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iv

essential cell biology second edition

Bruce Alberts Dennis Bray

Karen Hopkin Alexander Johnson

Julian Lewis Martin Raff

Keith Roberts Peter Walter

f'O Garland Science '-.:.:> Taylor & Francis Group

Garlad Sciece Vice President: Denise Schanck Managing Production Editor: Emma Hun Proofreader and Layout: Emma Hunt Senior Editorial Assistants: Kirsten Jenner, Frances Morgan Editorial Assistant: Lilith Wood Text Editors: Elizabeth Zayatz, Eleanor Lawrence Copy Editors: Richard Mickey, Christine Gregory Book and Cover Des.igo: Blink Studio, London Hlustrator: Nigel Orme Cover IUustration: Jose Ortega lndex.er: MerraH-Ross lntemational Ltd. Manufacturing: Nigel Eyre aod Marion Morrow

Bruce Albe rts received hs Ph.D. from Harvard University and is President of the Natiooal Academy of Sciences and Professor of Biochemistry and Biophysics at the University of Califomia, San Francisco. Dennis Bray received his Ph.D. from Massach1ses lnstitute ofTechnology and is curren tly a Medical Research Council Fellow in the Department ofZoology, University of

Cambdge. Karen Hopkin received her \1.0. in biochemistry from the Albert Einstein College o.f Medicine in the Bronx. and is a science writer in Cambridge, Massachusetts. Alexander Johnson received hs Ph.D. from Harvard University and is a Professor of Microbiology and lmmuology and Co-Director of the

Biochenistry and Molecular Biology Program at the Universi.ty of California, San Francsco. JuUan Lewis received his D.Pbil. from the University of Ox.ford and is a Pncipal Scientist at the London Research lnstitute of Cancer Research . Marln Raff received his M.D. from McGil1 University and is at the Medical Research Council Laboratory for Moleculik q;e Biology and Cell Biology Unit and in the Biology Departh'e1t at University College London. Keih Roberts received bis h.. from tbe University of Cambridge and is Associate Research Director at the John lnnes Centre, Norwich. Peter Walter receved his Ph.D. from The Rockefeller University e\ York and i.s Professor and Charman of the Department of Biochemisy and Biophysics a the Universiry of California, San Francisco, and a Investigator of the Howard Hughes Medical lnstitue.

by Bruce Alberts, Dennis Bray, Karen Hop.ki1, Alexander Johnson, Julian Lewis, Martin Raff, Keith Roberts, and Peter Walter.

AII righrs reserved. part of his book covered by the copyright hereon may be reproduced or used n any format any form or by any means-graphic, electronic, or mechanica1, including photocopying, recording, aping, or inforn1ation storage and retrieval systems-withou permission of the publisher.

Published by Garland Science, a member of the Taylor & Francis Group, 29 West 35th Street, Ne\York, 10001-2299, USA and 4 Park Square, Milton Park, Abingdon, 0Xl4 4RN, UK.

Pnted in Spain.

15 14 13 12 11 1 9 8 7 6 5 4 3 2 1

Essential Cel/ Biology 2 lteracive Artistic and Scientific Direction: PeerWaler Narrated by: )ulie Theriot Production Design and Development: Michael Morales

Llbrary of Congress Cata.loglng-in-PubUcation Data Essential cell biology I Bruce AJbers ... ( e a\.1. 2nd ed.

p.cm. ISBN 0-8153-3480X.-ISBN 0-8153-3481 -8 (pbk.) I . Cyology. 2. Molelar biology. 3. Biocl1emistry. I. AJbens. Bruce. QH581.2.E78 2003

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28 1:

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aa .

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chloroplasts

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Paramecium ( ) . ( . Barlow).

32 1:

z

a, z z {protozoas). Didnium, ,

z 150 m (10 ). . ' z { 1-30) . Didinium ,

. ( z ) . Didnum m z , z , .

z . 1-31 z. z :

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( 1-11). , , , m z

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n n a ' n n

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z zn , .

z , , v Homo sapiens

z zv . z

, n z Drosophila melanogaster ( 1-34) n . , n

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. (

.. Lewis).

36 1 :

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37

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S.pombe S. cerevisae

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.

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. ; vfi; , DNA: a . a , CDC2, , a .

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Proc. Natl. Acad. Sci USA 88:10885-10889, 1991).

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).

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n .

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2-4. (

) . . , , 60% . .

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(C-). vr (catios), r (as). z . , ()

z (Na + +) , ,

zu n (Mg2+ Ca2+ , ) . , Na + c- ' vn (ioic bod)

. a Na + c- ( , ' 1 mm 2 1019 ). uz f

( 2-8). NaCI, .

f n (ocoalet bods) . , : . f

, ( NaCI)

- (Na) (CI) (Na +) (C

() (NaCI)

()

54 2: u

'v (Na + c, . , . ,

.

zv

u . (molecule) (coalet bods),

. . - (2)

- , , z . z

, u , a . , n (bod legth) ( 2-9).

, z

- , , S, C- z . ,

. , z . , ,

, z . z, , z , , : , z z (i\. 2-5).

2-9. : . , .

, , ,

. . , ' a . , ,

.

!

( )

- n Et) ~ ( - ) !

0.074 nm

( )

r r 55

2-10. . ()

a , . () , a a ( ) ,

. , .v .. , 109. a (sticks and ball models)

.

, , . 1865 August Wlhelm Hofmann, " a

.

z , u u u

mzu l u uu . , z

( 2-10). , u n u. .

, u .

- - u. u u uu . u z (kcaVmole), u l u ' u u u. , 1 1 6 1023 u u (, 1 u ) , u uu 1 kcaVmole. 2-1 -

()

()

-

()

-

I -c -

56 2: u

: an der

Waals ( )

(nm)

0.15 0.25 0.30 0.35

x-"(f

u

2-12. a.

. +: , -: .

'

m2-3 : .

, z n .

z . , z

( z , . 2-1).

, . , ' C, , . , (polar structure) ( ) z ( ) ( ) . a z . , , -0-, z , --, . , , -C-H, -

, ( 2-12). n ru ' . .

' , , , z ( a) NaCI. n ' , , m ( 2-13).

m 70% . z . , zn z .

(20), . , n ' , . , '

a, ( 2-12 m 2-2). n -

58 2:

( } ( } ,

a , (hydroge bod). a a a : , u. , . , , , m z

. , , ,

z , . z, . ( } z

2-2. z . , ' - , z - . m , . ,

. , , z u , u u . , u (}. u . : , DNA, RNA 'i . ,

z u . ( 2-1},

C . , . . ua, z (. 11}. , z

2-13. .

m2-4

( ) : NaCI iu ,

' u

v' u . .

59

2-14. . () a

. {) a , a

. , a .

m z , , .

n , z

a \ f . ' , a, : , (+).

, . , z vf (3' ) ( 2-14). . ,

' \. f 30+ (acids).

30+ . 30+ -7

( 2-148). , 30+ + , + ' 30+.

z . , HCI, . , . -

()

~~~

()

a

~ + ~ + -

60 2: p

- (COOH) - ( . 2-2}. .

, , 30+ ( ) . , ,

30+ .

6 (base). z ' . '

30+, z , (-) , z

. , (NaOH) ( ailaiUn]

Na + -. NaOH z . z , ' . ( )

(2) . ' - : -2 + 20 --7 3 + + - (. 2-2).

- z ' 30+ , - 30+ . ' (10-7 ).

z vr. ' , 30+ z , pH {pH scale) ( 2-2). pH 7.0

pH. pH < 7, pH > 7.

rv pv

. . , , , z .

m2-5 . pH (

, pH=7,0), + ; , tz. . ,

' n 30+ pH; (:

18 zz 1 kg) .

61

z , z . , z . . , .

, C-C .

( 2-1). a . ' ,

, . f , ( - CH3), ( -} , ( -COOH}, ( - C=O}, ( -32-} ( -2} , . z m f

. a- z. (chemjcal groups) m 2-1.

a 100-1000, 30 . u . n v (, 1 }. '

. ' - , .

z m ( 2-2}. , .

f . n n -

62 2: u

22. rp .

(, )

70 1 1

0.4 0.4 1

0.2 26

( 1

20 250 100 100 50 ~300 ~3000

r , . , z ' r . , : , \

n , \ ( 2-15). r r , , z z

, v~ z ( 2-2).

r (sugars), vr , (CH20)v, 3, 4, 5, 6 7. , . , z C6H120 6 ( 2-16). r, fz : ,

u u

I]

AMINO:EA :

2-15. , , . , ,

. , ,

.

u 63

2-16. , . ' . () () , , .

- - . ()

. ( , .

, .

() (space-filling model),

. ( () : : , C: , 0: .

.

'

m2-6 ?

z 2-16 . . : .

yH20H

/-"- I c c '\. / " ' c- c

I ()

()

, . , f 2-3, z ' , z z, . ,

, D- L-, m . n , m . , z n f . 2-3 f n nf .

u u . n z , z, f z z.

, \ (, .) n\ f .

, - (3-50). , n ( ).

r . z -

- (codesatio reactio),

( 2-17). , , -

64 2: u

: : . : : , v ( 2-17).

( ), . , '

DNA, .

\zn . , , : ' , 13. : z ( z }

. : . , . , , r

rapvn , f z. , , n

: , . , -z

( 2-3}. , .

i, z: i, , z

\\J ( 2-4). i .

i . v yz:

. .

v v v vv (fatty acid), n\r ( 2-

18), . v , . (-COOH),

[

20

;

f

c

2-17. .

, .

( )~.

65

2-18. .

.

. . () . .

() . ( .

() () (

]

( ): ( -Coo-) t , n .

( 2-4). , , n r {amphipathic).

n vn: t KOl . , z , . , ,

c , . , n. .

n () () . t n

( 2-4).

, t z. n r.1.1.1v, ' ( 2-4). z , , ,

, ( 2-19). -

66 2: u

2-19. . u u , ' . () , a, .

r: a . () , , a ( ) ( Q. r : ' a . aaafa ,

, .

a z ,

. 13, a z .

, , ff. , z!. (

vn1, a ( rn ( 2-4).

r . m

. lf: , , , .

, , . , , {

2-4). , , , , z . m , .

. lvmn z m (. 16).

z m ! . u nz r ,

,

()

()

u 67

220. . , ,

' . , ,

.

2-21 . . () , pH 7,

. m

, () () (: , C: , 0: , : ).

~ ,~ .. u~\~u\~\~u~, ~~J.~.~J~~~hJ ..

u

m . z nlr8 lnv (lipid bilayer), ( 2-20,

11).

(amio acids) , : , , , - ( 2-21). m . .

, , .

nm

m ( 2-22). z -

"""'\"" ~ '"""'.. + ~ -H2N- C- COOH - H3N- C- COO

/ I 7 I

CH3 CH3 - \

(R)

vrv vrv

() () (

68 2:

. , m (Jll

v ) ( ). , , .

, n -

( 2-5). 20 ' , , z.

20 z,

n ' .

, m .

, , z , .

, ( ) 0- - ( 2-5). , . - . n - .

20 z . 20 z ( , , . 2-22). . ( 2-5).

4, .

DNA RNA

(ucleoside) z ' ( z). z z ( 2-6).

z (ucleotide). z z z, .

, z J: n ' (+) '

a m ia

I -

Phe H-9-CH2-G' O=C -

I - I

Ser H-C-CHOH I

O=C I - 0 I q

Glu H-C>-CH2 -CH2 -C O=C '-

- I /

Lys H-C-CH2-CH2-CH2-CH2-N-H+ O=t ' I a m auia

2-22. m . (

). . . m . , , , ,

, . m -

- - (. 2-5). , Phe-Ser-Giu-Lys ( FSE).

m2-7 ,

-

! - ;

' 69

2-23. () a-

. () . () (C: , : ,

: , 0: , : }. (dATP),

(} .

224. 1 . a ADP a

. (

( , }

( } , a . ADP a a . , a .

1

11

() i

()

-. . z . vrovn (C), vvn (}

ovpai?n (U} nvv n. ovavvn (G) avvn () vvn , . z n ( 2-6}.

. ' ' , vfv (adeosie triphosphate} ( 2-23},

' . nz m a .

avvprn ( 2-6}. n . . ( 2-24). 3, J

.

a ~ ~ - o--o-~-o-~-o-~-o-cH 2

an

--~--

~ a (PJ

- - + O-P-Q- P-0- CH2 ~

70 2: vp

m . vvvv .

() r (phosphodiester bod) ( 2-25).

, ,

( 2-6). -

. z z vt (riboucleic acids, RNA) , G, C . z z (

2 z r , . 2-6)

(deoxyriboucleic acids, DNA) u , G, C ( z RNA) ( 2-25). RNA , DNA .

DNA ( 2-7) .

' DNA RNA . , . DNA

( ), . , RNA .

' z z z ( G C n U) -

225. (DNA), .

, .

, 5' 3' . , (5' )

, (3' ) .

. , 5' . ,

: G-A-T-C.

G

c

u 71

'

mn2-8 m

01 ' ; n ;

2-26. . ( )

. .

( z , base-pairig) n z 5.

, z ( 2-26). z , u

. m , nz

( nv) ( 2-27 n : ;). , a n . , DNA RNA ( )

z . 1 J . z . , m vz. z

fn . , z , n n zn, n co2 , . n , (tubuli) , n n

(. 1-27), , DNA . , nn : -

a a '.

30%

f

70%

'-.:

f

:

vr, (4%) a (2%) DNA (1%)

R (%)

nl (15%)

(2%)

72 2:

. . , . ,

.

vv

(, ) m

, . m , ( 2-28 2-17). , z z .

, ,

z. , z m ' . , n . ' , u'v ( , 20

, . 2-5). , . , (sequece).

' 6 7. n , n ,

.

z . m, 200 2000 (20 20 20 .. . 200 ) , DNA 10,000 (

) ,

2-28. . a , a . ( a ) ().

-----ap

-- v -~-i

2-27. . a ( ) , .

m 2-9 ,

. .

' (CH2=CH2) m , ( ... -CH2-CH2-CH2 -CH2-CH2- .. ).

,

,

. ';

. + ..._

t-- Hp ---- 73

: ;

, . . 2000 ,

,

. , ,

(association forces) ( 2-29).

. , f i

( .. a , , ) . .

' , , , , m , . , - . . , m f.

. , ; ' -

, a ;

' . , ' . ,

()

tb\\1~ ( :,~,,/~~ ~~\\

() \ 229. ; 2000 , ' ,

() ' ().

~ I ,' ,i

()

()

l"

2-27. RNA

. , , , , .

410,000, r . , ,

.

z

, . , '

(coformatios), . , ,

z , : r niz ~ . a- , n ,

z n . , , RNA n

' ( 2-31). 1 ( 2-7) , . f , ,

n n . , n. z -

, m

76 2: v

.

, , . DNA. , z,

' . m ? an der Waa1s, \ , z . a der Waals

a a . az, z u . n 2-7.

u, u, u (. 2-2 2-7). u nn1p

. , .

v ' v v v v

, , , , z

, , ( 2-7). z

. , ' . , z, ' m , , .

, z

. , -

m2-10 v

;

'

77

\.\.~)1 / lf!l~

{{?J'' if~ ((~ '-/~ \)))

((C} ~) _r~~

~

' ,

~ :J.; ~ (, ~) ~ @

(( @~

((~~ @

~ ]

~ , , ~ 2-32.