1.3 Physical Foundations. Living Organisms Exist in a Dynamic Steady State, Never at Equilibrium with Their Surroundings

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1 1.3 Pyc Fu 21 FIGURE 1 23 Sm gub by m um. () Tw m cv: (R)-cv ( m pm ) ccc gc pm; (S)-cv (m cwy ) m k cwy. (b) Apm, c w u m usw, y gub by cp m b-g m, ug w y cgu w c cb m (R)-v (pm) 2 () (S)-v (cwy) L-Apy-L-py my (pm) (w) (b) L-Apy-D-py my (b) Mcu cgu c b cg y by bkg cv b. F cb m w u ubu ( c cb), ubu gup c b g w wy, gg m w c pp. y m bgcy cv. Mcu cm p m pc c b cg by bu g b, wu bkg cv b. Ic bw bgc mcu m vby pcc: y qu cmpmy mc bw cg mcu. 1.3 Pyc Fu Lvg c gm mu pm wk y v puc mv. T yc c ccu w c, k yc pc y cy, qu pu gy. Egy cum m bcum ympc p, g y cc cg. A g xp m qu gy, wu wc ucu c m vby bcm mg. I cu vu, c v vp gy c mcm cupg gy b m ug u my gycumg pc y mu cy u. g bcmy u, quv cmc m, m by wc gy xc, c, cum vg c. W c c cu gy cv k gy cv cx w mymc. Lvg gm Ex Dymc Sy S, v Equbum w T Suug T mcu c w vg gm k cc m gm uug. A Pmcum p, k c, ycy um bm, pp c cmp m uug, c y v c muy, ( ppxm) m c cmp c cy cgg uug. Aug ccc cmp gm cg ug m, ppu mcu w gm m c. Sm mcu, mcmcu, upmcu cmpx cuuy yz bk w cmc c vv c ux m gy ug ym. T mgb mcu cyg xyg m yu ug yu b mm w yz w p m; by x m y w v b g y pc by w mgb mcu. T guc yu g w yu m c m w ccug yu bm; b y v pcu guc mcu w v b

2 22 p 1 T Fu Bcmy cv mg cb x, pp w v b pc w uppy guc, yu b guc cc m c v w y. T mu mgb guc b m y c bcu y k c ju bc bkw, cump, cv m puc. T ccy cc u ymc y, y m qubum. Mg y qu c vm gy; w c c g g gy, bg cy w qubum w uug. W c bw xcy w m by y qubum. gm Tm Egy M m T Suug F cmc c ccug u, w c ym c puc p, v c m, mm mp, vyg w g pc. T ym uug g cu uv. I ym xcg m gy w uug, b. I ym xcg gy bu m w uug, c ym; xcg b gy m w uug, p ym. A vg gm p ym; xcg b m gy w uug. Lvg gm v gy m uug w wy: (1) y k up cmc u (uc guc) m vm xc gy by xzg m ( Bx 1 3, 2); (2) y bb gy m ug. T w mymc, vp m pyc cmy bu uy v bgc ym w, cb pcp cv gy: y pyc cmc cg, mu gy uv m c, ug m gy my cg. cumm uc gy, cpb cvg cmc, cmgc, mcc, mc gy w g ccy (Fg. 1 24). T Fw Ec Pv Egy gm y vg gm v gy, cy cy, m gy ug, wc m muc u c c u u. Pyc c bb g gy u v c m w cb x, mg gy-c puc uc guc ( ), c, uc g 2 mp: g (g-v uc 2 ) pyc c gm b gy y by xzg gy-c puc py pg c m- P gy Egy uc ccmp wk u vm (cmpx mcu uc ug, ) Sug () mc m w c u wk: cmc y mcc wk mc cc g g puc gc m (b) (c) Ic m (py) uug Mbm puc cmpu mp u mcu: 2, 3, 2, P 4 2 () Dc m (py) ym Smp cmpu pymz m m-c mcmcu: DA, RA, p FIGURE 1 24 Sm gy cv vg gm. Dug mbc gy uc, m ym pu uug (xp quvy py) c p gy cmpx u mcu c. () Lvg gm xc gy m uug; (b) cv m uu m gy puc wk; (c) u m gy uug ; () -puc mcu w gz g u, cg py uv. c m () c (c m) ym m cmpx mcmcu. W u quv m py p 13. ()

3 1.3 Pyc Fu 23 pc 2 m w, cb x, puc, wc cyc vm: gy (gy-yg x guc) Vuy gy uc c c b c w c m mcu, w w m g w ccmc p; uc, my gu w c by-v cc ccu. A c vvg c w xuc c: c xz ( c) uc (g c). g Mg Rqu Wk Egy DA, RA, p m mcmcu. I ug cmc gy m cv b bw ubu pym, c mu v gy ubu cc quc. I xmy mpbb m c mxu wu puy c g yp p, w uqu quc. T wu p c ppu mcu; bu ccg c w mymc, cy u w v-g uv: py uv cuy cg. T bg bu y mcmcu m mmc u, gy mu b upp ym ( c, c). T m cmp cmc ym xp py, S (Bx 1 3). Ay cg m ym xp py cg, S, wc by cv pv vu w m c. J. W Gbb, w vp y gy cg ug cmc c, w gy c, G, y c ym c b J. W Gbb, m qu: py,, cg umb k b; py, S; bu mpu, T ( g Kv). T gy G TS. W cmc c ccu c mpu, -gy cg, G, m by py cg,, cg k umb cmc b cv c bk m, py cg, S, cbg cg ym m: G T S P A pc ccu puy y G gv. Y c uc p gy mcu, uc p ucc c, wc gy m pv: mcu b m gy mxu mmc cmp. T cy u mymcy uvb, gy-qug (gc) c, c cup m c b gy (xgc c), v pc xgc: um gy cg gv. T uu uc gy cup bgc c gy by yy ppy b uc pp (ATP; Fg. 1 25; Fg. 1 15)., c P p ppy gup: Am c 888 pym P P 888 P P P P 2 G 1 pv (gc) G 2 gv (xgc) W c cup, um G 1 G 2 gv v pc xgc. By cupg gy, c b yz m m-c pym. Egy upg Lk Rc Bgy T c u bgc ( uy gy m vg ym) m by wc gy m u mbm g cpu cup c gy-qug c. I kg bu gy cupg, uu c mp mcc xmp, w Fgu A bjc p c p c mu p gy u v. I w p, g p gy p ppc gu. W ppp g--puy vc cup g bjc, m bjc, pu ww m g c m, ccmpg P P P Rb A 2 FIGURE 1 25 A pp (ATP). T mv m ppy gup ( pk) ATP, by bkg ppy b, gy xgc, c cup my gc c c ( xmp Fg. 1 26b).

4 24 p 1 T Fu Bcmy BX 1 3 WRKIG I BIEMISTRY Epy: T Avg Bg Dgz T m py, wc y m cg w, w u 1851 by Ru uu, mu c w mymc. A gu quv py vv c pbby c. wv, u c b u quvy by mp xmp, c mg pc py. T ky cp py m, m wy. 1: T Tk Rmz W kw m g m bg w c uu wk. Bu upp w u bu u k u w 100 ( ym ) kc ( uug ) w k c. A c, wk, bu p m k uug, g mpu uug ( kc) by my m mu u cmp qubum. A p p k kc pcy m mpu. T gy w c cc k w 100, py cpb g wk, pp. I quv gy p k kc (.., uv ) bu bcm cmpy mz ugu. T gy g vb wk bcu mpu w kc. Mv, c py kc ( uug) vb. W kw m vyy xpc v puy p bck m kc k mpu w 100 g. 2: T x Guc Epy y gy bu m. Abc (pc) gm xc gy m guc b m uug by xzg guc w 2, b m uug. T puc xv mbm, 2 2, u uug. I pc uug ug c py, w gm m y ug cg. Aug m py m p, py m k, u by qu x guc: W c p cmcy 2 ( g) Guc ( ) 7 mcu 12 mcu 2 ( g) 2 ( qu) T m c 1 mcu guc pu 6 mcu xyg, 7 mcu, m my p by x c w p 12 mcu ( ). Wv cmc c u c umb mcu w ubc cv qu gu puc, wc w m m mcu mvm mcu, u py, c. 3: Im Epy T wg pg m Juu, Ac IV, Sc 3, pk by Buu, w z mu c Mk Ay my. I mc m gm 125 Eg pb: c mu wk. T mu gy vb wk -gy cg, G; wy mw c mu gy, bcu m gy p c. T g v g bjc, g gy ( G) bjc ww g mu wk c b ccmp. w ppy cmc c? I c ym, cmc c pc puy u qubum c. W ym qubum, puc m xcy qu wc puc cv c. Tu cg cc c puc; y cv. T gy cg ym mv m qubum, w cg mpu pu, gv by -gy cg, G. T mgu G p pcu cmc c w m qubum ym y. Ec cmpu vv cmc c c c mu p gy, k umb b. I c ccu puy, puc v gy -

5 1.3 Pyc Fu 25 () Mcc xmp G > 0 G < 0 T m, Wc, k, u; m, vyg I bu w m. I w pg y vy, my mg. I y c cmpx quc v py, c py cc, mb, m p. Pm w Skp ug um u, vy c m. wv, 125 mkg up qu w w cmpy m, cc p, w wg bx, y wu v mg wv. b m T I w g I m 125 c m, bu y vy c py. Suc c v ccu m m gy; m b c gv py. I c, bc mmc c m y, wc bc pgmmg gc cmpu, cy mymc y. Lvg gm gy, m ucu, mmy c m u py-p. u m v m u W c k y Wk g bjc (b) mc xmp F gy, G Egc Rc 1: Guc P guc 6-pp L p gy p Exgc Rc 2: ATP ADP P Rc 3: Guc ATP guc 6-pp ADP G 1 G 2 Rc c G3 G 3 = G 1 G 2 FIGURE 1 26 Egy cupg mcc cmc pc. () T ww m bjc p gy c mcc wk. T p gy m vb by pu ww m, xgc pc (pk), c b cup gc upw mvm bjc (bu). (b) I c 1, m guc 6-pp m guc gc pp (P ) y puc g gy w c. F gc c, G pv. I c 2, xgc bkw pp (ATP) c v gc c w w c cup. T xgc c g, gv -gy cg ( G 2 ), gc c m, pv gy cg ( G 1 ). T c ccmp um c 1 2, -gy cg, G 3, mc um G 1 G 2. Bcu G 3 gv, v c xgc pc puy. c, u c gy, wc vb wk. Suc c xgc; c gy m c puc xp gv vu. Egc c qu pu gy, G vu pv. A mcc pc, y p gy xgc cmc c c b u ccmp wk. I vg ym m gy p cg py. I vg gm, mcc xmp Fgu 1 26, xgc c c b cup gc c v w uvb c. Fgu 1 26b ( yp gp c c c gm) u pcp cv guc guc 6-pp, p pwy x guc. T mp wy puc guc 6-pp wu b: Rc 1: Guc P guc 6-pp (gc; G 1 pv) (P bbv gc pp, P 4 2. D b cc bu ucu cmpu w; w cb m bk.) T c ccu puy; G

6 26 p 1 T Fu Bcmy pv. A c, vy xgc c c ccu c: Rc 2: ATP ADP P (xgc; G 2 gv) T w cmc c cmm m, P, wc cum c 1 puc c 2. T w c c b cup m c, wc w c w um c 1 2, w cmm m, P, m m b qu: Rc 3: Guc ATP guc 6-pp ADP Bcu m gy c 2 cum c 1, -gy cg c 3, G 3, gv, y guc 6-pp c ccu by c 3. T cupg xgc gc c ug m buy c gy xcg vg ym. A w, bkw ATP (c 2 Fg. 1 26b) xgc c v my gc pc c. I c, ATP mj c cmc gy c. K q G A Mu Rc Tcy Pc Spuy T cy cmc c g cmp c b xp qubum c. F c A bb 888 c D qubum c, K q, gv by [ K q [ A q] c [ D q] b q] [ B q] w [A q ] cc A, [B q ] cc B,, w ym c qubum. A g vu K q m c pc u c v b m cmpy cv puc. Gbb w G y cmc c uc -gy cg, G c ccc c pcc c m xp cc c puc: [ ] D] G G RT [ b (1 1) A ] [ B ] w [A ] cc A, ; R g c; T bu mpu. W c c qubum, vg c m c wk: G 0. F pc c, [A ] [A q ],, c puc, c [ [ ] c [D ] [ [A ] [B ] b [ A q] c [ q] [ q Dq] b K q B ] Subug 0 G K q [ ] c [D ] /[A ] [B ] b Equ 1 1, w b p G RT K q m wc w G mpy c wy (b K q ) xpg vg c c. Bcu K q xpmy mub, w v wy mg G, mymc c ccc c c. T u G G ju p m ( c p m). W K q 1, G g gv; w K q 1, G g pv. Fm b xpmy m vu K q G, w c gc wc c g cmp wc. cu bu p G : mymc c uc w w qubum c bu u g bu w qubum w b cv. T c gv by pm kc, pc w c p 6. Ezym Pm Squc mc Rc A bgc mcmcu muc mymcy b mmc ubu, y y kcy b: ucyz bkw ccu wy (v y c), m c m gm, mcu b. Vuy vy cmc c c ccu gc y bcu pc zym bcy, k cy, gy c pcc cmc c wu bg cum pc. T p m c() puc() m vby vv gy b, c cv b (Fg. 1 27), mu b umu y c pc. T bkg xg b m w gy qu,, xg b, cg g gy c puc. T g p c c gm p, c gy bw c gu cv gy, G. A zym cyz c by pvg m cmb : uc cmpm cmy, py, cg. T bg zym xgc, gy by bg uc cv gy c gy c c. A u cbu cy ccu w w m c b zym uc c c w pcc -

7 1.3 Pyc Fu 27 v c. T c by mgu pbby pucv c bw c. A u c v, cu p 6, zym-cyz c cmmy pc g m ucyz c. u cy, w w xcp, p. (I m c, RA mcu v cyc, cu p ) Ag w w xcp, c zym cyz pcc c, c c c cyz by zym. Tu zym qu by c c. T mupcy zym, pccy ( by cm bw c), ucpby gu gv c cpcy w cv b cvy. T cvy cuc cv gu cu pc. By wg pcc c pc gc pcu m, zym m w m gy c cu cv. T u zym-cyz cmc c c ucy gz my quc ccuv c, c pwy, wc puc c bcm c x. Sm pwy g gc u mp puc xc cmc gy cv m uu c; g gv, -gy-yg c g cbm. pwy w m pcu mcu cv m pgvy g m cmpx mcu, cug p ucc c. Suc yc pwy, F gy, G Rc (A) Acv b (, ) G c Rc c (A G uc Puc (B) FIGURE 1 27 Egy cg ug cmc c. A cv b, pg, mu b vcm cv c (A) puc (B), v ug puc m b c, c by g, gv -gy cg ( G). T gy qu vcm cv b cv gy ( G ). Ezym cyz c by wg cv b. Ty b m gy, bg gy c cvy uc cv gy m G uc G c. ( cv gy -gy cg, G.) B) G S u Ig S p bc c pwy (xgc) Smp ADP P 4 2 ATP wc vby qu pu gy, ccvy g bm. T v wk zym-cyz pwy cu cu mbm. ATP mj ccg k ( m) bw cbc bc cmp wk (w cmcy Fg. 1 28). T pwy zym-cyz c c m cu c p,, ug, ucc c vuy c vg gm. Mbm I Rgu Acv Bc Ecmy y vg c muuy yz u k cby,, p, ucc c mcu mp ubu, bu y pc pp qu by cu wk mpx bmcu Mcc wk mc wk Abc c pwy (gc) puc, pcu FIGURE 1 28 T c ATP mbm. ATP cmc m kg gy-g gyqug c pc. I c gu my cmy: /puc xgc c p/cum gc.

8 28 p 1 T Fu Bcmy c u y gv ccumc. F xmp, ug p c gw pcu p ucc c mu b m g qu, w gwg c qum pcu muc uc. Ky zym c mbc pwy gu c yp pcu mcu puc quy ppp cu qum c. pwy E. c y m c uc, cu p. T pwy v p cyz by v zym (A ug F p m pwy): A T zym 1 B D E F I c bg puc m uc p y, uu uc ccumu c cc b cyc cvy zym pwy, mmy wg puc uc. Suc bck b kp puc uz c mbc m bc. Aug ccp c pwy mp gzg u ug mbm, vmpc. T u mbc m c, my wc p m pwy. Mbm wu b b p mwk cc p pwy. A cg cc y mb wu v mpc pwy, g pp c wu uc w m ug c cu cmy. T k ug cmpx c mg m pwy quv m ug, bu w ppc, cu p 15, v bgu mp g v gu mbm. gu y w cy, zym, p c m mbc puc; ubc p 28. T xp g ( m DA cv p c) y zym y mbc c c. A y mu b k ccu w cbg v c cu mbm. Ambg cmp pcu w c gu ug jb y ju bgu. SUMMARY 1.3 Pyc Fu Iuc Lvg c p ym, xcgg m gy w uug, xcg cg gy m mv ymc y m qubum. Egy b m ug u by cvg gy m c w cmc b ATP. T cy cmc c pc w qubum c b xp -gy cg, G, wc w cmp: py cg,, py cg, S. T vb by qu G T S. W G c gv, c xgc g w cmp; w G pv, c gc g v c. W w c c b umm y c, G v c um G w p c. T pv wy cup c. T cv ATP P ADP gy xgc (g gv G), my gc cu c v by cupg m, ug cmm m, c. T -gy cg c, G, pyc c qubum c by qu G RT K q. M xgc cu c pc uu y bcu zym p cyz m. Ezym c p by bzg, ucg cv gy, G, cg c by my mgu. T cyc cvy zym c gu. Mbm um my cc c quc cv cu mb. Ec quc gu pv w c gv m xp gy y w cy. 1.4 Gc Fu Pp m mkb ppy vg c gm by puc mv cu g w y pc y. T cuy mp ccy, v m y, ucu mcu c gc m. Vy w c c cvz, v c cpp cv (Fg. 1 29), v uvv u y. Bu g vc gc uc vg gm v m y ucg v vy muc g p; my bc v y