cng t th tnh linh hot chc nng ca noron cng thp. V d, cc noron vn ng c thi gian tr l 2%o giy th ti a chng ch truyn c 500 xung in th hot ng trong mt giy. Cc noron trung gian c thi gian tr nh hn 1%o giy nn chng c th truyn ti a 1000 xung in th hot ng trong mt giy. R rng cc noron trung gian c tnh linh hot chc nng cao hn so vi cc noron vn ng.
VIII. C ch bn giao hng phn qua xinap

1. Cu to xinap
Hnh 7.3: Cu trc mt xinap
Cc v tr tn cng si trc ca mt noron tip xc vi cc noron khc v vi cc t bo c c gi l cc xinap. Cu trc mt xinap th hin trn hnh 7.3 gm mng trc xinap, khe xinap v mng sau xinap. Cc xinap l phn phnh to ca mt cc nhnh ca si trc noron trc. Trong cc xinap cha thnh phn quan trng nht, l cc bng xinap. Bn trong cc bng xinap cha cht mi gii. Gia mng trc xinap v mng sau xinap l khe xinap, rng khong 150A0 i vi xinap noron - noron, cn rng khong 500A0 xinap noron - c. Mng sau xinap c nhng th quan (receptor) chuyn bit nhn bit cht mi gii. 2. Bn giao hng phn qua xinap theo c ch vt l
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com Dng in hng phn mun truyn t noron trc sang noron sau phi vt qua mng trc xinap, khe xinap v mng sau xinap. C ba thnh phn ny u c in tr. Theo Katz, sau khi dng in hng phn vt qua ba in tr thuc cu trc ca xinap th in th hot ng t gi tr ban u khong 120mV, khi n mng sau xinap ch cn khong 0,01mV . Thc nghim xc nh, ngng kch thch mng sau xinap gy ra hng phn l t 20mV n 40mV. S liu do Katz a ra l khng ph hp vi thc nghim. gii thch c ch truyn xung in th hot ng qua xinap theo c ch vt l, cc nh khoa hc cho rng mng trc, mng sau v khe xinap c cu trc c bit nn c in tr rt b. Do vy, xung in th hot ng d dng vt qua ba thnh phn in tr trn nn khi n mng sau xinap gi tr in th hot ng tuy c b gim nhng vn ln hn 40mV. Vi gi tr vt ngng gy hng phn, n kch thch mng sau xinap lm cho mng sau xinap mt phn cc ri o cc nn li pht sinh xung in th hot ng cng c gi tr 120mV v tip tc c truyn i theo si trc ca noron sau. Tuy nhin, gi thit v ba thnh phn cu trc ca xinap c in tr nh, thc nghim cn cha xc nh c. 3. Bn giao hng phn qua xinap theo c ch ho hc Nm 1912 v 1921, Levi tin hnh th nghim buc hai tim c lp vo ng thng tim trong c cha dung dch sinh l hai tim thng vi nhau. Khi kch thch dy m tu ca tim mt th tim mt p chm v yu, c khi ngng p. ng thi tim hai cng p chm v yu, c khi ngng p ging nh tim mt. Nu kch thch dy giao cm ca tim mt th lm cho c tim mt v tim hai u p nhanh v p mnh. Levi xc nh dy m tu khi b kch thch pht sinh cht axetincolin c tc dng km hm cn dy giao cm khi kch thch s pht sinh cht adrenalin ch cn noradrenalin ngi c tc dng thc y tng nhp p ca tim. Th nghim ca Levi khng nh khi kch thch, hng phn xut hin vi s tham gia ca cht mi gii, truyn t tim mt sang tim hai. Cc xinap, khi noron trng thi tnh, c s tng hp axetincolin t axetat v colin. Lc u axetat kt hp vi coenzym A, to thnh axetin KoA. Nh xc tc ca enzyme colinaxetilase, xy ra phn ng gia axetin KoA vi colin to thnh axetincolin v coenzymA (KoA). Axetincolin sau khi tng hp s c tch ly li trong cc bng xinap c ng knh 0,02 - 0,03m, nm ri rc bo cht ca cc xinap. Khi dng in hng phn truyn n cc xinap gy tc dng kch thch lm cho cc bng xinap phng thch axetincolin vo khe h xinap. chut, mi xung in th hot ng khi truyn n cc xinap noron - c kch thch bng xinap gii phng vi triu phn t axetincolin vo khe h xinap. Cc phn t axetincolin vt qua khe h xinap mt khong 0,5ms. Axetincolin lm thay i tnh thm ca mng sau xinap v mng sau xinap rt nhy cm trc tc ng ca axetincolin. T s thay i tnh thm ca mng sau xinap dn n s mt phn cc v o cc, pht sinh in th hot ng c ln ging nh xung in th hot ng truyn n mng trc xinap. Nu l xinap noron - noron th xung in th hot ng pht sinh mng sau xinap, tip tc c truyn i theo si trc ca noron sau. ng thi mng sau xinap cng gii phng enzyme axetincolinesterase xc tc cho phn ng: Axetincolin + H2O Axetat + colin Mt phn t enzyme axetincolinesterase 25oC, trong 1 giy, c th thy phn c 300 ngn phn t axetincolin. Nh vy, mi phn t enzyme ch cn 1/300.000 giy l phn hy xong 1 phn t axetincolin nn mi gii phng ton b cht axetincolin c khe
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com xinap, trc khi c mt xung in th hot ng mi truyn ti, li c mt t axetincolin mi i vo khe xinap. Ngng gy kch thch mng sau xinap ca axetincolin ch cn nng v cng nh t 10-16 n 10-15M. Cc xinap gii phng cht mi gii l axetincolin l cc xinap kch thch v kch thch mng sau xinap lm pht sinh xung in th hot ng mi ging nh xung in th hot ng truyn n mng trc ninap. Trong c th sng cn tn ti cc xinap c ch gii phng cht mi gii c ch v lm c ch mng sau xinap, khng lm pht sinh xung in th hot ng mi mng sau xinap (tc khng to ra s hng phn mng sau xinap). Trng hp ny xy ra xinap noron - c tim ca ch c Levi pht hin khi kch thch dy m tu dn n gii phng cht c ch l axetincolin c tc dng c ch nhp p ca tim lm cho tim p chm v yu. Kt qu nghin cu khng nh hiu ng hng phn hoc c ch mng sau xinap khng phi do cht mi gii quyt nh m do bn cht ca cc th quan (receptor) mng sau xinap quyt nh. Do vy, axetincolin kch thch xinap noron - c nhng li c ch xinap noron - c tim. Hin nay cc nh khoa hc xc nh c mt s cht mi gii v tc dng ca chng (xem bng 7.3).
Bng 7.3: Cc cht mi gii v tc dng ca chng
Cht mi gii Axetincolin Adrenalin Noradrenalin Dopamin Serotonin Axit gamma - aminobutylic Glixin Axit glutamic Enxephalin v.v...
Tc dng Kch thch hoc c ch Kch thch hoc c ch Kch thch hoc c ch Kch thch hoc c ch Kch thch hoc c ch c ch c ch c ch c ch
Chng 8
QUANG SINH HC
I. Bn cht ca nh sng
nh sng l bc x in t trng lan truyn trong khng gian vi vn tc v cng ln (trong chn khng vn tc nh sng t 300.000km/s). nh sng c chia thnh 3 vng c bn: Vng nhn thy (vng kh kin) c bc sng () t 400 700nm Vng t ngoi c = 200nm 400nm Vng hng ngoi c = 700nm 1000nm Trong h sinh vt, cc loi c vng kh kin khng ging nhau. V d vi ngi vng kh kin c = 400nm 700nm nhng vi cn trng, vng kh kin li c = 320nm 500nm. nh sng va c tnh sng (c trng bi bc sng v tn s...) va c tnh cht ht (c trng bi cc lng t nh sng hay cn gi l photon). Photon c khi lng nh hn khi lng ca in t khong 1 triu ln (khi lng photon bng 1,785.10-27gam). Mi photon c mang mt gi tr nng lng c tnh theo cng thc: E = h. Hoc E = h.
v (8.1) E: Nng lng ca photon : Tn s nh sng bng s dao ng nh sng trong 1 giy
v: Vn tc nh sng bng 300.000km/s : Bc sng nh sng h: Hng s Planck bng 6,62.10-27erg.s n v dng xc nh nng lng photon l electron vn (hay in t vn), k hiu l eV hoc Kcal. Mt in t vn l nng lng cn thit cung cp cho 1 in t i qua th hiu mt vn vi khong cch gia hai in cc l 1Cm. Cng thc chuyn i gia cc n v o nng lng: 1eV=1,602.10-12erg v 1 Kcal=4,2.1010erg Thay gi tr hng s Planck v vn tc nh sng vo cng thc (8.1) s tnh c nng lng ca photon c bc sng tnh theo: 1244 E= (eV) (8.2) (nm) Nu quan nim mt photon tng tc vi mt phn t th chuyn mt phn t gam vt cht (1M) cha 6,023.1023 phn t ln trng thi kch thch cn c 6,023.1023 photon, gi l cn cung cp mt mol lng t (hay mt Einstein). Nng lng ca mt mol lng t tnh theo n v Kcal c tnh theo cng thc:
E=
28.650 (Kcal/M) (nm)
(8.3)
T cc cng thc trn thy rng nng lng ca nh sng t l thun vi tn s nh sng ( ) cn t l nghch vi bc sng nh sng (). Nng lng nh sng c bc sng ngn ln hn nng lng nh sng c bc sng di. V d nng lng tia t ngoi c bc sng =200nm, c gi tr 143,25Kcal/M cn nh sng kh kin c =750nm ch t 38,2Kcal/M.
II. Qui lut hp th nh sng

S hp th nh sng ca vt cht c biu hin ch cng nh sng b yu i sau khi xuyn qua lp vt cht nghin cu. Nm 1760, Lambert v sau nm 1852, Beer tm ra qui lut hp th nh sng bi vt cht. S bin i cng nh sng (k hiu l Chm sng ti Chm sng l dI) khi i qua lp mng vt cht (k hiu l dl) s t l vi cng nh sng chiu (k C hiu l I) v nng (k hiu l C) cng I Io nh h s k, c trng cho kh nng hp th ca vt cht. Biu thc ton hc ca nh l lut Lambert - Beer c th vit nh sau: Hnh 8.1: nh sng truyn qua dung dch dI = k.I.C (8.4) c nng C v chiu dy l. dl Du tr biu th s hp th nh sng bi vt cht. Phng trnh (8.4) c th vit di dng: dI = -k.C.dl (8.5) I Ly tch phn v tri theo cng nh sng t Io n I v v phi theo chiu dy t O n l ta c:
= k.C.dl Io I O
ln I
I Io
dI
= k.C.l
lnI - lnIo = -k.C.l I ln = k.C.l Io
(8.6) (8.7)
I = Io e kCl (8.8) Io: Cng nh sng ti C: Nng vt cht l: Chiu dy vt cht (tnh theo Centimet) Cng thc (8.6) c th vit di dng: lnIo - lnI = k.C.l
I: Cng nh sng l k: H s hp th
Io = k.C.l (8.9) I I t D = ln o , gi l mt quang hc ca dung dch. Khi cng thc (8.9) c th vit I nh sau: D = k.C.l (8.10) T cng thc (8.10) suy ra rng: mt quang hc ca dung dch t l thun vi nng dung dch v chiu dy ca lp vt cht m nh sng truyn qua. Khi l = 1Cm th mt quang hc (D) t l thun vi nng dung dch (C), th hin trn hnh 8.2. D Khi nng dung dch C = 1M v l = 1Cm th t (8.10) suy ra D = k. Nh vy, h s D3 hp th chnh bng mt quang hc ca dung dch c chiu dy 1Cm v nng D2 1M. ln
D1 O C1 C2 C3 C
Hnh 8.2: S ph thuc ca D vo C
Cng thc (8.10) c p dng tnh mt quang hc ca h ng nht (l h khng c ranh gii phn chia ra cc phn c nng khc nhau). i vi h d th nh h sinh vt, c nhiu phn c nng cht khc nhau th mt quang hc ca ton h s bng tng mt quang hc ca tng thnh phn ring r theo cng thc sau: D = D1 + D2 + D3 + ... Dn (8.11) D: Mt quang hc ca h D1, D2, D3 ... Dn: Mt quang hc ca lp 1, 2, 3... lp n. nh gi phn trm lng nh sng m mu vt hp th, ngi ta a vo khi nim truyn qua, k hiu l T, c tnh theo cng thc: I T= (8.12) Io T: truyn qua (n v l %) Io: Cng nh sng ti I: Cng nh sng l Khi mu vt hp th nh sng hon ton, tc I=0 th T=0. Nu mu vt hon ton khng hp th nh sng, tc I=Io th T=1 (hay 100%). Lng nh sng h hp th c tnh theo cng thc: (8.13) Ih = Io- I Ih: Cng nh sng b h hp th Io: Cng nh sng ti I: Cng nh sng l
T (8.12) c I = Io .T v thay I vo (8.13) ta c: I (8.14) Ih = Io - (Io.T) = Io(1-T) h = 1 - T Io
Ih gi l hp th. Khi mu vt hon ton khng hp th nh sng, tc T=1 th Io hp th bng khng. Nu mu vt hp th hon ton nh sng, tc T=0 th hp th I bng mt. Cc gi tr mt quang hc (D), truyn qua (T), hp th h c o I o trn my quang ph k. Trn thc t, thng s dng phng php xc nh quang ph hp th ca vt cht. Phng php ny o s ph thuc ca mt quang hc (D) vo bc sng () hay tn s () ca nh sng chiu. Mi cht c mt quang ph hp th c trng ring. Tc l, mi cht ch hp th cc i mt s bc sng nht nh. V d quang ph hp th ca mt s phn t sinh hc quan trng: - Quang ph hp th ca protein t gi tr cc i bc sng =280nm. - Quang ph hp th ca carotin t gi tr cc i bc sng =480nm. - Quang ph hp th ca rodopxin t gi tr cc i bc sng =550nm. - Quang ph hp th ca dip lc a t gi tr cc i hai bc sng 1=440nm v 2=700nm. - Mt ngi c th phn bit c 300 mu sc khc nhau nhng ch yu hp th ba mu c bn l mu (c =600nm), mu xanh (c =550nm) v mu da cam (c =450nm).
T s
III. Cc giai on c bn ca qu trnh quang sinh hc

Cc qu trnh quang sinh hc thng c nh gi theo hai quan im sau: - Quan im mt l nhng phn ng m cc sn phm cui cng ca n c d tr nng lng cao hn so vi nng lng ca cc cht ban u tham gia vo phn ng, c gi l nhng phn ng to nng lng. V d nh qu trnh quang hp thc vt. - Quan im hai l cc phn ng quang sinh hc trong nh sng ng vai tr l ngun nng lng hot ha cc phn t khi tham gia vo phn ng ha sinh hoc l di tc dng ca nh sng dn ti cc phn ng ph hy bin tnh mc phn t, t bo, m hay c th. Xt theo quan im no, qu trnh quang sinh hc cng u tri qua nhng giai on ni tip nhau nh sau: 1. Hp th lng t nh sng bi cc sc t hoc t bo (nh t bo que, t bo nn) gy nn trng thi hng phn hay cn gi l trng thi kch thch. 2. Kh trng thi kch thch in t ca phn t. Giai on ny xy ra cc qu trnh sau: - Thi hi nng lng qua cc qu trnh quang l (nh pht hunh quang hay ln quang). - Thi hi nng lng qua cc qu trnh quang ha dn ti hnh thnh nn nhng sn phm quang ha khng bn vng u tin. i vi qu trnh quang hp l cc sn phm NADPH v ATP. - Thi hi nng lng bng cch ta nhit ra mi trng.
3. Din ra cc phn ng trung gian khng cn ti s chiu sng (gi l phn ng ti) vi s tham gia ca cc sn phm quang ha khng bn vng ni trn to thnh sn phm quang ha bn vng (vi qu trnh quang hp, l hydratcacbon). 4. Hiu ng sinh hc cui cng nh cc biu hin sinh l cm nhn mu sc, s vt, s sinh trng v pht trin v.v... Giai on hp th lng t nh sng v giai on kh trng thi kch thch in t ca phn t, c trng chung cho tt c cc phn ng quang sinh vt (xem hnh 8.3). S2 (Singlet 2) 1 S1 (Singlet 1) A a b 2 3 T (Triplet) c 4 So (Singlet)
Hnh 8.3: S biu din cc mc nng lng ca phn t v cc bc chuyn gia cc mc nng lng .
- ng a, A khi phn t hp th nng lng nh sng chuyn t mc nng lng c bn ban u l So ln mc nng lng cao hn l S1 hoc S2. - Theo c hc lng t th khng c bc chuyn pht x (tc pht ra sng nh sng) khi phn t chuyn t mc S2 v mc So v cng khng c bc chuyn thng t mc So ln mc triplet (gi l bc cm). - Khi phn t chuyn t mc S1 v mc c bn So s pht hunh quang (ng b) hoc chuyn t mc triplet v mc c bn So s pht ln quang (ng c). - Cc ng 1, 2, 3, 4, l phn t thi hi nng lng di dng ta nhit ra mi trng. Qu trnh phn t hp th nng lng nh sng chuyn ln trng thi kch thch v sau tr v trng thi ban u, l mt qu trnh bt thun nghch.
IV. S pht quang

Phn t sau khi hp th nng lng nh sng chuyn ln trng thi kch thch c mc nng lng l S1 hoc S2 u c gi tr ln hn mc nng lng ban u ca phn t l So. Trong khong 10-13 giy, phn t mc nng lng E2 phi gii phng mt phn nng lng d tha qua con ng thi nhit ra mi trng tr v trng thi kch thch c mc nng lng thp hn l S1 (con ng 1 hnh 8.3). Khi phn t mc nng lng S1 n s tr v mc nng lng c bn So qua cc con ng sau: Qu trnh ta nhit (ng 2, 3, 4 trn hnh 8.3) Pht hunh quang (ng b trn hnh 8.3) S1 Pht ln quang (ng c trn hnh 8.3)
Vn chuyn nng lng Cung cp nng lng cho cc phn ng quang ha 1. S pht hunh quang Khi cc phn t chuyn t trng thi kch thch c mc nng lng thp nht l S1 xung trng thi c bn l So th s pht ra nh sng hunh quang. S pht ra nh sng hunh quang c th biu din di dng: S1 So + h'
v ': Tn s nh sng hunh quang ' = '

v: Vn tc nh sng; ': Bc sng hunh quang Thi gian phn t dng trng thi singlet S1 l t 10-9 n 10-8 giy, cho nn cng c xem l thi gian ko di ca s pht ra nh sng hunh quang. Do vy, s pht hunh quang ch xy ra trong thi gian chiu sng mu vt, cn khi ngng chiu sng th s pht hunh quang s tt. c trng ca nh sng hunh quang l ph hunh quang. Ph hunh quang l ng cong ph thuc ca cng hunh quang vo bc sng nh sng hunh quang ('). nghin cu ph hunh quang, cc nh nghin cu dng h thng cc knh lc, c b tr theo s sau: h 1
2
h',
Hnh 8.4: S ghi ph hunh quang.
1) Knh lc ch cho nh sng c m dung dch nghin cu hp th (vi dung dch protein th = 280 nm) i qua. 2) Dung dch nghin cu ph hunh quang (dung dch protein). 3) Knh lc ch cho nh sng hunh quang do phn t cht nghin cu pht ra. 4) My ghi ph hunh quang. S pht hunh quang tun theo cc qui lut sau: * Qui lut Stock: Ph hunh quang lun dch chuyn v pha bc sng di hn so vi im hp th cc i ca ph hp th. Bi v phn t hp th nng lng nh sng kch v thch c E = h. = h. (ng A trn hnh 8.3) mt mt phn nng lng do thi nhit (ng 1 trn hnh 8.3) tr v mc S1 v khi chuyn v mc So pht ra nh sng v hunh quang c nng lng E' = h' = h. . y E > E' nn < ', tc l nng lng nh ' sng kch thch ln hn nng lng nh sng hunh quang nn bc sng nh sng kch thch ngn hn so vi bc sng nh sng hunh quang. V d chiu dung dch protein nh sng kch thch c =280nm th cc phn t protein s pht ra nh sng hunh quang c bc sng di hn l =340nm. * Qui lut Vavilp: S pht ra nh sng hunh quang ca mt cht no (chng hn
Chlorophyll) khng ph thuc vo bc sng nh sng kch thch v nh sng hunh quang c pht ra khi phn t chuyn t mc nng lng S1So. Khi phn t hp th nng lng nh sng chuyn ln cc mc nng lng cao hn S1, qua con ng thi nhit cui cng u chuyn v mc S1 sau pht nh sng hunh quang. V d Chlorophyll c th hp th c nh sng xanh (=440nm) v nh sng (=700nm) nn khi chiu dung dch Chlorophyll d l nh sng xanh hay nh sng th ph hunh quang ca Chlorophyll vn khng thay i. * Qui lut Levin: Ph hp th v ph hunh quang i xng quanh mt bc sng o (xem hnh 8.5). Qui lut ng vi phn t c cu trc n gin. D 1 2
1 o 2
Hnh 8.5: Ph hp th (1) v ph hunh quang (2) i xng qua o
2. S pht ln quang Khi phn t hp th nng lng nh sng chuyn t mc nng lng c bn So ln mc kch thch S1 v khi tr v trng thi ban u c th bng s thi nhit (ng 2, hnh 8.3) hoc pht hunh quang (ng b, hnh 8.3), hoc thi nhit chuyn v mc triplet (ng 3, hnh 8.3), sau phn t chuyn t mc triplet v mc So v pht ra nh sng ln quang (ng c, hnh 8.3). S pht ln quang c th biu din di dng: TSo+h*
v *: Tn s nh sng ln quang * = * . Ln quang cng c c trng bi ph ln quang. Ph ln quang l ng cong ph thuc ca cng nh sng ln quang vo bc sng nh sng ln quang (*). Ph ln quang lun dch chuyn v pha nh sng c bc sng di hn so vi ph hp th v ph v v v hunh quang. Nguyn do v E ht = h. > E 'hq = h. > E lq = h. * suy ra bc sng nh ' sng hp th () nh hn bc sng nh sng hunh quang v bc sng nh sng hunh quang (') li nh hn bc sng nh sng ln quang (*) (xem hnh 8.6). S pht ln quang ko di t 10-4 n 10-2 giy, tc l lu hn so vi s pht hunh quang. Do vy khi tt nh sng chiu nhng s pht ln quang vn c th xy ra.
D 1
'
Hnh 8.6: Ph hp th (1), ph hunh quang (2) v ph ln quang (3) (<'<*).
3. S vn chuyn nng lng Phn t hp th nng lng nh sng chuyn t mc nng lng thp l So ln mc nng lng cao l S1. Khi phn t chuyn t mc nng lng S1So, n c th qua con ng thi nhit, pht hunh quang hay ln quang, hoc n c th chuyn nng lng cho phn t khc. Gi s phn t trng thi kch thch l cht cho nng lng (k hiu * l M1 ) cn cht nhn nng lng l phn t trng thi c bn (k hiu l Mo) th qu trnh chuyn nng lng c th vit di dng:
* M1 +MoM1+ M * 0
M * l phn t trng thi kch thch v c mc nng lng cao hn so vi nng lng 0 ca phn t Mo. Qu trnh chuyn nng lng l mt qu trnh vt l, khng km theo s bin i ho hc v khng cn M1 va chm vi Mo. S chuyn nng lng c th xy ra theo nhiu c ch nhng y ch xt c ch chuyn nng lng theo cng hng cm ng. c s chuyn nng lng theo c ch ny cn c mt s iu kin sau:
* - Cht cho nng lng M1 phi c kh nng pht hunh quang, c trng bi cng pht quang l J. * - Ph hunh quang ca cht cho M1 phi c vng chung vi ph hp th ca cht nhn Mo (xem hnh 8.7), c trng bi mt quang hc l D. D J * - Khong cch gia cht cho M1 v cht nhn nng lng Mo phi nh hn gi tr ti hn cho php. mt s iu kin khi khong cch trung bnh gia cht cho v cht nhn nng lng t khong cch t 20 n * 100 A th M1 c th chuyn ton b nng lng cho Mo, tc l hiu sut chuyn nng lng bng mt (=1). Vi h c cha nhiu phn t c kh nng hp th nh sng chuyn ln trng thi kch thch th kh nng xy ra s chuyn nng lng i vi phn t protein t ch vi phn trm cn axit nucleic t 30%
o
* Hnh 8.7: Ph pht quang ca cht M 1 l J v ph hp th ca cht Mo l D c vng chung (phn gch ngang)
v h c nhiu phn t hp th nh sng th c th t ti 100%.
V. Phn ng quang ha Phn ng quang ha l nhng phn ng xy ra trong h di tc dng ca nh sng. Phn ng quang ha no cng u tri qua hai giai on. Giai on u l giai on c chiu sng xy ra s hp th nng lng nh sng to nn nhng phn t b kch thch, cc ion v cc gc t do. Giai on th hai l giai on ti (khng cn s chiu sng) tip tc xy ra cc phn ng vi s tham gia ca cc sn phm quang ha c hnh thnh t giai on sng to nn sn phm quang ha bn vng. Tc phn ng quang ha c xc nh bng nng phn t cht tham gia vo phn ng trong mt n v thi dC ). Phn t ch c th tham gia vo phn ng khi hp th lng t gian (k hiu l dt nh sng chuyn ln trng thi kch thch (tc c hot ha). Do vy, tc phn ng quang ha phi bng s lng t nh sng c hp th trong mt n v thi gian dN ). T c phng trnh: (k hiu l dt dC dN = (8.15) dt dt S lng t nh sng c hp th trong mt n v thi gian t l thun vi s lng t nh sng chiu vo h (k hiu l I), nng phn t cht hp th (C) v tit din b mt b chiu sng (S). T c phng trnh: dN = S.C.I (8.16) dt T (8.15) suy ra: dC = S.C.I (8.17) dt Tuy nhin, khng phi tt c nng lng photon u c cc phn t hp th chuyn ln trng thi kch thch v sau phn t kch thch tip tc tham gia vo phn ng m c mt s phn t kch thch khng tham gia vo phn ng m tr v trng thi ban u bng cch thi nhit ra mi trng hay pht quang. V th c khi nim hiu sut quang ha (hay sut lng t, k hiu l ) c xc nh theo cng thc:
S photon c phn t hp th tham gia vo phn ng (N2) S photon c h hp th (N1) Hay:
N2 N1
(8.18)
Trn thc t gi tr bao gi cng nh hn 1 v ch t vi phn trm n vi chc phn trm. V d nh hiu sut quang ha ca phn ng kh hot tnh ca men khi b chiu sng ch t t 10-3 n 10-2 (tc t 1 0 00 1%), ngha l mt phn t men phi hp th t
100 n 1000 photon mi b mt hot tnh xc tc. S d hiu sut quang ha c gi tr thp nh vy l do ch c mt s t phn t sau khi hp th photon tham gia vo phn ng. T dn ti khi nim tit din quang sinh () v c cho rng ch khi photon p trng tit din quang sinh ca phn t mi dn ti phn t tham gia vo phn ng. By gi hiu sut quang ha c tnh theo cng thc: = (8.19) S <S <1 Phng trnh (8.17), thay khi nim tit din b mt b chiu sng bng tit din quang sinh s c vit di dng: dC = - .C.I (8.20) dt Du tr biu hin nng phn t kch thch khi tham gia vo phn ng quang ha s gim dn theo thi gian. Phng trnh (8.20) c th vit di dng: dC = - .I.dt (8.21) C Nu ly thi gian bt u xy ra phn ng quang ha l khng giy, ng vi nng ban u ca cc phn t tham gia vo phn ng quang ha l Co th sau thi gian l t, s phn t kch thch cha tham gia vo phn ng l C. Ly tch phn hai v (8.21) ta c:
dC C = .IO dt Co
C t
ln
C = - .I.t Co
C = e .I.t C = C o .e .I.t Co
(8.22)
Nng phn t kch thch tham gia vo phn ng quang ha s gim dn theo thi gian. nh gi tc phn ng quang ha c ph thuc vo bc sng nh sng chiu (), dC ngi ta o tc phn ng quang ha nhng bc sng khc nhau nhng c dt cng mt cng nh sng l I (tc l cng s lng t nh sng chiu vo mu vt). dC /I s ph thuc vo bc sng nh sng chiu c gi l o quang ph Khi gi tr dt hot ng ca phn ng quang ha. Quang ph hot ng ca phn ng quang ha c dng trng vi quang ph hp th ca cc phn t tham gia vo phn ng quang ha. V d: Trong quang ph hot ng ca qu trnh quang hp, xut hin nh cc i vng bc sng 480nm - 500nm m carotin cng c nh hp th cc i ti vng bc sng ny, chng t carotin c tham gia vo qu trnh quang hp. Hoc khi chiu nh sng ln dung dch men th hin ph hot ng ca phn ng quang ha dn ti kh hot tnh men trng vi ph hp th ca axit amin thm. iu ny chng t axit amin thm trong thnh phn cu to ca phn t men tham gia vo phn ng quang ha nn dn ti lm cho phn t men b mt hot tnh xc tc.
Nu cht A hp th nng lng nh sng nhng tham gia vo phn ng quang ha li l cht B th ph hot ng v ph hp th ca cht B phi trng nhau. Nh vy c ngha l cht A chuyn nng lng cho cht B (A+hA*BB*) cht B chuyn ln trng thi kch thch (B*) v tham gia vo phn ng quang ha. Nu trong h c nhiu cht, khi chiu sng v nghin cu ph hot ng, sau so snh vi ph hp th ca nhng cht bit s xc nh c cht no chuyn nng lng (nn khng c ph hot ng) v cht no tham gia vo phn ng quang ha (nn c ph hot ng).
VI. Phng php o hp th trong vng nh sng trng thy v t ngoi

Khi o hp th, cc nh nghin cu hay dng my quang ph k (cn gi l my so mu). S cu to my quang ph k c m t trn hnh 8.8.
Hnh 8.8: S cu to my quang ph k
1) Ngun chiu sng 2) Knh lc bc sng nh sng 3) Cc ng mu (cuvt) 4) T bo quang in 5) Mn hnh thit b o gi tr mt quang hc (D) hay truyn qua (T) Nguyn tc o: Cho dung mi (cht dng pha dung dch nghin cu) vo ng thy tinh s 3 v chnh thit b o s 5, cho kim ch s khng. Sau dung mi ra, trng sch, sy kh, ngui ri dung dch nghin cu vo, trn mn hnh thit b o s 5, kim s ch gi tr mt quang hc D hay truyn qua T. Lu c dung mi v dung dch nghin cu u o bc sng c chn trc. V d o mt quang hc ca dung dch ADN th phi chn bc sng =260nm. Mi cht hp th cc i mt bc sng nht nh (k hiu l max) v khi chiu nh sng max, i vi mi cht c mt h s hp th xc nh (k hiu l kmax). Song cng c nhng cht hp th cc i mt s bc sng khc nhau nh triptophan, tirozin, phenilalanin v do vy chng cng c cc h s hp th tng ng (xem bng 8.1).
Bng 8.1: hp th v h s hp th bc sng max.
Cc cht Tirozin Phenilalanin Triptophan Hixtidin Xistein Adenin Adenozin Guanozin Guanin Xitozin Xitidin
max (nm) 274; 222; 193 257; 206; 188 280; 219 211 250 260,5 259,5 252,5 246 267 271
kmax (1,4; 8; 48).10-3 (0,2; 9,3; 60).10-3 (5,6; 47).10-3 5,9.10-3 0,3.10-3 13,4.10-3 14,9.10-3 13,6.10-3 10,7.10-3 6,1.10-3 9,1.10-3
Uraxin Timin Timidin ADN ARN
261,1 264,5 267 260 258
10,1.10-3 7,9.10-3 9,7.10-3 6,6.10-3 7,4.10-3
Phng php o quang ph hp th trong vng t ngoi v nh sng nhn thy c p dng xc nh nng . V theo nh lut Lambert - Beer th mt quang hc (D) t l thun vi nng dung dch (C). V vy, mun xc nh nng ca mt cht cn nghin cu th trc tin phi c th chun D = f() ca cht nghin cu chun (t lm hay ly t ti liu tham kho (xem hnh 8.2). Sau ly dung dch cht cn nghin cu o c gi tr mt quang hc l Dx, da vo th chun s suy ra c nng Cx. xc nh nng vi khun (Cvk) cch lm D(650nm) cng tng t, ch vic thay nng phn t D=f() gam (M) bng s lng vi khun trong 1ml v o bc sng =650nm (xem hnh 8.9). V d: T th chun D = f(=260nm) ca dung dch ADN chun bit c nu D260 (mt quang hc o =260nm) bng 1 s tng ng vi nng ADN l 50 Dx microgam/ml, nu D260 bng 0,5 tng ng vi nng ADN l 25 microgam/ml, nu D260 bng 0,1 tng ng vi nng ADN l 5 O Cx C microgam/ml.
Hnh 8.9: Nng vi khun (Cvk) xc nh bng cch o mt quang hc (D)
Phng php o quang ph hp th cn c th cho php xc nh thnh phn ca phn t. T bng 8.1 bit h s hp th kmax ca 4 baz Nit l Adenin, Timin, Guanin, Xitozin. Da vo bc sng max trong bng 8.1 s o c mt quang hc tng ng ca mi baz Nit. p dng cng thc (8.10) D = k.C.l v khi chn cuvt c dy l = D 1Cm th suy ra D = k.C. Khi bit D v k s tnh c nng C = . k * Phng php phn tch hp th nguyn t - Phn tch nh tnh: S hp th ca nguyn t vt cht i vi nh sng mang tnh cht chn lc, ngha l mi nguyn t ch hp th mt bc sng nh sng nht nh. iu ny cho php nhn dng c nguyn t c mt trong mu em phn tch khi so snh quang ph hp th ca mu vi quang ph hp th ca cc nguyn t ho hc bit trc. - Phn tch nh lng: p dng nh lut Lambert - Beer v s dng cng thc (8.10) s xc nh c nng ca cht nghin cu khi o c mt quang hc D v h s hp th nguyn t k nh sng chiu c bc sng xc nh (nh sng n sc). Phng php phn tch hp th nguyn t c chnh xc cao hn nhiu so vi phng php dng quang ph k.
VII. Quang hp
1. Phng trnh quang hp tng qut Trn tri t ch c hai ngun nng lng m th gii sinh vt c th s dng l nng lng ho hc ca cc cht v c c ngun gc t tri t v nng lng nh sng c ngun gc t v tr. Nng lng nh sng l ngun nng lng v tn, c ngha quyt nh ti s sng trn tri t c thc vt v nhiu loi vi sinh vt quang dng chuyn ha thnh nng lng ho hc trong cc phn t hu c. Nh qu trnh quang hp m mi nm trn tri t c 5.1010 tn cht hu c c tng hp, hp th 2.1012 tn CO2 v thi ra 13.1010 tn O2 vo kh quyn. Phn ng tng qut ca quang hp l phn ng tng hp cc cht hu c (hydratcacbon) t cc cht v c (kh CO2 v H2O) di tc dng ca nh sng (k hiu l h).
6CO2 + 6H2O diep C6H12O6 + 6O2 luc Qu trnh quang hp din ra thc vt bc cao v thc vt bc thp. Mt loi quang hp khc din ra vi khun theo phn ng:
CO2 + 2H2S h CH2O + H2O + 2S i vi vi khun, qu trnh quang hp khng thi ra oxy m thi ra kh lu hunh. 2. Nng lng trong qu trnh quang hp nh lng vai tr ca nh sng c th tnh tng nng lng cc lin kt cc cht tham gia vo phn ng v cc sn phm cui cng ca qu trnh quang hp theo phng trnh tng qut sau y:
O=C=O+HOH h HC=O+O=O
H cc cht tham gia c: 2 ni C=O c nng lng l: 2x190 Kcal/M 2 ni O H c nng lng l: 2x110 Kcal/M Nng lng tng cng l: 600 Kcal/M sn phm quang hp c: 1 ni O=O c nng lng l: 1x116 Kcal/M 1 ni C=O c nng lng l: 1x190 Kcal/M 2 ni C H c nng lng l: 2x92 Kcal/M Nng lng tng cng l: 490 Kcal/M So snh nng lng trc v sau phn ng ta c: 600Kcal/M-490 Kcal/M=110Kcal/M Nh vy nng lng nh sng cung cp cho qu trnh quang hp cng chnh bng 110 Kcal/M ph v lin kt c (cht tham gia) v hnh thnh lin kt mi (sn phm). Theo tnh ton l thuyt 1 photon nh sng (=690nm) c nng lng khong 42Kcal/M v nh vy c 110Kcal/M, thc vt ch cn hp th 3 photon nh sng l thc hin qu trnh quang hp. Trn thc t, theo dn liu ca nhiu tc gi to ra 1 phn t oxy,
ti thiu phi cn t 4 n 12 photon. Nu ly gi tr trung bnh s photon cn thit cho qu trnh quang hp l 8 th hiu sut trung bnh ca qu trnh quang hp s l: 3 photon 42Kcal = = 0,375 hay t 37,5% 8 photon 42Kcal Hiu sut cc i ca qu trnh quang hp c th t c l: 3 photon 42Kcal = = 0,75 hay t 75% 4 photon 42Kcal
3. Sc t cm quang thc hin c qu trnh quang hp, cc t bo thc vt, cc vi khun phi c cc sc t cm quang tip nhn nng lng nh sng. C ba nhm sc t chnh l: Porphyrin, Phycobilin v Carotenoid. - Sc t nhm Porphyrin: Tt c cc loi thc vt v vi khun c quang hp u cha Chlorophyll cc dng khc nhau. T bo thc vt cha Chlorophyll cn vi khun quang hp cha Bacteriochlorophyll. Chlorophyll cc loi khc nhau khng ng k v cu trc phn t nhng li khc nhau v quang ph hp th bc sng max. Cc c th tin hnh quang hp thi oxy nh thc vt bc cao, to v vi khun Cyanobacteria hp th cc i bc sng 650nm v 680nm. Vi khun xanh lc ch yu cha Bacteriochlorophyll c, d, e li hp th cc i bc sng 725nm; 740nm; 760nm cn vi khun ta cha Bacteriochlorophyll a hp th cc i n bc sng 950nm v cha Bacteriochlorophyll b hp th cc i bc sng 1020nm n 1100nm. - Phycobiliprotein: L cc sc t mu hoc xanh lam, c mt vi khun lam, to , to nu... Nh c Phycobiliprotein m t bo vi khun lam hp th nh sng c bc sng vng 450-700nm. Cc sc t nh Phycoerythrin hp th cc i bc sng 565nm, Phycocyanin hp th cc i bc sng 620nm, Allophycocianin hp th cc i bc sng 654nm... - Carotenoid: Nhm Carotenoid c th chia lm ba loi: khng vng, n vng hoc hai vng. thc vt bc cao c mt Carotenoid thuc loi hai vng, vi khun ta c Carotenoid thuc loi khng vng cn vi khun xanh c n 80-85% Carotenoid thuc loi mt vng. Nh c tp hp cc sc t quang hp a dng nh vy nn cc sinh vt c quang hp hp th c hu ht ph nh sng mt tri chiu xung tri t (bc sng t 350 n 1100nm). S khc nhau v vng nh sng b hp th cc nhm sinh vt khc nhau lm cho chng khng phi cnh tranh vi nhau v ngun nng lng chiu sng. 4. S chuyn nng lng trong quang hp Tp hp cc sc t tham gia vo qu trnh quang hp, gia chng c xy ra s chuyn nng lng. Cc nh khoa hc tnh ton trong gran ca lc lp c nng Chlorophyll rt cao t 0,1 n 1M/dm3 v khong cch trung bnh gia cc phn t sc t ch khong t
10 n 20 A . S chuyn nng lng gia cc phn t sc t xy ra theo c ch cng hng cm ng. thc vt bc cao v to, bng phng php ph v t bo, tch ring ly lc lp v dng phng php in di, sc k tch c phc hp sc t - protein. Trong thnh phn phc hp sc t - protein c khong 50-60% Chlorophyll ca t bo thc vt hay to. Trong phc hp sc t ny c hai dng Chlorophyll pht hunh quang cc i bc
sng 680nm v 695nm. V vy, ngi ta cho rng c hai knh vn chuyn nng lng, bt u t s nhn nng lng nh sng bi Chlorophyll b sau vn chuyn nng lng cho Chlorophyll a hp th cc i bc sng 680nm (k hiu l Chl.a (680)) v Chlorophyll a hp th cc i bc sng 695nm (Chl.a (695)). Chlorophyll c kh nng hp th nh sng chn lc, chuyn nng lng hp th c t nh sng sang cho h vn chuyn in t quang hp chuyn thnh ha nng. Nhm sc t Carotenoid cng c kh nng hp th nng lng nh sng ri chuyn nng lng cho h Chlorophyll thc hin qu trnh quang hp. Nhm sc t Phycobilin cng c kh nng hp th nng lng nh sng vng bc sng ngn v chuyn nng lng cho Chlorophyll thc hin qu trnh quang hp. V d: vi khun lam, qu trnh vn chuyn nng lng din ra theo chiu hng sau: Phycoerythrin Phycocianin Allophycocianin (565nm) (620nm) (654nm) Allophycocianin B Chlorophyll a. (671nm) (680nm) Cc s ghi trong ngoc di l bc sng hp th cc i (max) ca mi sc t. 5. C ch ca qu trnh quang hp Qu trnh quang hp c chia ra 2 giai on (hay 2 pha, c gi l pha sng v pha ti). * Pha sng quang hp y l pha u tin ca quang hp, c s tham gia trc tip ca nh sng. Sc t quang hp l Chlorophyll (Chl) hp th nng lng nh sng do photon chuyn cho chuyn ln trng thi kch thch (Chl*) c mc nng lng cao hn so vi trng thi ban u (Chl). E = h Chl Chl* in t ca phn t Chlorophyll trng thi kch thch, khi tr v trng thi ban u chuyn nng lng cho h chuyn in t nh v trong lc lp tng hp nn ATP. pha sng quang hp c s tham gia ca phn t nc. Di tc dng ca nh sng vi s tham gia ca h sc t v h oxy ha trong lc lp m phn t nc b phn li, gi l s quang phn li nc. Quang phn li nc xy ra qua nhiu giai on nh sau: 4H2O Sac to 4H+ + 4OH 4OH - 4e - + 4OH 4OH
h
Cl O2 + 2H2O
Mn + +
Kt qu: 2H2O 4H+ + 4e - + O2 Oxy c thi vo khng kh cn proton (tc H+) v in t (e-) s tham gia vo cc qu trnh tip sau. in t c vn chuyn qua h vn chuyn in t quang hp tng hp nn ATP cn H+ kt hp vi NADP- to nn NADPH2. Tm tt phn ng ca pha sng nh sau:
2H2O + 2ADP + 2H3PO4 + 2NADP Sac to 2ATP + 2NADPH2 + O2 Sn phm ca pha sng l ATP, NADPH2 s tip tc tham gia vo phn ng tip theo ca pha ti tng hp nn cht hu c vi s tham gia ca CO2. * Pha ti quang hp Cc phn ng ca pha ti din ra khng c s tham gia trc tip ca nh sng chiu m c s tham gia ca ATP, NADPH2 v CO2. Sn phm u tin ca pha ti quang hp l tng hp nn glucose (C6H12O6). C 3 con ng tng hp glucose t CO2 ng vi 3 nhm thc vt khc nhau l thc vt C3, thc vt C4 v thc vt CAM (Crasulacean Acid Metabolism), c thc hin theo cc chu trnh C3, C4, CAM. VIII. Tia t ngoi v cc hiu ng sinh hc ca n 1. Tia t ngoi v hiu ng tc dng ca n mc t bo Trong thin nhin ngun bc x t ngoi l mt tri. Trn ng i ti tri t, phn ln nng lng ca tia cc tm b tng Ozn hp th v ch cn li phn nh sng c bc sng t 200nm n 400nm (nanmt) tc dng ln c th sinh vt. Tia t ngoi thng c chia ra 3 vng chnh sau: - Vng cc tm sng di c = 320nm n 400nm - Vng cc tm sng trung c = 280nm n 320nm - Vng cc tm sng ngn c = 200nm n 280nm Khc vi tia gamma () v tia Roentgen (X), tia t ngoi vng 200nm n 400nm khng c kh nng gy ra qu trnh ion ha cc phn t sinh hc. Tia t ngoi ch gy ra hin tng kch thch lm cho in t ca cc phn t sinh hc t mc c bn chuyn ln mc kch thch c nng lng cao hn, tc l hot ha cc phn t . Do vy, cc phn t sinh hc c hot ha, chng d dng tham gia vo cc phn ng ha sinh dn ti gy kch thch hoc lm tn thng c th sinh vt. Mt khc, kh nng xuyn su ca tia t ngoi km, ch vi milimt, do n ch gy nn cc hiu ng sinh hc lp t bo ngoi cng. Tia t ngoi c tc dng lm xm da, gy pht ban hoc gy bng da. Tia cc tm sng trung kch thch s tng hp vitamin D c tc dng chng ci xng... Tia cc tm sng ngn c tc dng lm thay i cu trc c trng ca lipit v protein, c tc dng dit khun. Hu ht cc cng trnh nghin cu trn cc i tng nh vi khun, vi sinh vt, t bo thc vt v ng vt, u pht hin qui lut chung l kh nng gy ra t vong nhiu, khi chiu tia t ngoi c bc sng nh hn 300nm v t vong nhiu nht bc sng 260nm. iu ny c lin quan ti s tn thng ca phn t ADN v n hp th cc i bc sng 260nm. Tia t ngoi cn gy ra hiu ng c ch s phn bo hoc kch thch s phn chia t bo ty thuc vo bc sng nh sng chiu. V d nh Coviadin khi chiu tia t ngoi c bc sng 256nm ln cc t bo nm men gy ra s c ch phn chia t bo. Vraigerchi chiu tia t ngoi c bc sng 265nm v 280nm dn ti ph hy qu trnh pht trin phi. Nh vy, s c ch qu trnh phn chia t bo c lin quan ti s hp th tia t ngoi ca phn t ADN bc sng 260nm v phn t protein bc sng 280nm. Nm 1914, Lep l ngi u tin pht hin ra hiu ng kch thch khi chiu x trng cu gai vi liu lng thp. Sau ny cc nh nghin cu tip tc pht hin trn cc i tng vi khun, vi sinh vt, to n bo, di tc dng ca tia t ngoi liu lng thp thy tc phn chia t bo tng ln t 110% n 170%. Theo Kimball,
hiu ng kch thch phn chia t bo c lin quan ti s hp th ca mt s loi protein v lipit. Pomper v Atvut li gii thch rng di tc dng ca tia t ngoi, c mt s t bo b tn thng nng tit ra mt s cht c kh nng kch thch s phn chia t bo, lm cho s phn chia t bo din ra nhanh hn. IX. Tc dng ca tia t ngoi ln axit nucleic Nhiu cng trnh nghin cu trn cc i tng nh vi sinh vt, thc vt v ng vt u i n kt lun l di tc dng ca tia t ngoi, tnh cht vt l ca phn t axit Nucleic b thay i. Khi chiu tia t ngoi ln dung dch ADN, Sugar nhn thy nht ca dung dch gim cn trng lng phn t ADN khng b thay i. Song nu chiu liu tia t ngoi cao hn s dn ti ph hy chui Polynucleotide, to nn cc Oligonucleotide, gii phng Timine v phtpht v c. Khi chiu tia t ngoi ln dung dch ARN cng gy ra hiu ng tng t nh vi dung dch ADN. Trung tm hp th tia t ngoi ca axit Nucleic l cc baz Nit. Khi chiu tia t ngoi ln dung dch axit Nucleic thng dn ti xy ra cc phn ng nh quang oxy ha, quang thy phn, quang nh hp. V d: Phn ng quang oxy ha xy ra vi Adenine theo dng sau: Adenine Hypoxantin N = CNH2 N = COH h hV + H CH CN O CH CN +NH3 2 CH CH N CN N CNH Phn ng quang thy phn xy ra vi Uraxin theo dng sau: Uraxin O HN
hV h + H O
2
Uraxin O HN CH2 CHOH
O NH O NH Phn ng quang nh hp xy ra gia hai phn t Timine theo dng sau: Timine Timine Nh hp Timine-Timine (T-T) O O O O CH3 CH3 NH h HN NH HN -CH3 H3ChV + H O
2
+ O NH NH O
O= NH H H NH
=O
Phn ng quang nh hp c th xy ra gia hai phn t cng gc hoc c th xy ra gia hai phn t khc gc. Khi chiu tia t ngoi ln dung dch Xitozine v Uraxin s xy ra s kt hp gia hai phn t cng gc, to nn nh hp Timine - Timine (T-T), to nn nh hp Uraxin-Uraxin (U-U), hoc gia hai phn t khc gc, to nn nh hp TimineUraxin (T-U). Setlau khi chiu tia t ngoi ln axit Polydezoxyinozin v axit Polydezoxyxitidin thy t l to nh hp T-T chim 50%, to nh hp T-X chim 40% cn nh hp X -X chim 10%. Phn ng quang nh hp gia hai phn t Timine l tc nhn chnh dn ti lm tn thng cu trc v ph hy tnh cht sinh hc ca phn t ADN khi b chiu tia t ngoi. Tm li, khi chiu tia t ngoi ln ADN xy ra cc qu trnh ni tip nhau nh sau: - u tin l tho xon hai chui ca phn t ADN vng c Timine (gi l tn thng vng). - Xy ra phn ng quang nh hp gia hai phn t Timine v c nh cu hnh bin dng ti vng b tn thng ca phn t ADN. - Hn ch hoc ph hy chc nng ca phn t ADN (nh mt kh nng tch chui nhn i ADN). X. Tc dng ca tia t ngoi ln protein Khi chiu tia t ngoi ln dung dch protein s dn ti lm thay i c, nht, lng... Chng t xy ra s thay i cu trc phn t protein. Cc protein hoc polypeptide c hot snh sinh hc nh enzyme, hormone, khng th, khng nguyn... khi b chiu tia t ngoi s gim hot tnh hoc mt hn hot tnh. Khi chiu tia t ngoi ln protein cc nh nghin cu nhn thy ph hp th ca protein (max=280nm) c lin quan ti ph hp th ca mt s axit amin nh tirozin, triptophan, xistein. Khi chiu tia t ngoi c bc sng 260nm n 280nm ln dung dch enzyme s dn ti kh hot tnh enzyme. Vi bc sng chiu trn th ch c tirozin v triptophan hp th. V vy, ngi ta cho rng ch yu do tirozin v triptophan hp th tia t ngoi nn b ph hy cu trc, dn ti lm mt hot tnh ca enzyme. Thc nghim xc nh c ch enzyme l tripsin th cn phi ph hy mt trong bn gc triptophan v mt trong su gc xistein cn kh hot tnh ca pepsin ch cn ph hy mt gc triptophan. Qu trnh hp th tia t ngoi ca axit amin theo dng sau: AH h AH* (trng thi kch thch) (axit amin) Tip theo xy ra qu trnh quang ion ha ca axit amin c hot ha (kch thch): AH* AH+ + e- A + H+ + eA + B A - B Axit amin do s quang ion ha mt mt proton (H+) v mt in t (e -) sau kt hp vi mt cht khc (k hiu l B) to thnh phc cht (A-B). Chnh phc cht ny lm bin i cu hnh phn t protein dn ti lm mt hot tnh enzyme. Tm li, s tn thng cu trc v s kh hot tnh ca phn t protein ch yu lin quan ti s hp th tia t ngoi ca cc axit amin thm nh tirozin, triptophan, phenilalamin. Ngoi ra, proton (H+) c hnh thnh do s quang ion ha axit amin cng lm t lin kt hydro gp phn thay i cu trc phn t protein.
Chng 9
PHNG X SINH HC
I. Cc ngun tia phng x
Trong t nhin, tia phng x c chia thnh hai loi sau: - Tia phng x c bn cht l sng in t c bc sng cc ngn (<10 A ) gm tia Roentgen (hay cn gi l tia X) v tia gamma (). - Tia phng x c bn cht l ht nh ht anpha (), proton (p), notron (n), dng in t (e-), dng pozitron (e+)... Nguyn t ho hc c k hiu A X , trong X l nguyn t ho hc, A l s khi bng Z tng ca proton (p) v notron (n) c trong ht nhn nguyn t (A=p+n) cn Z l nguyn t s bng s proton v xc nh in tch dng ca ht nhn. Trong bng tun hon Mendeleev cc ht nhn c nhiu hn 126 notron v nguyn t s ln hn 82 th u l 232 238 nhng nguyn t phng x. C 4 h phng x l Thri ( 90 Th ), Urani ( 92 U ), Urani
o
235 92
U , Neptuni
237 93
Np .
1. Tia phng x c bn cht l sng in t 1.1. Tia Roentgen Tia Roentgen c <10 A , c hnh thnh t ng phng Roentgen (hnh 9.1). Chn khng cao Thy tinh c bit
o
A 2
50100KV Hnh 9.1: ng phng Roentgen A: Ant (cc dng); K: Katt (cc m). 1: Cc in t; 2: Tia Roentgen c pht ra. Khi Katt c t nng v di tc dng ca in trng t ngun in cc ln 100KV, cc in t s thot ra khi Katt v "bay" cc nhanh v hng cc Ant. Khi in t p vo Ant, n s b dng li mt cch t ngt v nhng phn ln nng lng cho cc in t ca nguyn t cht lm cc Ant. Khi cc in t trng thi kch thch c mc nng lng cao (gi l E2) khi tr v trng thi ban u c mc nng
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com lng thp (gi l Eo) s pht ra tia Roentgen c tn s: =
E2 Eo (9.1) (h: hng s h
Planck) Khi tia Rnghen s c nng lng: E = h. (9.2)

C =
C: Vn tc nh sng v l bc sng nh sng. Khi in t p vo Ant ch c 0,2% nng lng pht ra tia Roentgen cn 99,8% nng lng chuyn thnh nhit, t nng Ant. ng phng tia Roentgen hot ng di in p 400KV, pht ra tia Roentgen c bc sng ngn nht l 0,03 A cn trung bnh l 0,06 A . 1.2. Tia gamma () Tia gamma c <1 A , c pht ra do s phn r ca ht nhn nguyn t Coban thnh nguyn t Niken theo phn ng sau: 60 Co 60Ni+1+2 Tia 1: C nng lng 1,17MeV (Mega electron Vn) Tia 2: C nng lng 1,33 MeV ( 1 MeV = 106 eV ) 2. Tia phng x c bn cht l ht 2.1. Tia anpha () Tia c pht ra do s bin i ca ht nhn nguyn t Radi (Ra) thnh nguyn t Radon (Rn) v pht ra tia theo phn ng:
226 88 Ra 222 86 Rn + pht ra ht
Ht chnh l ht nhn nguyn t Heli 4 He . Thng thng mi nguyn t phng x 2 pht ra ht c mt mc nng lng xc nh. Song cng c trng hp nguyn t phng x pht ra hai loi tia c nng lng khc nhau. Nh v d trn, s bin i ca 226 222 ht nhn nguyn t 88 Ra 86 Rn s pht ra tia c E=4,77MeV chim 94,3% v E=4,59MeV (Mega electron Vn) chim 5,7%. Ht mang in tch dng nn c kh nng ion ha cao nhng n li c khi lng tng i ln nn kh nng xuyn su yu. Trong khng kh, ht c nng lng t 410MeV, qung ng i ca ht ch t t 35cm. Vn tc ca ht c th t ti 3 vn km/giy. 2.2. Tia bta () C hai loi: - Bta m (-): Khi ht nhn nguyn t tha notron, n s chuyn v trng thi n nh bng cch chuyn notron thnh proton v pht ra in t (goi l bta m) v notrino () l mt th trung ha in, c nng lng rt thp. Phn ng xy ra nh sau: np+e-+
V d: S phn r ca phtpho phn ng:

32 15 P 32 16 S + e
( P ) thnh lu hunh ( S) s pht ra tia bta m theo

32 15 32 16
in t (e ) pht ra c nng lng cc i Emax=1,7MeV. Trn thc t tnh nng lng 1 trong phn r - ngi ta qui c ly gi tr nng lng trung bnh E bng nng 3 1 lng cc i. Trong v d trn ta c E = .1,7MeV0,57MeV. 3 in t mang in tch m v chuyn ng vi vn tc 3 vn km/giy. - Phn r bta dng (+): Khi ht nhn nguyn t tha proton, n s chuyn v trng thi n nh bng cch chuyn proton thnh notron v pht ra pozitron (e+), gi l phn r bta dng km theo th notrino () theo phn ng: p n + e+ +
()
V d: S phn r ca phtpho phn ng:

30 15 30 P 14 Si + e + +
( P ) thnh Silic ( Si) s pht ra tia bta dng theo

30 15
30 14
Pozitron c khi lng bng khi lng in t v mang in tch dng. So vi in t th pozitron rt khng bn, n rt d kt hp vi in t (gi l s hy cp) v pht ra hai lng t gamma c nng lng 0,51MeV. 2.3.Proton (p) Nm 1919, Rutherford pht hin ra hin tng ht i qua mi trng khng kh lm xut hin ht proton theo phn ng:
+ 14 N 17 O + p(proton ) 7 8
Proton chnh l ht nhn ca nguyn t hydro (H+), mang in tch dng.

2.4.Notron (n) Notron v proton l thnh phn cu trc nn ht nhn nguyn t. Notron l th trung ha in (tc khng mang in tch) v c khi lng xp x bng khi lng ca proton. Ngun notron thu c khi dng ht bn ph cc nguyn t nh nh Bery 9 Be theo 4 phn ng:
( )
+ 9 Be 12 C + n (notron ) 4 6
Cc ht notron c E t vi MeV gi l notron nhanh, E nm trong khong 10KeV500KeV gi l notron trung gian, E<10KeV gi l notron chm cn E=0,025eV gi l notron nhit (thng b hp th).
3. Nhng n v o lng c bn - n v Roentgen (k hiu l R): Roentgen l liu chiu tia Roentgen hay gamma, to ra c 2,08.109 cp ion trong 1cm3 khng kh iu kin tiu chun (p=1at, to=0oC) hay to ra c 1,61.1012 cp ion trong 1gam khng kh cng iu kin tiu chun.
Ngoi ra cn dng n v Culng trn kg (C/kg) vi cng thc qui i 1C/kg khng kh = 3876R. - Liu hp th phng x (Radiation absorbed dose: K hiu l Rad): Rad dng tnh liu hp th i vi tt c cc tia phng x. Rad l liu hp th 100ec bi 1 gam vt cht khi b chiu bi bt k tia phng x no. Cng thc qui i: 1ec=0,625.1012eV - n v Gray (k hiu l Gy): Gray l liu hp th 1J/kg mi trng vt cht khi b chiu bt k tia phng x no. Cng thc qui i: 1Gy=100Rad - Liu tng ng: Cng mt nng lng hp th nh nhau nhng li chiu cc tia phng x c bn cht khc nhau th s gy ra i vi ngi v ng vt c v nhng hiu ng sinh hc khc nhau. Nguyn nhn l do cc tia phng x c hiu ng sinh hc tng i (HUSHT) nh sau:
Tia phng x 1. Tia hay tia X 2. Tia bta () 3. Notron chm v notron nhit 4. Notron nhanh 5. Proton 6. Tia anpha ()
HUSHT 1 1 5 10 10 10-20
n v ca liu tng ng l Rem. (Roentgen Equivalent for Man) Rem l liu lng ca bt k tia phng x no, gy ra hiu ng sinh hc ging nh hiu ng sinh hc khi ta chiu tia Roentgen hay gamma vi liu lng l 1 Roentgen. Cng thc qui i gia Rem v Rad: Liu chiu (Rem) = Liu chiu (Rad) . HUSHT (9.1) Vi tia Roentgen hay c HUSHT = 1 th 1Rad=1Rem cn vi proton th 1Rad=10Rem. - phng x: n v o phng x l Curie (Ci). Curie l phng x ca mt ngun, trong mt giy c 3,7.1010 ht nhn nguyn t b phn r. n v nh hn l mili Curie (1mCi=10-3Ci) v micro Curie (1Ci=10-6Ci).
II. Tng tc ca tia Roentgen v tia i vi vt cht

Ty theo mc nng lng ca tia m n tng tc vi vt cht theo 1 trong 3 hiu ng esau: eh 1. Hiu ng quang in Quang in t Photon Hiu ng ch yu xy ra i vi tia X v c nng lng t 0,010,1MeV. V photon c nng lng thp nn khng th xuyn su m ch va chm vi in t (e-) vnh ngoi. Photon truyn ton b nng lng cho in t v nh bt in t ra khi qu o Nguyn t vt cht ca n tr thnh in t t do, gi l quang in t. Nng lng ca quang in t Hnh 9.2: Hiu ng quang in c xc nh: eE=h - Eo (9.2) in t h: Nng lng ca photon Compton e h E0: Nng lng cn thit nh bt h' Photon in t ra khi vnh (theo hnh 9.2 th in t vnh n=2). Quang in t c nng lng li tip tc gy ra s ion ha cc nguyn t vt cht khc. Nguyn t vt cht
Hnh 9.3: Hiu ng Compton
2. Hiu ng Compton Hiu ng ch yu xy ra vi tia phng x c nng lng ln hn 0,1MeV5MeV. Do c nng lng cao hn so vi hiu ng quang in nn photon khng nhng nh bt in t ra khi qu o ca n (gi l in t Compton), photon b mt mt phn nng lng v b lch hng (gi l tia th cp c nng lng l h'). in t Compton v tia th cp ty thuc vo nng lng m chng c, li tip tc gy ra s ion ha tip theo hay b mt dn nng lng trn ng i ca n. 3. Hiu ng to cp electron (e-) v pozitron (e+) Hiu ng to cp xy ra vi tia X v tia c emc nng lng E>1,022MeV. Khi photon s xuyn su vo ht nhn nguyn t, nh bt h + ra 1 electron (e ) v 1 pozitron (e ). Hai ht ny Photon c khi lng bng nhau nhng mang in tch e+ tri du nn d dng kt hp vi nhau, gy ra s
hy cp, gii phng ra nng lng E=0,511MeV di dng tia . Tia c to thnh li tip tc tng tc vi vt cht theo hiu ng quang in hay Compton.
Nguyn t vt cht
Hnh 9.4: Hiu ng to cp
III. Tc dng ca tia phng x c bn cht ht i vi vt cht

1. Tng tc ca bc x i vi vt cht Khi ht i qua mi trng vt cht n s tng tc vi cc nguyn t ca vt cht v nh bt in t ra khi nguyn t. in t b nh bt ra mang in tch m, gi l ion m cn nguyn t b mt in t nn mang in tch dng, gi l ion dng. l hin tng ion ha vt cht. Trong mi trng khng kh, nng lng cn thit to ra mt cp ion mang in tch tri du l 32,5eV. Mt electron vn (1eV) l nng lng ca mt in t c c khi qua th hiu 1 vn vi khong cch gia hai in cc l 1 xentimt. Nu nng lng ca ht cha nh bt in t ra khi qu o ca n th ht ch lm cho in t chuyn t mc nng lng thp ln mc nng lng cao, tc l ht gy ra hin tng kch thch in t. Qu trnh ht trc tip gy ra hin tng ion ha th gi l s ion ha trc tip cn in t sau khi b nh bt ra nu c nng lng cao li gy ra s ion ha nguyn t tip theo, gi l s ion ha gin tip. 2. Tng tc ca ht bta () i vi vt cht Ht cng ging nh ht , khi va chm vi nguyn t ca vt cht s gy ra hin tng ion ha hoc hin tng kch thch. Kh nng ion ha ca ht yu hn nhiu so vi ht v ht c in tch t hn so vi ht . V d: Ht v ht c cng mt mc nng lng l 1MeV nhng ht to ra c 7500 cp ion trn 1mm ng i cn ht ch to c 53 cp ion trn 1Cm ng i ca n. 3. Tng tc ca notron (n) i vi vt cht Ht notron trung ha in nn khng c kh nng ion ha v v th n c th xuyn qua nhiu lp v in t tin n gn ht nhn nguyn t theo 3 kiu sau y:
- Kiu khuych tn n hi, ch yu xy ra i vi notron trung gian c E=1keV500keV. Trong khuych tn n hi, notron truyn mt phn nng lng cho ht nhn nguyn t, sau notron b i hng v nng lng gim dn. - Kiu khuych tn khng n hi xy ra i vi notron nhanh c E=1MeV10MeV. Sau khi tng tc vi ht nhn nguyn t, lm cho ht nhn nguyn t b kch thch cn bn thn notron cng b i hng v nng lng gim dn. - Kiu thu ot notron xy ra i vi notron nhit c E=0,025eV. Khi ht nhn thu ot notron n s tr thnh ht nhn mi v trng thi kch thch. Sau ht nhn b v ra (hay b phn hch) thnh hai ht nhn km theo s pht ra ht notron.
IV. C ch chung v tc dng ca tia phng x ln c th sng

Tc dng ca tia phng x ln c th sng ln lt phi tri qua hai giai on sau: - Giai on ha l: Giai on ny c thi gian tn ti rt ngn, t 10-16 n 10-6 giy. Trong giai on ny cc phn t sinh hc chu s tc dng trc tip hoc tc dng gin tip ca tia phng x. Thuyt tc dng trc tip cho rng i tng b chiu x (c th l c th, m hay c quan, t bo, phn t nghin cu) s trc tip hp th nng lng ca tia v dn n tn thng hoc t vong. Thuyt tc dng gin tip li cho rng i tng b chiu x khng trc tip hp th nng lng ca tia m chng tng tc vi cc sn phm ca qu trnh phn ly phng x nc nn dn n tn thng hoc t vong. i vi nhng th nghim invitro th quan nim trn d phn bit cn vi th nghim invivo, cc nh nghin cu li qui c: Khi b chiu x nu l tc dng trc tip th cc phn t hu c s trc tip hp th nng lng ca tia v b tn thng cu trc nn dn n tn thng chc nng. Nu l tc dng gin tip th cc phn t hu c s khng trc tip hp th nng lng ca tia m tng tc vi cc sn phm ca qu trnh phn ly phng x nc v b tn thng cu trc nn dn n tn thng chc nng. Trong giai on ha l mt s phn t sinh hc quan trng nh enzyme, nucleoprotein b tn thng, ngi ta gi l nhng tn thng ha sinh. - Giai on sinh hc: Giai on ny thng ko di t vi ngy n hng chc nm sau khi b chiu x. Trong giai on sinh hc nhng tn thng ha sinh khng hi phc c s ko theo nhng tn thng chuyn ha, dn n nhng tn thng hnh thi v chc nng. C th tm tt c ch chung v tc dng ca tia phng x ln c th sng theo s sau:
Chiu tia phng x
Tc dng trc tip
Tc dng gin tip
Giai on ha l (10 6 s)
H2O O2 To ion v gc t do. Phn t b thch thch Tc dng ln cc phn t sinh hc quan trng v cc c quan t ca t bo
Ri lon chuyn ha v chc nng t bo Giai on sinh hc
Gy ra cc hiu ng sinh hc (tn thng hoc t vong)
Hnh 9.5: S tng qut v tc dnginh hc ca tia phng x
V. Tc dng ha hc ca tia phng x
Trong c th sng, nc va l mi trng va tham gia vo thnh phn cu trc ca t bo. ngi, nc chim trung bnh 65%, c bit c m nc chim 85%. Do vy, khi chiu tia phng x ln c th sng s gy ra hin tng ion ha cc phn t nc: H2O+hH2O+ + eCc ion
H2O + e- H2OH2O+ H+ + OHo gc t do H2O- OH- + Ho H2O + h H2O+ + e- H2O* Ho + OHo Gc t do Ho c thi gian sng ngn t 10-610-5 giy. Trong thi gian ny n c th tham gia vo cc phn ng: Ho + Ho H2 Ho + OHo H2O Khi c oxy ha tan trong nc s xy ra cc phn ng: Ho + O2 HO o 2
HO o + HO o H2O2 2 2
Ho + HO o H2O2 2
(hidro peroxit)
Hidro peroxit l mt c t i vi t bo. Gc t do OHo tham gia vo cc phn ng: - Phn ng oxy ha: Fe++ + OHo Fe+++ + OH- Phn ng tch nguyn t hidro ra khi phn t hu c: CH3 - CH2OH + OHo CH3 - CHOH + H2O - Ph v lin kt i: CH2 = CH + OHo OH - CH2 - CH CN CN Nh vy, di tc dng ca tia phng x xy ra qu trnh phn ly phng x nc dn ti hnh thnh nn cc trung tm hot ng l nhng gc t do, chng c kh nng tham gia vo cc phn ng rt cao. Ngoi ra, trong iu kin c oxy s dn ti hnh thnh nn H2O2 l mt c t i vi t bo.
Simpo PDF Merge and Split Unregistered Version - http://www.simpopdf.com VI. nhy cm phng x ca sinh vt nhy cm phng x ca sinh vt trn tri t vi mc tin ha khc nhau th rt khc nhau. Nhn chung nhng loi tin ha cng cao, s bit ha cng phc tp th c nhy cm phng x cng cao. nh gi nhy cm phng x ca sinh vt, cc nh sinh hc phng x a vo khi nim liu bn t vong (Lethally Dose) k hiu l LD50/30 l liu gy cht 50% ng vt th nghim trong 30 ngy theo di k t sau khi b chiu x. nhy cm phng x ca sinh vt th hin: Loi no c LD50/30 cng nh th nhy cm phng x cng cao cn ngc li LD50/30 cng ln th nhy cm phng x cng thp. Sau y l nhy cm phng x ca mt s sinh vt, xp theo th t t cao n thp.
Bng 9.1: nhy cm phng x ca mt s sinh vt (t cao n thp)
TT 1 2 3 4 5 6 7 8 9 10 11 12
Sinh vt
Ngi Ch Heo Kh Chut nht trng Chut cng trng Th ch, nhi, c Ra Rn Cn trng Virus
LD50/30 300R 300R 335R 500R 550R 600R 900R 103R 1500R 8.103-2.104R 104R - 106R 2.106R
Trong cng mt c th, cc t bo v m cng c nhy cm phng x khc nhau. Sau y l nhy cm phng x ca t bo v m xp theo th t t cao n thp: 1. Bch cu lympho; 2. Hng cu non v bch cu ht; 3. Ty bo; 4. T bo m tinh hon, m rut non, t bo trng, t bo ca cc tuyn, t bo ph nang, t bo ng dn mt; 5. T bo m lin kt; 6. T bo thn; 7. T bo xng; 8. T bo thn kinh; 9. T bo no; 10. T bo c.
VII. Cc hiu ng sinh hc lin quan ti s chiu x

1. Hiu ng tch ly Cc sinh vt khi b chiu x u th hin hiu ng tch ly. V d: Liu gy t vong i vi ng vt c v l 103R. Nu chiu x 5 ln, mi ln chiu 200R th sau khi chiu ln th 5 ng vt hp th 103R nn dn n t vong. Nh vy nhng tn thng sau mi ln chiu x sinh vt khng hi phc c hon ton m vn cn lu li nn sau mi ln chiu x tn thng cng nng thm, cui cng vt qu gii hn chu ng s dn ti t vong. 2. Hiu ng nghch l nng lng Cc tia phng x c kh nng gy ra cc hiu ng sinh hc rt ln ngay c khi chiu x liu khng cao (xt v mt nng lng th c gi tr nh). V d: Liu gy t vong cho ng vt c v l 103R, tng ng vi 84000ec hay 0,002cal/g, vi nng lng ny ch tng nhit 1 lt nc ln 1oC. gii thch hiu
ng nghch l nng lng, cc nh nghin cu u da vo thuyt tc dng trc tip hay thuyt tc dng gin tip. 3. Hiu ng pha long Khi chiu x liu lng xc nh ln dung dch enzyme n th hin: Nu tia phng x tc dng theo c ch trc tip vi quan nim mt ht (hay 1 photon) "bn trng"mt phn t enzyme s lm cho n b mt hot tnh th s phn t enzyme b mt hot tnh c lin quan ti nng enzyme lc ban u. Nu nng long th s phn t enzyme b mt hot tnh t cn nu nng cao th s phn t enzyme b mt hot tnh nhiu. Nu tia phng x tc dng theo c ch gin tip th s phn t enzyme b mt hot tnh ch lin quan ti s lng gc t do c hnh thnh trong dung dch m khng lin quan ti nng ca enzyme (tr trng hp nng enzyme qu long hoc qu cao). Cc nh sinh hc phng x chiu x vi khun E.Coli trng thi bnh thng c nc (c trng cho c ch tc dng gin tip) v trng thi kh (c trng cho c ch tc dng trc tip), kt qu thu c nh sau:
Bng 9.2: S phn t sinh hc ca t bo E.Coli b ph hy khi chiu x.
Phn t sinh hc ADN ARN Protein Lipit
No 2,1.104 4,2.104 4,7.106 4,1.107
N1 48 96 230 182
N2 14 28 370 43
N1+N2 62 124 600 225
No: S phn t trong 1 t bo. N1: S phn t b ph hy theo c ch gin tip. N2: S phn t b ph hy theo c ch trc tip. Vi kt qu ny th c ch tc dng gin tip chim u th hn so vi c ch tc dng trc tip (t l 3 tc dng gin tip: 1 tc dng trc tip). 4. Hiu ng oxy Hiu ng oxy th hin l trong khi ang chiu x, nu tng nng oxy th nhy cm phng x tng ln cn nu gim nng oxy th nhy cm phng x li gim xung. Nu ta tng hay gim nng oxy trc hoc sau khi chiu x th nhy cm phng x s khng thay i. V d: Chiu x chut bch liu 1200R iu kin oxy chim 21% th chut cht 100%. Ngc li, iu kin oxy chim 5% nu vn chiu x liu trn th chut sng 100%. Ngoi oxy th oxt Nit (NO) cng lm tng nhy cm phng x. S tng nhy cm phng x khi tng nng oxy ch trong mt gii hn nht nh. Nu nng oxy tng qu 20% so vi nng bnh thng th nhy cm phng x khng tng ln na. Hiu ng oxy th hin r i vi tia X, tia , tia nhanh cn khng th hin i vi tia , tia proton. Gray gii thch hiu ng oxy theo c ch tc dng gin tip ca tia phng x. Theo Gay, di tc dng ca tia phng x hnh thnh nn mt s lng ln cc gc t do v c (Ho, OHo) v hu c (Ro). Trong iu kin chiu x c oxy s hnh thnh nn cc peroxt v c (H2O2) v hu c (RO2) l nhng c t git cht t bo.
Alecxander li gii thch hiu ng oxy theo c ch tc dng trc tip. Khi chiu x chnh cc phn t hu c trc tip hp th nng lng ca tia v hnh thnh nn cc gc t do hu c (Ro). Trong iu kin chiu x c oxy to thnh peroxt hu c (RO2) l mt c t git cht t bo. 5. Hiu ng bo v phng x Dayli (1942) tin hnh chiu tia X ln dung dch enzyme thy rng: Nu thm vo dung dch cht thiour hay lu hunh th s lng phn t enzyme b mt hot tnh s gim xung. Baron (1949) pht hin xistein c kh nng hn ch t vong nu tim cho chut liu t 950mg1200mg/kg vo thi im 5 pht trc khi chiu x liu 800R. Sau ny cc nh khoa hc tip tc pht hin ra nhiu cht c kh nng ging nh xistein v c gi l nhng cht bo v phng x. nh gi hiu lc ca cc cht bo v phng x (BVPX), cc nh nghin cu a vo yu t gim liu lng (YTGLL) v c tnh theo cng thc: YTGLL = LD50/30 (l s dng cht BVPX) LD50/30 (l khng s dng cht BVPX) (9.3)
LD50/30: Liu gy cht 50% ng vt c s dng cht BVPX (hay khng s dng cht BVPX) trong 30 ngy theo di sau khi b chiu x (gi l liu bn t vong). Nhng cht BVPX bao gi cng c YTGLL>1. Cht c k hiu WR-2721 do M sn xut l cht BVPX c YTGLL cao nht, t 2,6. Theo c ch tc dng gin tip, cc nh nghin cu cho rng cc gc t do c hnh thnh do chiu x d dng phn ng vi cc cht BVPX hn l cc phn t hu c. Chnh v vy bo v c cc phn t sinh hc, nn hn ch c s t vong. Theo c ch tc dng trc tip, cc nh nghin cu li cho rng cc phn t sinh hc tuy trc tip hp th nng lng tia nhng li truyn cho cc cht BVPX tr li cu trc ban u nn cng hn ch c s t vong. Hoc cht BVPX lm gim nng oxy trong c th hay gii phng cht BVPX c sn trong c th, u c tc dng hn ch s t vong. VIII. Cc thuyt gii thch c ch tn thng do tc dng ca phng x 1. Thuyt "bia" Thuyt "bia" do Desauer (1922), Crouser (1924) v Lee (1935) a ra. Trn c s mt s th nghim chiu x dung dch enzyme, dung dch t bo thy rng: nng dung dch va phi khi thay i liu chiu x t thp n cao th s phn t enzyme b mt hot tnh cng tng ln lm cho ng cong t l % sng st c dng ng thng (hnh 9.6). Khi nng dung dch qu long th cc ht ca tia phng x s tng tc vi nhau hoc khi nng dung dch qu cao lm cho cc ht ca tia phng x s va p ln 2 hay ln 3 vi phn t enzyme b mt hot tnh (hay t bo cht). Khi t l % sng st s c dng hnh ch S (hnh 9.7).
% sng st 100 100
% sng st
Liu chiu
Hnh 9.6: T l % sng st dng ng thng
Liu chiu
Hnh 9.7: T l % sng st dng ch S
Cc tc gi cho rng s t vong ca t bo hay l s mt hot tnh ca enzyme xy ra khi chiu x l do ch cn va chm mt ln gia ht ca tia phng x vi "bia" ca t bo hay phn t enzyme. Theo cc tc gi th "bia" chnh l nhn t bo hay trung tm hot ng ca phn t enzyme. Thuyt "bia" ch gii thch c c ch tc dng trc tip ca tia phng x ln dung dch enzyme, dung dch protein, dung dch ADN... cn khng gii thch c hiu ng oxy. 2. Thuyt c t Da trn c s ng vt khi b nhim ha cht c hi s dn ti s t vong nn cc nh sinh hc phng x cho rng khi b chiu x, trong c th to thnh cht c no v chnh c t ny l nguyn nhn dn ti s t vong. Thc nghim xc nh trong c th b chiu x c hnh thnh c t l peroxt nhng n khng phi tc nhn u tin m l sn phm ca qu trnh phn ly phng x nc, c to thnh giai on cui ca qu trnh tn thng phng x. 3. Thuyt gii phng enzyme Trn c s th nghim hot tnh enzyme t bo sau khi b chiu x tng ln r rt nn Bacq v Alecxander (1952) a ra thuyt gii phng enzyme. Khi t bo trng thi sinh l bnh thng, nng enzyme trong t bo c kim sot theo c ch iu ha cm ng. Khi t bo b chiu x th tn thng trc tin l mng t bo, mng nhn v mng cc bo quan. V d: ADNase c trong ty th v lp th cn ADN c trong nhn nn khi mng nhn v mng ty lp th b tn thng lm tng phn ng phn gii ADN. Sau ny Duver cho rng lizoxom l bo quan cha loi enzyme, nu mng lizoxom b tn thng s gii phng ra cc enzyme git cht t bo. Thuyt gii phng enzyme cha gii thch c mng t bo c kh nng chu ng c liu chiu x rt ln ti 10KR, trong khi liu t vong ca ng vt c v ch 1KR? Mt khc, khi chiu x khng phi hot tnh enzyme no cng tng ln, chng hn ADNase, ATPase, ARNase hot tnh tng khng nhiu, thm ch khi tng khi gim. 4. Thuyt phn ng dy chuyn
Da vo thc nghim l cc loi m k thut c th xy ra phn ng dy chuyn v phn ng dy chuyn ny nhnh cng nh tc phn ng dy chuyn tng mnh di tc dng ca tia phng x nn Taruxop (1952) a ra thuyt phn ng dy chuyn. Taruxop cho rng: T bo trng thi sinh l bnh thng c mt h thng cc enzyme chng oxy ha nn cc thnh phn lipit ca t bo khng b oxy ha theo phn ng dy chuyn m oxy ha c s kim sot. Khi b chiu x, h thng enzyme chng oxy ha b ph hy nn xy ra phn ng dy chuyn: RH + h Ro + Ho Ro + O2 RO2 RO2 + RH ROOH + Ro ROOH RO + OHo Ro, Ho, OHo l nhng trung tm ca phn ng dy chuyn cng vi c t RO2 git cht t bo. Thuyt phn ng dy chuyn gii thch c hiu ng nghch l nng lng, nhng li khng gii thch c hin tng t bin di truyn c lin quan ti phn t ADN? 5. Thuyt cu trc chuyn ha Trn c s gia vi cu trc v qu trnh trao i cht ca t bo c mi lin quan mt thit vi nhau m Cudin a ra thuyt cu trc chuyn ha (1972). Thuyt ny cho rng tia phng x tc ng ln c vi cu trc ln qu trnh trao i cht ca t bo. S thay i ca c hai qu trnh ny c nh hng n nhau, dn n xy ra nhiu hiu ng gy tn thng hoc git cht t bo. Thuyt ny ging thuyt "bia" ch xem t bo l mt h d th, c nhy cm phng x khc nhau tng pha (dch nhn, mng nhn, cc bo quan...). Thuyt ny khc thuyt "bia" ch xem s xut hin tn thng nh l xc xut va chm ca tia phng x vi cc thnh phn quan trng ca t bo. Cudin c bit nhn mnh s bin i tnh thm ca mng nhn, ng vai tr quan trng trong qu trnh b tn thng do tia phng x gy ra. Song thc nghim li xc nh mng nhn khng phi l cu trc u tin b tn thng phng x. V th gi thuyt ca Cudin v nhng gi thuyt trn vn cha gii thch y c ch tn thng phng x u tin din ra c th sng. IX. Tc dng ca tia phng x ln phn t sinh hc Trong cc t bo sng lun c cc phn t v c v hu c nhng quan trng nht l cc cao phn t sinh hc nh ADN, protein, lipit, gluxt... Tn thng cc phn t hu c s dn n tn thng mc t bo, m, c th. Cc phn t hu c c th nhn nng lng ca tia phng x mt cch trc tip hoc gin tip. Biu hin chung cc phn t b tn thng do chiu x l: - Gy hin tng t mch dn ti lm gim trng lng ca phn t hoc khu mch s lm tng trng lng phn t. - Lm thay i tnh cht ha l dung dch b chiu x nh thay i nht, thay i h s lng, thay i im ng in ca phn t nghin cu... - Gy tn thng cu trc hoc ph hy cu trc phn t. - Lm thay i hoc ph hy chc nng sinh hc ca phn t. Tn thng ca phn t ADN di tc dng ca tia phng x l s thay i cu trc bc 1, 2, 3, nh hng n tnh cht di truyn ca phn t. Setlov v Doyle (1954) khi chiu
x ADN kh trong iu kin chn khng s gy ra s ph hy cu trc v lm mt kh nng ha tan ca ADN. Vi mu ADN kh, khi chiu x th hiu ng oxy th hin rt yu cn trong mu m lng oxy bng lng nc th hiu ng oxy th hin rt r. Khi khng c oxy th ch yu xy ra s t mch. i vi protein, khi b chiu x th thng dn ti cc hin tng: - Ph v lin kt peptit trong mch chnh hoc ph hy cu disunfit, dn ti lm gim trng lng phn t. - Xy ra hin tng khu mch l s dnh kt gia cc phn t protein vi nhau lm cho nht dung dch protein tng ln. - Ph hy cu trc phn t dn ti lm mt chc nng sinh hc ca n nh i vi enzyme th b mt hot tnh xc tc. Trong nghin cu hay dng enzyme hay vi khun l nhng i tng c xem nh mc phn t hay t bo v d nh gi l cn hay mt hot tnh sinh hc. Khi chiu x cc i tng ny, cc nh nghin cu rt ra qui lut logarit ca t s sng st (hoc hot tnh sinh hc) s t l nghch vi liu chiu x theo cng thc: N (9.3) ln = k.D No No: S phn t ban u N: S phn t cn hot tnh D: Liu chiu x k: Hng s (k t mt gi tr nht nh i vi mt loi phn t sinh hc xc nh, b chiu x bi mt loi bc x v trong iu kin chiu x nht nh). Cng thc (9.3) c th vit di dng: N=No.e - kD (9.4) Khi liu chiu x cng cao th s phn t cn hot tnh s cng gim. nh gi kh nng ph hy phn t ca tia phng x trong nghin cu ngi ta tnh tr s G (gi l hiu sut ha phng x). l s phn t ca cht nghin cu b ph hy khi hp th nng lng 100eV trong qu trnh b chiu x. Tr s G ca cc baz Nit trong phn t ADN khi b chiu x tia X hay tia c trnh by trong bng 9.3.
Bng 9.3: Tr s G ca cc baz Nit trong phn t ADN khi b chiu tia X hay
Tr s G Trong phn t ADN Trong dung dch c O2 Adenin 1,09 0,22 Guanin 1,09 0,17 Timin 1,89 0,40 Xitozin 2,15 0,38 Khi chiu x, s phn t b ph hy ch chim mt phn rt nh trong tng s cc phn t c trong t bo nhng cng gy ra t bin di truyn, lm tn thng hnh thi v chc nng, nu nng c th git cht t bo.
Baz Nit
X. Tc dng ca tia phng x ln qu trnh phn bo

Chu k sng ca t bo gm 4 giai on pht trin chnh: - Giai on u tin k hiu l pha G1 (Gap c ngha l pha hay giai on) khi t bo mi c hnh thnh sau qu trnh phn bo. pha ny t bo thc hin sinh tng hp protein v tch ly cc cht cn thit cho s tng hp ADN pha sau ng thi t bo tng v th tch. Pha ny ko di nht, c khi chim gn mt na chu k sng ca t bo. - Giai on tip theo gi l pha S (Synthesis ngha l tng hp) l giai on tng hp ADN. pha ny hm lng ADN trong nhn t bo tng ln gp i. - Giai on th ba gi l pha G2, l pha hu tng hp. T bo ch yu tng hp cc cht chun b cho qu trnh phn bo. - Giai on th t k hiu l pha M (Mitose c ngha l phn bo nguyn nhim) xy ra qu trnh phn bo. Bergone v Tribondeau t kt qu nghin cu rt ra nh lut, gi l nh lut phng x: " nhy cm ca t bo trc tia bc x t l thun vi kh nng sinh sn v t l nghch vi mc bit ha t bo". Trng hp ngoi l, t bo lympho bit ha hon ton nhng li c nhy cm phng x cao nht. Khi chiu x liu thp, i vi nhng t bo cha bc vo pha phn bo th b ngng hn qu trnh phn bo cn t bo bc vo pha phn chia th s phn bo s b chm li hoc ngng hn qu trnh phn bo. Nu t bo pha S tc l ADN c nhn i, nu t bo ngng phn chia v tip tc pht trin s dn ti hin tng a bi th. a s t bo ng vt v thc vt c liu c ch tm thi qu trnh phn bo l 50R. c bit trng ca mt s ng vt khng xng sng bin nh trng cu gai, liu c ch phn bo l 104R. Serclo s dng Poloni chiu ln bo t cy dng x thy rng: Hin tng c ch qu trnh phn bo xy ra c khi ch chiu x nhn hay ch chiu x nguyn sinh cht. Song liu gy c ch s phn bo khi ch chiu x nguyn sinh cht cao gp 20 ln liu khi ch chiu x nhn. Kt qu tng t cng thu c khi chiu x ln trng ca ong hay tm. Khi chiu x, s tn thng ca ht nhn c ngha quyt nh ti s tn thng hay gy cht t bo. iu ny li c Axtrakhob (1947) xc nh qua th nghim: Ly ht nhn ca t bo b chiu x em cy vo t bo khng chiu x b ly mt nhn th t bo ghp c nhn b chiu x cn nguyn sinh cht khng chiu x ch sng c 3 ngy. Trong khi t bo ghp c nhn khng chiu x v nguyn sinh cht b chiu x sng c ti 3 tun. Vai tr ca nhn trong hot ng sng ca t bo, thc cht l vai tr ca phn t ADN - phn t lu gi thng tin di truyn ca t bo. Do vy, khi chiu x, tn thng nhn s nh hng nghim trng ti hot ng sng ca t bo.
XII. Mt s ng dng ca ngun tia phng x ion ha

* Phng php chiu x gy ra bin d to ging mi Qua qu trnh s dng ngun tia phng x tc dng ln i tng sinh vt, gy ra bin d to ging mi, cc nh khoa hc rt ra nhng nhn xt sau: - Kt qu chiu x chu nh hng vo cc iu kin ca ht khi chiu x nh nhit , m, oxy, pH... Do vy, vi mt l em chiu x, phi chn cc ht ng nht v trng lng, kch thc, m, nhit , nng oxy, pH v.v... - Liu chiu x gy ra t bin to ging mi thng s dng t 10KRad n 100KRad.
- Sut liu lng nh hng n s lng t bin. Liu gy t bin gp i t bin xut hin ngu nhin gi l "liu gp i". Thc t xc nh khong t 15-30 Rad trong mt ln chiu cn nu chiu liu thp v thi gian chiu lu th "liu gp i" s ln ti 100 Rad. Khi tng liu chiu nh hn 1 KRad th s dng sut liu l 10-100 Rad/pht cn khi tng liu chiu ln hn 1 KRad th s dng sut liu l 500 Rad/pht. - Sau chiu x, tn s gy ra t bin c li l rt thp v a s trng hp t bin xut hin khng di truyn c cho th h sau. Qui trnh chiu x chn ging mi, tun t theo cc bc sau: 1. Chiu x ht gy ra t bin 2. Th nghim chn lc nhng t bin c li 3. Xc nh thuc tnh mi c li v mc n nh qua cc th h. 4. Nhn ging. * ng dng phng php chiu x tiu dit cn trng c hi C nhiu phng php tiu dit cn trng gy hi nhng s dng phng php chiu x li rt hiu qu v khng gy ra nhim mi trng. Phng php s dng l chiu x ln cn trng to ra nhng thuc tnh bt li cho cn trng nh cht yu, con c v sinh, con ci t trng hay c t l sinh con c cao... Kt qu thu c i vi rui gy hi cho ngnh chn nui pha Nam nc M v Mehico bng cch th 50 triu con c v sinh, cui cng ging rui ny b tiu dit. * S dng phng php chiu x kh trng Phng php kh trng dng tia bc x ion ha rt hiu qu v sch, c bit i vi nhng vi khun v nm mc, kh trng bng nhit cao khng tiu dit ht c. Trong cng tc kh trng bng nhit, ngi ta hay dng liu t vong 90% (k hiu l LD90) l liu cn thit dit 90% s vi khun em chiu x v cn gi l liu thp phn. Liu thp phn (LD90) rt thch hp cho vic tnh ton liu kh trng. Vi chng vi khun c No l s t bo ban u cn N l s t bo vi khun cn sng st sau chiu x liu D th liu chiu c tnh theo cng thc: D=LD90.(logNo-logN) (9.5) kh trng thc phm, nm 1960, Smit v Nate xut phng php xc nh trc tip liu LD90 vi qun th ban u l 109 bo t. Cch thc hin: chn 20 hp tht (hay nc hoa qu) cn kh trng, c cy thm vo mi hp 1ml hn dch cha 108 bo t Clostridium botilium ri ng hp li. Sau khi chiu x liu D xc nh ri cc hp vo t m 37oC trong mt thi gian (chng hn 1 thng). Sau 1 thng, kim tra xem c hp no b hng nh c c t hoc cn b nhim khun. Trng hp c c t, tch chit c t v tim phc mc chut th c tnh. Gi M l tng s bo t tim vo c 20 hp tht cn D l liu chiu x v N l s hp c cha c t. Liu thp phn c xc nh theo cng thc: D LD 90 = (9.6) log M log N
Trong thc t dit nm mc v vi khun, ngi ta hay p dng liu t 300KRad n 1 triu Rad. * S dng phng php chiu x bo qun lng thc, thc phm Phm vi p dng v liu s dng nh sau: - Chiu x liu nh hn 100KRad chng ny mm khoai ty, hnh, ti v.v..., lm tri cy chm chn nh chui, cam, qut..., tiu dit cn trng bo qun ng cc nh la, ng, lc, u tng v.v... - Chiu x liu t 100KRad 1 triu Rad bo qun tht ti, c ti, trng v.v... - Chiu liu t 1 triu Rad 5 triu Rad kh trng cc cht ph gia, gia v, ch phm enzyme v.v... * S dng ngun phng x trong y hc ht nhn Trong y hc ht nhn thng s dng tia gamma (), liu dng t 100 Rad 103 Rad iu tr bnh ung th. chn on bnh cc bc s thng s dng cc cht ng v phng x ghi hnh c quan, t c hng iu tr. V d chn on bnh v tuyn gip, bc s s cho bnh nhn s dng thuc c nh du It phng x (I131). Sau mt thi gian, phn ln It tp trung tuyn gip v pht ra tia , dng my ghi hnh nhp nhy s cho hnh nh ca tuyn gip. Nu tuyn gip to s mc bnh cng nng tuyn gip cn tuyn gip nh s mc bnh thiu nng tuyn gip. Cng vi xc nh mt s ch tiu sinh ha, cc bc s s c hng iu tr c th cho bnh nhn.
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ON SUY NGH (ch bin) L VN TRNG

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M U L SINH, S HNH THNH V PHT TRIN

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II. Mt s khi nim v i lng c bn

T CO chuyn tip thnh CO2 theo phn ng: CO +

s lt CO2 thi ra sau khi xy ho 1g thc n

LN ca xy 4,686 4,739 4,801 4,862 4,924 4,985

VII. Phn bit nguyn tc hot ng ca c th sng vi my nhit

VIII. nh lut II nhit ng hc

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Q v nhit lng cung cp cho h khng ch lm thay i entropi ca h m cn T

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nhn nhit l dSB=

A b gim). S thay i entropi ca ton h c xc nh: Q Q Q(TA TB ) dS=dSA+dSB= = TB TA TA .TB

IX. Tnh cht thng k ca nh lut II nhit ng hc

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1 . chnh l tnh thng k ca nh lut II nhit ng 50 2

X. nh lut II nhit ng hc p dng vo h sinh vt

Hiu sut () 36% 30% 55% 75% 90%

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XI. Nng lng t do

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= k.t lnC - lnCo = - kt

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III. Phn ng bc hai

gii phng trnh trn, ly tch phn 2 v:

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V. Phn ng thun nghch

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Ly tch phn hai v phng trnh (2.15) ta c:

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Khi hng s tc phn ng thun c xc nh theo cng thc: k1 = x* ax * ln a * t (ax * a * x ) (2.17)

Thay k1 vo phng trnh (2.14) s tnh c hng s tc ca phn ng nghch:

VI. Phn ng song song

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da a = (k1 + k 2 ).dt ao o ln a = (k 1 + k 2 ).t a = a o .e ( k1 + k 2 ).t ao (2.21)

Thay a vo phng trnh (2.19) ta c: db = k 1 .a o .e ( k1 + k 2 ).t .dt v1 = dt db = k1. a o .e ( k1 + k 2 ).t .dt

T phng trnh (2.19) suy ra: v1 db dc k1.a db k1 = = = k1.dc=k2.db v 2 dt dt k 2 .a dc k 2 Ly tch phn 2 v ta c:

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VII. Phn ng ni tip

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Thay gi tr 1 vo phng trnh (2.29) s thu c: kC C b = 1 o .e k1 .t + 2 e k 2 .t k 2 k1 Khi to = 0 th Cb = 0 2 = k1C o 1 = 2 k 2 k1

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Hnh 2.2: S thay i nng cht A, B, C theo thi gian

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VIII. Phn ng vng

IX. Phn ng bc khng

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XI. Phn ng dy chuyn

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Nu tch s .t rt ln hn 1 th nghim c tnh theo cng thc:

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XII. Nhit v tc phn ng

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