Äîêóìåíò âçÿò èç êýøà ïîèñêîâîé ìàøèíû. Àäðåñ îðèãèíàëüíîãî äîêóìåíòà : http://www.phys.msu.ru/upload/iblock/231/2009-00-00-kirjanov.pdf Äàòà èçìåíåíèÿ: Thu Nov 19 11:41:45 2009 Äàòà èíäåêñèðîâàíèÿ: Mon Oct 1 23:52:50 2012 Êîäèðîâêà:

. . .





I I

: 03.00.02 ­



­

2009


. . : .­. ,

: . , .-. , : . . .

"17" 2009 . 17 . 00 . 501.002.11 . . : , ­1, 119991, , , 1, 2, , 5­19. . . . "17" 2009 .

501.002.11, .­.

. .



. . , , , . , . . . . . , , ­, . , . , , . . , . , . , I . , + P700 . . II. , . , , 1


. . , . , . , , . , , . , , . : 1. , . . . 2. II I , . 3. , + , P700 , . 4. II . , . , , I, , ­ . , , II. , 2


II. . , , , . . . II. II . ( ­ ). . 10­ , C O2 assimilation in plants: genome to biome (, 2008), (XVIII , , 2005) (XIX , , 2009), ­ 2008 (, 2008) ­ 2009 (, 2009), II, XIV, V XVI , (, 2005, 2007, 2008 2009 ), VI ­ ­ (, 2006). . , , . 137 . 5 , 35 123 .

3



. , . , , . , . . . , , ­ . , , , . , . . 1.1 . 1.2 , . 1.2.1 . 1.2.2 , . 1.2.3 . 1.2.4 . 1.3 . , , , , . , . 1.4 . . 2.1 , , . . ( , , ), , 4


. 1: , . ­. , , . 1. : ­ ; ­ ; ­ ; 5 ­ ­5­; ­ ­ 1,5­; ­ 3­ ; ­ ; i i ­ (i = 1, 2); i , i ~ ~ + i ­ , ~ (i = 1, 2); Q Q- ­ Q; U U - ­ U ; F F - ­ F . , , . . 2.2 . 2.3 . 1 , , . 1, : [2 ] M2 - y1 , [2 ] y1 , [2 ] N2 - y2 - y3 , ~ + [2 ] y2 , [2 ] y3 , [1 ] M1 - y5 , [1 ] y5 , [1 ] N1 - y6 - y7 , [1 ] y6 , ~ ~ ~ ~ + - - [1 ] y7 , [Q] Q0 - y4 , [Q ] y4 , [U ] U0 - y9 , [U ] y9 , [F ] F0 - y8 , ~ [F - ] y8 , [] y15 , [] A0 - y15 , [] y11 , [5] y12 , [] y13 , [] y14 [] y10 . 5


Mi Ni ­ (i = 1 I i = 2 II); Q0 , U0 F0 ­ Q, U F A0 ­ . : y4 Q0

y1 = a2

f

1-A

K1

(M2 - y1 ) - a2b y1 + g2b y2 (M2 - y1 )-

- g2f y1 (N2 - y2 - y3 ) + s12 y5 (M2 - y1 ) - s21 y1 (M1 - y5 ), y2 = g2f y1 (N2 - y2 - y3 ) - g2b y2 (M2 - y1 ) + b2b y3 y4 - - b2f y2 (Q0 - y4 ), y3 = b2f y2 (Q0 - y4 ) - wy3 - b2b y3 y4 , y4 = b2f y2 (Q0 - y4 ) - b2b y3 y4 + rb y9 (Q0 - y4 )- - rf y4 (U0 - y9 ) - p( y5 = a1
f O2 ,2) y4

,

1+A

K1

a2f y4 a1f Q0

(M1 - y5 ) - a1b y5 + g1b y6 (M1 - y5 )- (1)

- g1f y5 (N1 - y6 - y7 ) + s21 y1 (M1 - y5 ) - s12 y5 (M2 - y1 ), y6 = g1f y5 (N1 - y6 - y7 ) - g1b y6 (M1 - y5 ) + b1b y7 y8 - - b1f y6 (F0 - y8 ), y7 = b1f y6 (F0 - y8 ) - b1b y7 y8 - py7 y9 , y8 = b1f y6 (F0 - y8 ) - b1b y7 y8 - p1 y8 y10 - p3 y8 (U0 - y9 ) - p y10 = p13 y14 y15 - p1 y8 y10 , y y y y
12 14 15 (O2 ,1) y8

,

y9 = rf y4 (U0 - y9 ) - rb y6 (Q0 - y4 )y9 + p3 y8 (U0 - y9 ) - py7 y9 ,
11

= p1 y8 y

10 15

2 - 2p10 y11 ,

2 = p10 y11 - p11 y12 y15 , y13 = p11 y12 y

- p12 P [CO2 ]y13 ,
12

= 2p12 P [CO2 ]y

13

- p13 y14 y15 , + p13 y14 )y15 .

= (pf f 2 wy3 + pf f 1 py7 y9 )(A0 - y15 ) - (pAT P + p11 y

(1) , . 1. pf f 1 pf f 2 . N1 , N2 , M1 , M2 , Q0 , F0 , U0 , A0 pCO2 . AK 1 . (1) . AK 1 , s21 , s12 , p(O2 ,1) p(O2 ,1) , 6


. , , . 2.4 . BDF (backward differentiation formula) . 2.5 2.6 . . , , ­ ­5­ 3­ , : y1 = · · · = y9 = y11 = y13 = 0, y12 = 1, y14 = 2, y15 = 0.5. 2.7 , ­ , . 2.8 , , , , , . , (1) Ei , i = 1, . . . , 4 Zi , i = 1, . . . , 11 : y1 (t) = 10-8 M2 E1 (T ), y2 (t) = 10 y4 (t) = Q0 Z2 (T ), y5 (t) = 10
-8 -7

N2 E2 (T ), y3 (t) = N2 Z1 (T ),

M1 E3 (T ), y6 (t) = 10-7 N1 E4 (T ),

y7 (t) = N1 Z3 (T ), y8 (t) = F0 Z4 (T ), y9 (t) = U0 Z5 (T ), y10 (t) = Z6 (T ), y11 (t) = Z7 (T ), y12 (t) = Z8 (T ), y13 (t) = Z9 (T ), y14 (t) = Z10 (T ), y15 (t) = Z11 (T ), T = 103 t. (1) : 1010 109 1011 C, g2b = C1 , b2f = D1 , N2 N2 N2 109 1010 = D12 , a1f = 10F, a1b = 109 G, g1f = H0 , M2 N1 7

a2f = 10A, a2b = 109 B , g2f = b
2b

=

103 D1b , s21 N2


g1b =

109 1011 103 F1 , b1f = G1 , b1b = G1b , s12 N1 N1 N1 102 102 103 P1 , p = P , p3 = P3 , rf = p1 = F0 N1 U0 p10 = 103 P10 , p11 = 103 P11 , p12 = 103 P p(O
2

109 103 D21 , w = P2 , M1 N2 103 102 R , rb = Rb , U0 E0 3 12 , p13 = 10 P13 , = , Q0 = E0 N2 .

,1)

= 103 P

O 21

, p(O

2

,2)

= 103 P

O 22

, , . , 1 . , , , 1 . , , , y1 y4 . 3.1 , I: a2f = 0.0011, a b
2f 2b

= 3 · 109 , g
2b

2f 1f

= 4.5 · 1011 , g = 0.028, a
1b

2b

= 5 · 1010 ,
1f

= 2.5 · 1011 , b
1f

= 4500, a

= 5 · 109 , g

= 1011 , = 0.05, (2)

g1b = 5 · 1010 , b pf
f1

= 4 · 1011 , b
10

1b

= 10, w = 10, p = 100, rf = 6.28,
11

rb = 0.2, p3 = 1, p1 = 70, p = 0.0018, p
ff2

= 3.5, p

= 0.003, p

12

= 1.6, p

13

= 0.0009, N1 = N2 = Q0 = P [CO2 ] = 1,

F0 = U0 = 10, A0 = 50, M1 = M2 = 100. 3.2 , (2) . . 2 y9 , y7 , y8 , y11 , y13 , y15 0.0002, 0.0008, 0.0011, 0.00125 0.0014 a2f . , II. (2), . 3.3 , [1]. 8


1.5 1 y9, ~[U-] 0.5 0.1

0.12 y11, ~[]



0.08

0.05

0.04

0 1 0.8 y7, ~[~+] 1 0.6 0.4 0.2 0 0.3

0 0.04 y13, ~[] 0.03 0.02 0.01 0 12 y15, ~[]

y8, ~[F-]

0.2

8

0.1

4

0 0 100 200 300 400 500 600 ,

0 0 100 200 300 400 500 600 ,

. 2: y9 ( ), y11 ( ), y7 ( ), y13 ( ), y8 ( ) y15 ( ) , a2f = 0.0002 ( ), a2f = 0.0008 ( ), a2f = 0.0011 ( ), a2f = 0.00125 ( ) a2f = 0.0014 ( ). 0.1.

9


I . . I R ) y7 0.8 [1] -5 , p11=0.0085 p11=0.0067 0 +17 . 3. 0.6 p11=0.0034 0.4 P [1] 0.2 Q 0O , , ­. +17 °C P R 0 °C Q - -5 °C O 0 100 200 300 , y1...15 (t), i = 1, . . . , 15, . , - . 3: I p11 ( ), - [1] , . - y7 0.0085, 0.0067 0.0034. - ( ), - p11 ( ­ ). 5­, - (2), . - . (2), . , -5 (. 3 ), O, P, Q R, , [1], . 0 . P R, +17 . . 3 . , p11 = 0.0034, , O, P, Q, R . p11 = 0.0067 Q , p11 = 0.0034, p11 = 0.0085 . P, Q R p11 = 0.0034 p11 = 0.0067 , -5 0 . 3.4 10


, , , : a1f = 0.1, a2f = 0.01, a1b = 109 , a2b = 2.5 · 109 , g1f = 1010 , g2f = 1.8 · 1010 , g1b = g2b = 107 , b1f = 1011 , b2f = 6 · 1010 , b1b = 10, b2b = 107 , w = 450, rf = 10, rb = 1.6, p = 250, p3 = 6.3, p1 = p10 = 10, p11 = 0.05, p12 = 1, pf
f1

(3)

= pf

f2

= 0.018,

N1 = N2 = F0 = P [CO2 ] = 1, Q0 = U0 = 10, M1 = M2 = 100, A0 = 50. - 610-11 y1 0.05 y4,y9 - 410-11 t=20 II - 210-11 0.04 0100 . 0.006 y 4 0.004 0.03 0.002 . 4. y9(y1) 0 , y1 (t), y9 0.02 0.04 y4 (t) y9 (t) 0.03 0.02 0.01 0.01 t=20 0 10 . 0.1 y4(y1) 0.05 a1f t=0 . 4. 0 0 0 0.01 0.02 0 210-11410-11610-11 y1 , y4 y9 y1 . y4 y9 ( , . 4: y1 (t) ( . 4) - ), y4 (t) ( ), y9 (t) ( ) y4 (y1 ) ( y1 , - y9 (y1 ) ( ). . 4) . (3), y1 - 5 · 10-11 - . - . . . I (y1 ) 10 ( . 4). y4 11


10% , y9 2 (. 4, , , ). y1 (t) , y4 (t) y9 (t) ­ (. 4 ). , , ­ . . [2] , II . , . , . 3.5 , II : y1 (FO ) a2b + a2f a2f M2 . + g2f N2 + M1 s12 (4)

, . , , , (4) , . , , . . 3.6 , II: y1 (y4 ) = A(y4 ) = g2f M2 (C (y4 ) + D(y4 )) - E (y4 ), B (y4 ) = (g2f M2 (C (y4 ) - D(y4 )) + E (y4 )) -
2 2 4a2f g2f b2f 2

A(y4 ) + B (y4 ) , 2b2f g2f (a2b + M1 s21 )(Q0 - y4 )

(5)

(6)

M2 N2 w(Q0 - y4 )2 . (7)

(6)­(7) 12


C (y4 ) = a2f b2f (Q0 - y4 ), D(y4 ) = g2b N2 (w + b2b y4 ), E (y4 ) = (a2b + g2f N2 + M1 s21 )(M2 g2b (w + b2b y4 ) + b2f w(Q0 - y4 )).

(8) (9) (10)

(5) , , II, . II . , FO Fm II. 3 , , . , . , , . 4.1 , . : v v v v v v v
7,1 7,2 7,3 8,3 8,4 9,1 9,3

=v =v =v

8,1 8,2 9,4

= b1f · y6 · (F0 - y8 ) , = -b
1b

1 + F - 1 + F ~ ~+
7 ,1

v

-

;

· y7 · y8 ,

1 + F ~+ 1 + U ~+

-v

- 1 + F ; ~
7 ,2

= -p · y7 · y9 ,

-v

- 1 + U ; ~ F
-

7,3

= -v

9,2

= -p3 · y8 · (U0 - y9 ) , F
- v

+ U - F + U

v

8,3

-

;

= -p1 · y8 · y10 ,

+ - F + ;
8,4

= rf · y4 · (U0 - y9 ) , = -rb · y9 · (Q0 - y4 ) ,

U + Q- - U
9,1

v

-

+Q ;

U

-

+ Q - U + Q- .
9,3

v

yi (i = 7, 8, 9) . 5. (2) , . , 13


­ . 2 . ­ . 5 yi = 0. , yi ( ) yi (t). 10-12 10-4 v9,1 v9,2 U (. 5 ) , , y9 (0) = 0. 10-4 U - . 1 ~ v7,1 (. 5 ), F , I, II (a2f /a1f 0.039). v7,1 y7 , . 2. 10-2 1 , ~+ (v7,2 , . 5 ) (v7,3 , . 5 ) . , II, , I, Q , F . ­ Q- U (v9,1 , . 5 ) 10-2 , , U F - (v9,2 , . 5 ). F - U 10-3 . 10-2 1 y7 I (. 2 ) v8,4 F - ­ (. 5 ). 1 30 y7 , , v7,1 , v7,2 v7,3 . y7 , y8 y9 (. 2, , ). U , v9,2 , ­ v9,1 II. U - v7,3 (. 5 ), , , 1 . ~+ + 1 v7,2 (. 5 ). ~ y7 . 1 60 . ~+ , . 2, , 1 ~+ - U (. 2 ) . 14


1 3 2 1 0.025 0 -0.025 -1 -2 -3 3 2 1 0.04 0 -0.04 -1 -2 -3 3 2 1 0.005 0 -0.005 -0.15 -1 -2 -3 10 v
-12

100

200

300 v

400
9,2

500

600


y9

v v

9,1

9,3

v

9,4



1 y7


v

7,1

v v y8
7,3

7,2

v

8,1

v
8,3

8,4 8,2

v 1 , 100 200 300

10

-9

10

-6

10

-3

400

500

600

. 5: y9 ( ), y7 ( ) y8 ( ) , v9,1 , v9,2 , v9,3 , v9,4 ( ), v7,1 , v7,2 , v7,3 ( ), v8,1 , v8,2 , v8,3 , v8,4 ( ) . , , ­ 10-12 1 1 600 . ­ yi (t) vi,j (t) , . [-0.025, 0.025] [-0.04, 0.04] , ­ [-0.15, -0.005] [-0.005, 0.005]. .

15


Q- II. , y4 (t) y1 (t), . 2 ( ) . , F - U - . 1 ~+ 100 . I F U . 1 ~+ , , , F - , ­ 1 U - ( ~+ + v7,3 y7 ). 1 1 . ~ ~+ ­ , I, II 1 ~+ 1 . ~+ , p p3 . , I F U . I H2 . 2 2 ~+ ~ + - U + 1 U + 1 , , II ~ ~ , I. I , (. 2 ). a2f y13 (t) y15 (t) (. 2 ) . , (. 2 ) , 5 (. 1). a2f = 0.0008 a2f = 0.0002, 5 . , , ­ . , (. 2 ) ­, F - . 16


. a2f > 0.0011. 4.2 , II. , , , , II. , , (5), . , , ( FO ) ( Fm ) II. . 6 y1 (y4 ), (5) (1). (2) , . (5) (2). , y1 = y2 = y3 = 0, , . 0.0016 < y4 < 0.0035, . y1 (y4 ) . , . . 7 y1 (y4 ), (5). , . 2. y1 (y4 ) . 7 y4 . y4 = 0 , y4 = Q0 ­ . y1 (y4 ) y4 . (5) y1 y4 . , y4 (y1 ). II , FO Fm 17


110

-11

y1~[*] 2
-12

810

610

-12

410

-12

10

-6

10

-5

10

-4

0.002 10-3 - y4~[Q ]

0.003

0.004

. 6: y1 (y4 ), (5) ( ) (1) ( ). (1) (5) (2). , , . 10-3 c .
y 410
-11 1

210-11

0 0 0.2 0.4 y
4

0.6

0.8

1

. 7: y1 (y4 ), (5), a2f = 0002 ( ), 0008 ( ), 0011 ( ), 00125( ) 0014 ( ). (2). .

18


II. 4.3 . , . : 1. , , ­. 1 . . 2. I. I . 3. , I . I . 4. , II . .


[1] .. . .... . .­. / . . . . ­ ., 1980. [2] Genty B., Briantais J.M., Baker N.R. The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence// Biochimica et biophysica acta. 1989. ­ V. 990. ­ N. 1. ­ P. 87­92.

19


:
1. .., .. II // . 3. . . 2009. ­ 6. ­ . 65­69. 2. .. II : // II , . . . ... 2005. ­ . 68­69. 3. .. 10 10 : // XVIII . . . 2005. ­ . 24-25. 4. .. // VI - . . . .. . 2006. ­ . 93­102. 5. .. , // XIV , . . . . . . . 2007. ­ . 75-76. 6. .., .. // ­ 2008 . . . . . . . . 2008. ­ . 134­138. 7. .. 10 100// V , . . 20


: . . . , 2008. ­ . 7. 8. .., .. // XVI , . . . . . . . 2009. ­ . 10­11. 9. .., .., .. // 2009. . . . . . . . 2009. ­ . 113­116. 10. .., .., .. // XIX , . . ., . . . 2009. ­ . 25.

16.11.2009 . . 1.0 .­. . 1.0

70

60 â 90/16

, . ­, ­, . 21, . 5