Physio L

.
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1 of 130
.

-MSc,
--MSc.
2 of 130
..............4
...................................................................4
.............................................................................................................4
..............................................................................................................5
...........................................................................................................5
...........................................................................................................6
....................................................................................................6
.................................................................................................................7
............................................................................................................7
....................................................................8
..........................................................9
..........................................................9
................................................................10
H ...........................................................11
............................................................11
.................................................................12
.............................................................13
..................................................................13
...........................................................14
H ..........................................................15
......................................................15
..................................................................16
.......................................................................................................16
1. ..........................................................16
2. .................................16
3. ...................................................17
4. ........................................................................17
5. ........................................................................18
6. .........................................................................19
7. ...........................................19
...............................................................20
...........................................................20
............................................................21
...............................................................21
....................................................................22
1. O .........................................................................22
2. ..................................................................22
3. ....................................25
4. ...............................31
5. ......................41
6. ...................47
7. ........................................48
8. ............................................................48
B .......................................................................................................49
...................................................................56
3 of 130

...................................................................56
..............................................................57
................................................................57
.................................................................59
.........................................................59
..........................................................60
......................................................60
..........................................................61
..........................................................61
........................................................................................................61
1. ...................................................62
2. .........................62
3. ...........................................63
4. ..................................................................64
5. .................................................................64
6. ...................................................................64
7. ...................................64
....................................................65
........................................................................................................65
1. ...............................................65
2. ......................66
3. ......................................66
4. .............................................................66
5. ............................................................66
6. ..............................................................68
7. ...............................68
............................................................68
...............................................................69
..............................................................70
.................................................................70
...........72
1. . & ....................................................72
2. ................74
3. .........................................91
4. ........................................100
H ...........................................................101
H .............................................................101
...........................................................101
..............................................................102
.........................................................................................................102
...............................................................................103
.......................................................................................................103
................................................................................................103
.............................................................................................................103
........................................................................................................105
..............................................................................................105
.........................................................106
4 of 130

............................................................107
........................................................107
..............................................................108
...........................................................108
H ..............................................................110
................................................................111
.............................................................111
.....................................................112
.........................................................................................................112
...........................................................................113
.......................................................................................................113
................................................................................................114
.............................................................................................................114
........................................................................................................115
..............................................................................................115
.....................................115
...................................................................................................122
1 .............................................................................................................122
2 .............................................................................................................122
3 .............................................................................................................122

...........................................................124
2.1 ........................................................................................................124
2.2 .................................................................................................124
2.3 ....................................................................................125
2.4 ..............................................125
2.5 .......................................................126
2.6 ................................................................................126
2.6.1 .............................126
2.6.2 .................................................127
2.7 ..................................................................................128
1 ........................................................................................................128
2 ........................................................................................................128
3 ........................................................................................................128
4 ........................................................................................................128
5 of 130

.

.

.

.

,
.
,
.

.
pH.

.
pH ,
.
.
6 of 130

.
.
. pH 6
4,5. pH
.
,
.
.

Ca, Mg, K .
,
.
,
, .

(grana)
.

. ,

.
, .
7 of 130

.

.

ATP.
-
.

, .

.
, NADH, NADPH, - .
.
> 1.000 mg.kg-1

. ,

.

1 6%
.

,
. 8.000-
8 of 130

12.000 ppm 3.000-8.000 ppm
.
.
. ,
.
, . ,

.
.
.
/. .

.

.
.
,
.

.
, .

.
9 of 130

, , Ca Mg
.
.

, Ca
Mg P S.
.
10 of 130

pH

P

.
.
Mo, Mo

.

.
.
.
7-50 mg.kg-1
.
10-27 g.kg-1. To
.

0,45-10%,
400 g.L-1.
0,1 1200 g.L-1
200 g.L-1. l(OH)4-, Al(OH)2+
11 of 130

Al(OH)3. H 10
1000 mg.kg-1 200 mg.kg-1.

.
.
,
, .
0,5 3 mg.L-1
. ,
.
,
.

pH
6,0 7,0. ,
,
-,
.

100 mg.kg-1 .
500 mg.kg-1

.

,
1/1000 .
pH 4,0
.
12 of 130
10

10 g.kg-1, 500 g.kg-1.

0,5 mg.L-1.
H

.
3 9 mg.kg-1.
40%
.
AsO42-, H2AsO4-, 3AsO4, AsO43-, H2AsO3-.
0,01 1,5 mg.kg-1.

.
.
.

2 mg.L-1 . 2
mg.kg-1 .
.
, -, -
- .

.
,
.

13 of 130
11

,
.
.
.
.

,
.
,
.
.
,

.
,
.

0,5 1,5%. 0,08%
.

,
.
.
20 g.kg-1.
14 of 130
12

. ,
.
VO43- VO3-
.
100
mg.kg-1 .
0,5 mg.kg-1
. 0,3-1 mg.kg-1.
2 mg.kg-1
.
<5 50 mg.kg-1.

100 mg.kg-1. ,

.

. 2 g.kg-1.

pH (
pH). M
.

, ().
.
100-3000 mg.kg-1 500 mg.kg-1. pH
15 of 130
13

,
.
,
.
.
,
. .
200 mg.kg-1 500
mg.kg-1. 15
mg.kg-1.

.
0,34 1,6. Be2+
Be(OH)+.
0,1-15
mg.kg-1 1,6 mg.kg-1.
0,4-1 g.L-1.
1 ng.L-1.

0,001-0,4 mg.kg-1, 0,001 mg.kg-1.
,
. Leguminosae Cruciferae
.
.
,
ADP
P. .
.
16 of 130
14

2 mg.L-1
, .
10-50 mg.kg-1 .
200 mg.kg-1
.
,
.
H
.
Bi3+ Pb2+
.
,
H 20
g.kg-1. 27 mg.L-1
. BiO+
Bi(OH)2+.

.
.
.

. Pinus sibiricus .
.
0,02
0,13 mg.kg-1.
17 of 130
15
1.

. ,
. ,

.
2.
:
,
film ,
.
.
-, .
.
18 of 130
16
3.

, ,
.
.
RNA --.
,
. ,
,
.
.
:
-, -
.
-
.

,
.

.
4.
20 mg kg-1
(ppm) . .
,
, (
).
19 of 130
17

.
, 1 6
ppm, 20 70 ppm.
latex 80 100 ppm.

, 5 10 .
, , , ,
, Brassica ( , , )
, .
5.

.

. , ,
. , .
,
, .

. , ,
.

. .
.
, . ,
.
,
.
, .
20 of 130
18
6.
.
, .
,
.
5 ppm .
10 ppm,
.

.
.
7.
7.1

,
.
.
7.2

. Ca:B .
21 of 130
19

7.3

. , Ca:B 500:1,
100:1.
,
.
.

pH.

(0,064 nm) (0,062 nm)
. ,
.
.
, 1,6--
.
,
,
.
.
.
0,5 mg.kg-1. Ga3+
Ga(OH)4-.

4 ,
22 of 130
20

(- 63 kcal.mol-1 56.7 kcal.mol-1) (0,134 nm
0,139 nm ). Ge4+
Ge(OH)4.
0,1 mg.kg-1
0,01 mg.kg-1.
40 .
.
,
. 5 mg.kg-1 GeO2
.

pH .
,
, 1 mg.kg-1 5-2500 g.kg-1.

.
,

. .
15 mg.kg-1 .

.

.
23 of 130
21

500 g.kg-1.
.

.
10 mg.kg-1.
.
,
.
.
10 mg.kg-1. 20 mg.kg-1 .
150 mg.kg-1
.

1. O
(SO42-)
(SO2) .

,
(Mills and Jones
1996).
2.
2.1
,
.
,

24 of 130
22

.
5 kg .
2.2
:
(1)
, , ,
(2) ,
(3)
pH < 4.0
(4)
pH > 7.
,
pH
.
.
2.3 T
90% ( )
:S = 10:1.

,
.
2.4
,
.
0.4 1.3 kg . ,
(2S)
25 of 130
23

. ,
, -. ,
2S - .

S4- S +
(e-). , S4-
.

.
Al- Fe-OH

.
pH
6.5.
.

.
,

, .
2.5

(24% S),
(13% S), (17.5% S) (18.6% S).
To -

.
26 of 130
24

,
pH .
2.6
,
. 3 5 mg
S L .

.
pH . pH

.
,
1 kg.-1 4.5 kg.-1
.
,
:S, SO4-S : -S .
N:S = 15:1 ,
3:1 17:1.
3.
3.1

. ,
.

.
.
,
27 of 130
25

(SH). , ,

, .
,
.
3.2

, .
..

. ,

(APS) ( 1).
.
,
.
O
O S O
ATP
O
AMP
O S O
O
SO42-
O
ATP
PPi
APS
1
.
28 of 130
26

.

. , APS
( 2) ( PAPS, 3.3) (
) (-SH)
APS PAPS (Schmidt and Jger 1992).
in vivo.
,
(SO32-)
(S2-).

(Brunold 1993).
H -,
.
()
,
() ,
(Miyazaki and Yang 1987).
29 of 130
27
O
R
AMP
S O
O
O
AMP
S O
R-SH
( PAPS)
HS
CH2
CH
COOH
NH2
(-SH)
. .

( 2).

(Fankhauser and Brunold 1978).
, (Fankhauser and Brunold 1979).

(sulfite) (Schmidt 1986).
,
30 of 130
28

(Schmutz and Brunold 1982).

. (Willenbrink
1964, Fankhauser and Brunold 1978). ,
,
.
()
.
3.3
,
,
.
APS,
(PAPS) ( 3).
PAPS .
,
.
3.4

,
(Rennenberg 1989).
, .

(Rennenberg 1989).
31 of 130
29
O
AMP
O S O
APS
O
ADP
O S O
APS
ATP
ADP
PAPS
. ,
,
.

.
3.5
H (Stulen
and Dekok 1993):
1. ,
32 of 130
30

2.
ATP,
3. S,
4. .

APS (Brunold and Schmidt 1978),
APS
R-S-SO3- ( 2).
(Brunold and Suter 1984).

,
.
.
4.
4.1
0.1% 0.55%
. Gramineae

Leguninosae Cruciferae .

0.18-0.19%, 0.25-0.30% 1.10-1.70%

Gramineae, Leguninosae Cruciferae (Deloch
1960).
4.2

(arschner 1995). (-S-S-)
(SH) .
.
, .
33 of 130
31

.
CO2,
, , N2
.
- (CoA),
. CoA

. CO2
.
-.
, 2% (R-S).
90% (Dekok and Stulen 1993)

(ergmann and Rennenberg 1993).

.
,
.
4.3
,
.
, .
(.. R1-C-S-C-R2),
(.. R-SH)
.

( 1)
.
34 of 130
32

, N:S 40:1
30:1 (Dijkshoorn and Wijk 1967).

.
(-
)
.

,

K ,

(Tomati and Galli
1979). H

(
) (Levitt 1980).
1
,
(Willenbrink 1967).
mg / 100 g

(g/mg )

+S
-S
5.8
0.9
2.8
27.0
13.5
3.8
48.0
3.5
6.5
5.2
4.4
H ( 4).
- ,
-
.
35 of 130
33

(Hell and Bergmann 1988). ,

-. -
(Rennenberg and Lamoureux 1990).
g2+
NH2
CH
CH2
CH2
CH
CH2
COOH
CH2
(GSH)
2 x GSH
SH
GSSG
4
.

. 50%
(Rennenberg and Lamoureux 1990).
(.. )
. 0.7
mmol.kg-1 ,
36 of 130
34

(Nieto-Sotelo and Ho 1986). H
.
-.
, -

,
.
H
(Schtz et al 1991)
(Schmidt and Jger 1992).

.

(Caknak and
Marschner 1992).
stress, (Smith et al
1990b).
(
),

(Schrder et al 1990).
4.5

,
(Rauser 1990).
(Grill et al 1987, Rauser
1990). O
37 of 130
35

- ( 2 10) .
-.

,
(Grill et al 1987).
2
24 0 3
M Cd2+ , .
10 cm (Tukendord and Rauser 1990).
Cd2+
()
(nmol / g )
0
3
43
44
421
156
3
230
Cd

(nmol/g
)
0
13.1
H ,
.
2.
3 .
.
, 1-2 .
0.05

(Tukendorf and Rauser 1990).
Silene vulgaris

(Verkleij et al 1990).

,
, (Rauser 1990).
38 of 130
36

4.6
O
, .
12 Da,

.
:
() /
() NADP/ (Schrmann 1993).
,
. ,

-1,6-
Calvin
(PSII) CO2.
4.7
To
, , ( )
( 1).
,
,
. ,
-

: (), -

.
- (-CO-CH3)

( 5). C2
39 of 130
37

,
, .
CH3
CH3
, TPP
CoA-SH
S-CoA
CO2
OH
+Mn2+
CO2
CO2
-
.
4.8
O
(Schnug 1993).
.

(Ernst 1993). A
,
.
40 of 130
38

4.8.1
A
S---.
Allium ( 6).
80% Allium
. , Allium cepa S--CH2-CH2-CH3).
- (R =
O
R
CH2
CH
COOH
R'
NH2
S-()-

6
.
H .

, - .
4.8.2
15 taxa ,
Brassicaceae (Schnug 1993).
- ( 7), R
.
41 of 130
39
S
R
R:
CH2
R:
CH
CH2
> Brassica nigra
CH2
> Sinapis alba
7
-.
, (Hglund et al 1991,
Wink 1993). ,
- Brassica napus.
,
.

(..
),

(Schnug 1993).
4.9
To - , .. ,

42 of 130
40
C6, .. ( 8) .

, 5%
(Schmidt 1986). T
.

,
.
CH2
CH
CH2
CH2OH
C6
8
.
5.
5.1
:
,
,
,
.
43 of 130
41
5.2

,
. ,
. ,
. ,
. ,
. , .
,
.
, .

.
5.3

, , .
,
. ,
.
,
4.4 S 2.0
(Edelbauer 1980).
5.4

,
(Karmoker et al 1991).
44 of 130
42

, (Burke et
al 1986). O
.
(Burke et al 1986), (Dietz
1989).
5.5

(Burke et al 1986, Dietz 1989).
A ,
. ,
,

( 1). ,
,
( ).
5.6
,

( 3).
(Freney et al 1978, Karnoker et
al 1991).

. ,
, .

(SO4-S) (Freney et al 1978).
45 of 130
43

3

(Ergle and Eaton
1951).
SN
(% )
mg SO42-.L-1
g /
SO4-S
oS
3-
0.1
1
10
50
200
1.1
2.4
3.4
4.7
4.7
0.003
0.003
0.009
0.10
0.36
0.11
0.12
0.17
0.26
0.25
1.39
1.37
0.06
0.03
0.10
2.23
2.21
1.19
0.51
0.45
0.96
1.28
2.56
3.25
3.20

.

(Freney et al 1978).

.
, .
(Robson and Pitman 1983).

.
5.7

(DeBoer and Duke 1982). ,

46 of 130
44

(Anderson and Spenser 1950).

(ara et al 1987). H

,
.
5.8
,
( 1).
,
( 1),

( 4).
4

(Wrigley et al 1980).
-S
(% )
+S
0.25
-S
0.10

nmol / 16 g -
21
7
11
5
27
34
33
93

(Randall and
Wrigley 1986). ,
47 of 130
45

(Castle and
Randall 1987).

30%,
36 71% (audet et al 1986). ,

23% (Grabau et al 1986b).
H
.

(Arora and Luchra 1970). ,

,

(Ewart 1978).
5.9

Brassicaceae .

( 5).

,
.
,

. ,
,
(Schnug 1993).
48 of 130
46

.
5

Brassica juncea (Schnug
1993).

mg S /
1
15
45
405
1215
g
80
208
285
261
275

mg /100 g
3
8
31
53
52
6.

. ,
.
,
.

.
,
( ,
).
.
49 of 130
47
7.
() (F)
(As), (),
(), (Pb), (Se) (Fe).
7.1
.
,
S:N = 1:15 ( ) 1:14
1:17 . S:N
, S:
.
0.5% 3%.
,
.
,
,
.
8.

0.7 mg SO2.m-3.
(Sekiya et al 1982a)
SO2 (Sekiya et al 1982b) ,
(H2S)
.
2S ,
50 of 130
48

SO2 ,
(Sekiya et al 1982b).

, ,
.
SO2
.
SO2
0.5 0.7 mg SO2.m-3 .

SO2.
, SO2
,

(Schnug 1993).
,
(Murphy
and Boggan 1988) .
Anderson A.J. and Spenser D. (1950). Sulphur in nitrogen metabolism of

legumes and non-legumes. Aust. J. Sci. Res, Ser B3, 431-449.
Arora S.K. and Luchra Y.P. (1970). Metabolism of sulphur containing
aminoacids in Phaseolus aureus Linn. Z. Pflanzenernaehr. Bodenk. 126,
151-158.
Baudet J., Huet J.C., Lesaint C., Mosse J. and Pernollet J.C. (1986) Changes in
accumulation of seed nitrogen compounds in maize under conditions of
sulphur deficiency. Physiol. Plant. 68, 608-614.
51 of 130
49

Bergmann L. and Rennenberg H. (1993). Glutathione metabolism in plants. In
Sulfur Nutrition and Assimilation in Higher Plants. (L.J. DeKok, I.
Stulen, H. Rennenberg, C. Brunold and W.E.Rauser, eds), pp. 109-123,
SPB Academic Publishing, The Hague, The Netherlands.
Brunold C. (1993). Regulatory interactions between sulfate and nitrate
assimilation. In Sulfur Nutrition and Assimilation in Higher Plants. (L.J.
DeKok, I. Stulen, H. Rennenberg, C. Brunold and W.E.Rauser, eds), pp.
662-675, SPB Academic Publishing, The Hague, The Netherlands.
Brunold C. and Schmidt A. (1978). Regulation of sulfate assimilation plants. 7.
Cysteine inactivation of adenosine-5-phosphsulfate sulfotransferase in
Lemna minor L. Plant Physiol. 61, 342-347.
Burke J.J., Holloway P. and Dalling M.J. (1986). The effect of sulfur deficiency
on the organization and photosynthetic capability of wheat leaves. J. Plant
Physiol. 125, 371-375.
Brunold C. and Suter M. (1984). Regulation of sulfate assimilation by nitrogen
nutrition in the duckweed Lemna minor L. Plant Physiol 76, 579-583.
Cakmak I.and Marschner H. (1992). Magnesium deficiency and high light
intensity enhance activities of superoxide dismutase, ascorbate peroxidase
and glutathione reductase in bean leaves. Plant Physiol 98, 1222-1227.
Castle S.L. and Randall P.J. (1987). Effects of sulfur deficiency on the synthesis
and accumulation of proteins in the developing wheat seed. Aust. J. Plant
Physiol. 14, 503-516.
DeBoer D.L. and Duke S.H. (1982). Effects of sulphur nutrition on nitrogen
and carbon metabolism in lucerne (Medicago sativa L.). Physiol. Plant. 54,
343-350.
DeKok L.J. and Stulen (1993). Role of glutathione in plants under oxidative
stress. In Sulfur Nutrition and Assimilation in Higher Plants. (L.J.
DeKok, I. Stulen, H. Rennenberg, C. Brunold and W.E.Rauser, eds), pp.
125-138, SPB Academic Publishing, The Hague, The Netherlands.
52 of 130
50

Dietz K.J. (1989) Recovery of spinach leaves from sulfate and phosphate
deficiency. J. Plant Physiol. 134, 551-557.
Dijkshoorn W. and van Wijk A.L. (1967). The sulphur requirement of plants as
evidenced by the sulphur-nitrogen ratio in the organic matter. A review of
published data. Plant and Soil 26, 129-157.
Ergle D.R. and Eaton F.M. (1951). Sulfur nutrition of cotton. Plant Physiol.
26, 639-654.
Ernst W.H.O. (1993). Ecological aspects of sulfur in higher plants: the impact
of SO2 and the evolution of the biosynthesis of organic sulfur compounds
on populations and ecosystems. In Sulfur Nutrition and Assimilation in
Higher Plants. (L.J. DeKok, I. Stulen, H. Rennenberg, C. Brunold and
W.E.Rauser, eds), pp. 295-313, SPB Academic Publishing, The Hague,
The Netherlands.
Ewart J.A.D. (1978). Glutamin and dough tenacity. J. Sci. Food Agric. 29, 551556.
Fankhauser H. and Brunold C. (1978). Localization of adenosine-5phosphosulfate sulfotransferase in spinach leaves. Planta 143, 285-289.
Fankhauser H. and Brunold C. (1979). Localization of O-acetyl-L-serine
sulfhydrolase in Spinacia oleracea L. Plant Sci Lett. 14, 185-192.
Freney J. R., Spencer K. and Jones M.B. (1978). The diagnosis of sulphur
deficiency in wheat. Aust. J. Agric. Res. 29, 727-738.
Hglund A.J., Lenman M., Falk A. and Rask L. (1991). Distribution of
myrosinase in rape-seed tissues. Plant Physiol. 95, 213-221.
Grill E., Winnacker E.L. and Zenk M.H. (1987) Phytochelatins, a class of
heavy metal binding peptides from plants are functionally analogous to
metallothioneins. Proc. Natl. Acad. Sci. (USA) 84, 439-443.
Grabau L.J., Blevins D.G. and Minor H.C. (1986). Stem infusions enhanced
methionine content of soybean storage protein. Plant Physiol. 82, 10131018.
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Hell R. and Bergmann L. (1988). Glutathione synthetase in tobacco suspension
cultures: catalytic properties and localization. Physiol. Plant 72, 70-76.
Karmoker J.L., Clarkson D.L., Saker L.R., Rooney J.M. and Purves J.V. (1991).
Sulphate deprivation depresses the transport of nitrogen to the xylem and
the hydraulic conductivity of barley (Hordeum vulgare L.) roots. Planta
185, 269-278.
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54

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57 of 130
55

0,02 0,5 mg.kg-1
0,2 mg.kg-1.
30 700 g.kg-1 200 g.kg-1.

.
10 mg.kg-1 .
pH.
.
.

.
,
.
.
2-6 mg.kg-1
3 mg.kg-1. .
0,5 mg.L-1
, (0,1 mg.L-1)
.
I- IO3-.
58 of 130
56
0,01-3mg.kg-1
0,2 g.L-1
Cd2+, Cd(OH)+
0,1-1 mg.kg-1
,
.
.

0,2 9 mg.kg-1. K
3 mg.kg-1 .
,
.

.
.
pH.

,
. ,

.

. 1,65
0,8.
59 of 130
57

. H
0,3 26 mg.kg-1
4 mg.kg-1,
0,005 g.L-1. Cs+.
T .
.
.

.
0,03 0,4 mg.kg-1.

.
60 of 130
58

+
. -,
.

.

.
.
,
.

.
.

.

.
.
.

1,5 4% : (w:w) = 1:1.

pH
.

.
61 of 130
59

,
.
.
5-10 mg.kg-1.
, .
1-40 mg.kg-1
0,1-1 mg.kg-1 (0,5 N HOAc)
0,01-0,18 mg.L-1
Co2+ CoCO3
0,1 0,57 mg.kg-1
0,03 0,27 mg.kg-1
: 0,2 mg.kg-1

. To 12

, .
.
0,1
30 mg.L-1 .
6-143 mg.kg-1 .
.
62 of 130
60

.
10-55 g.kg-1 50 g.kg-1.

.
. Mn2+

. .
n2+ Mn3+

, ,
, RNA,
.
Mn ,

Hill .

.
.
40 200 mg.kg-1
(ppm), 10 50 ppm.
1.
2.
63 of 130
61

3.
4.
5.
6.
7.
1.
.
(Mg2+)
Mg2+
.

(, , )
. pH
pH
5.5.
2.

.

. ,
.
64 of 130
62
3.
3.1

15-20% .
70%
, .
3.2
..
) ( .. Mg2+
Mn2+) .
ATP ADP.

--,
CO2.
Mg2+ +

.
3.3
ATP
ATP
.

.

.
65 of 130
63
4.
0.15%
0.40%
5.
,
.
,
.
,
.
,
.
6.
7.
7.1

, , .
Ca:Mg = 2:1 K:Mg = 8:1
.
66 of 130
64

7.2

7.3
,
. ,

.

.
.
1.
2.
3.
4.
5.
6.
7.
1.
MoO4- .
67 of 130
65
2.
MoO4- .
3.
.

.

.
4.
0.15 0.30 mg.kg-1
(ppm). ,
.
5.

.
.

.
,
. Brassica,
68 of 130
66

.
, ,

.
.

. ,
.
69 of 130
67
6.
,
15 ppm .
5 ppm .
7.
7.1
.
,
.
7.2
.

.

.
.

.
70 of 130
68

. ,
.
pH
pH .
.
0,05 mg.L-1.
.
.
( 3%) .
M 30 mg.L-1
, 10 mg.L-1 . 100 mg.L-1
.
350 mg.kg-1 .
. 2 mg.kg. 2-6 g.kg-1
,
.

0,358 nm
0,331 nm.

.

.

,
.

71 of 130
69

,

.
, : )
, )
)
.

,

.
,
.
.
.

. 0,3
3,5 mg.kg-1 1,5 mg.kg-1 .
,

.

, .
.
.
72 of 130
70

.
3-300 g.kg-1
50 g.kg-1.
, , .
73 of 130
71
1. . &
2 .
. ,
? ?.
494 kJ mol-1
.
(1)
, .

(1) : :: :
(2) ,
.

(2) : : :
Pauling (3)

. Pauling

spin.
74 of 130
72

(3) : :
:
(2s)2(*2s)2(2px)2(2py=2pz)4(*2py=*2pz)2
2 - .
2( )
:
1.
*2p, 2-,

2( ). 2- -
1,5.
2. 22-,
22.
- 22- 1 (
).
3. 0,5. ,
.
4. 2 4 .
.

. ,
24 2-,
.
75 of 130
73
2.
2.1

.
,
, .
,
.

.
,
. ,

.
, ,
, .

.

[(NAD(P)H,
]
.
(original) , CO2 2 (
1):
-G
(1) (.. )
(.. CO2, 2)
76 of 130
74

.

,
. ,
.
:
1.
([]=e-+H+)
, CO2:
(2) CxHyOz ye- +yH+ + xCO2
2. 2 [] 2:
(3) 2e- + 2H+ + 1/2O2 H2O
?
(.. NAD+/NADH). K

().
( CO2 2)
.
. (3)
,
.
2, ?
.
(2) (3).
. ,

. T ,
77 of 130
75

2.
,

. ,
.

,
.
-,
(-) ,
.
2.2
,
.

:
(4) [CH2O] + H2O + O2 CO2 + 2H2O (G0 -480 kJ/mol CO2)
C6H12O6 + 6O2 + 6H2O
12H2O + 6CO2 (G0 = -2880 kJ/mol )

. , -
CO2 -.
2 ,
.
78 of 130
76

. 2,
CO2 2

.
? ( )
( )
NAD+/NADH .
2.3
,
(
) :
(5) C6H12O6 + 2ADP + 2Pi + 2NAD+
2C3H4O3 + 2ATP + 2NADH + 2H+
(G0 = - 80 kJ/mol )
? 2
4 . , 2
.
(3-- 1,3
-- + 2[]), NAD+ [].
Calvin
.
( ) 2.
NAD+. E
NADH

[], NAD+.
79 of 130
77

2.4

2
.
- [],
.
.
.
.
,
.
(
).
:
(6) C6H12O6 + 2ADP + 2Pi
2C2H5OH + 2ATP + 2CO2
(G0 = -160 kJ/mol )

(7) C6H12O6 + 2ADP + 2Pi
2C3H6O3 + 2ATP
(G0 = -120 kJ/mol )

(NADH)

.
,
.

2,
30g/L, Saccharomyces
120 g/L .
80 of 130
78

CO2.
(..
Granineae). ,
? .
2.5

,
.
.
CO2.
(
) ,
2 ( ). []

, H2O.

, ?
? ? .
(TCA)
Krebs.

1 :
(8) CH3CO-SCoA + ADP + Pi + 3H2O
2CO2 + 8[H] + ATP +CoASH
(G0 -100 kJ/mol )

.
.
TCA
,
81 of 130
79

, .
TCA ,
?? .
TCA
,
. (
DH)
2 2.
,
TCA
FAD.
, Fe
()
() .
.
, .
b c.
2 . ,
, ?
b ( ).
, (
).

. .

in vitro
( ),
2
.
82 of 130
80

,
.
,
c. ,

.
. ?
. CO
2 Fe2+ .
quantum .
2
CO
CO.
2.6
(CN-)

(Fe3+).

. ,
CN-.
CN-
(antimycin), .
.
CN-
.
Michaelis 2 (Km 20mol/L)
(Km 0.1mol/L). H
,
83 of 130
81

Araceae,
.
.
.
CN- ? ?
,

.
CN-
.
, ,
CO2.
CN-
.

2 CN-.
, CN-
2. , CN
CN- ? ?.
2
CN-. ,
CN-.
2.7
:
, b560
c552,
3.
1 ATP 2
(G = 32 kJ/mol = 165 mV z=2). H
84 of 130
82

:
(9) 3NADH+FADH2+3H++11ADP+11Pi+2O2 11ATP+3NAD++FAD+4H2O
(G0 = -230 kJ/mol O2)
TCA
, ,
:
(10) C3H4O3 + 2.5O2 + 3H2O
3CO2 + 5H2O
(G0 = -1150 kJ/mol )

(11) 15ADP + 15Pi
15ATP
(G0 = 440 kJ/mol )

:
(12) C3H4O3 + 2.5O2 + 15ADP + 15Pi
3CO2 + 2H2O + 15ATP
(G0 = -710 kJ/mol )

, 40%
.
TCA 38 .
20 .

( CO2 ).

2
( /). 2
e- (1/2 2). , 2 , 3
NAD+/NADH.
CN- /=1 ( ) 0 (
),
85 of 130
83

. , /
. ,
50-75% .
ADP/ATP
, 2 ( ).
2

. ,

2,4 .

2.

, Mitchell.
,
:
1. +
2.
+ ..
3-4 + 2 e-.
, +/ 1/3 ?
NADH.
-250 mV, 80%
.
3.
+ +.
86 of 130
84

4.
.
.
3+/.
5. +
.
(
),
.
I, III IV
.
.
,
,
TCA .
+,
-. ADP .
, -, -
.
,
. ,
NAD(P)H
NAD(P)H ?
. ? ,
( ).
87 of 130
85

2.8.

NAD(P)H ?
, 2. ,
NAD(P)H
,
NADH-
c,
b,
2.

.

, NAD(P)H.
CN-
.

.
. ,
Fe3+ Fe2+
.

.
2.9 -

, ,
.
-
.
: (1) (
88 of 130
86

) (2)
NADPH. NADH,
NADPH , NADPH

(.. ).
-
. .
, CO2 1/3 CO2 TCA .
2.10

, .
, ( CO2 O2)
.
CO2
. CO2
.

CO2 2 ,
2.
2 10-20
ml/L O2 m(O2)
.
2 1bar. ,
DCMU CN-. CO2
, 2.

, TCA .
89 of 130
87
2.11

DCMU. CO2
.

ADP,
.
CO2 .
. ,

-. CO2 Calvin,
-, :
CO2 - 2,
- 3--.
/ - (RUBISCO):
+CO2
(13) P-Rubis
[C6] 2--
(14) P-Rubis
- + -
+2
: CO2
, 2
. 2
90 of 130
88

CO2
-. 2 CO2

0.4:1 (25
C).
RUBISCO
,
.
?
.
, RUBISCO
. ,
2.

. ?
-,
.

. ,
.
: (1)
2
( 22
) (2) .

. CO2
.
, .

.
91 of 130
89

, 2
CO2 .
CO2. H
C2 .
. ?
? ? Calvin.
CO2 30-60%
2. , ,
2.

2, ( /=3)
NAD+/NADH. ,

.

.

-. ,
.
,
,
C2 .
RUBISCO (..
CO2 2),
(.. )
.
(NH4+)
92 of 130
90

.

.

- .
(
)
.
3.
3.1 : CO2 2
( CO2 2)
, ,
.
,
2 .
.
2
C2,
CO2 2.

.

,
.
.
CO2 (molCO2/s),
2 (molO2/s).
.
,
?
93 of 130
91

.
CO2 2
. ,

in vivo
? .

, (..
, ) .
,
. , 2
(? ?).
.
.
,
.
.
, ,

( ).
, . ? P
.
Michaelis 2
6.10-7 mol/L (20 oC). 400
2 . ,

94 of 130
92

2 2,
. ,
(
), 2 .
2, .
2 104 ?
,
(.. ).
()
,
.
2
,
2. Q10 1,
2 2-3,
2
, ,
.
,
2 .
3.2
CO2 2
, . :
mol CO2 . t-1
(15) =
mol O2 . t-1
95 of 130
93

.

CO2:
1. :
C6H12O6 + 6O2 6CO2 + 6H2O
(16) = 6/6 = 1.00
2. .. :
C6H8O7 + 4.5O2 6CO2 + 4H2O
(17) = 6/4.5 = 1.33
3. .. triolein:
C57H104O6 + 80O2 57CO2 + 52H2O
(18) = 57/80 = 0.71
4. ( ):
C6H12O6 2CO2 + 2C2H5OH
(19) =
5. :
(20) = 13.5/36.5 = 0.37

0.97-1.17,
.
( ) (.. ).

<1.
.
Ricinus A=0.4. ? 60
96 of 130
94

, 0.6, . 20 ()
. 16.

2 .
1.3-1.5. ?
.
,
2.
1.7 NO3-.
..
.
3.3
2 -
.

2,
,
. ,
,
(, ).
, -
. ,
,

.
,
,
.
97 of 130
95

,
2.
( ) ,
,
? . 2,
( ), .
?
.
,
. ,

. 2
.
, ,
.
3.4

T

.
2
.
mRNA, .
,
,
.
Pasteur .
2
, ,
. 25 oC
98 of 130
96

1 bar 10-50 ml/L O2.

Pasteur. ,
, 2
( ).
, Pasteur ,
.
(.. Sinapis alba) Pasteur. ?
CO2 2.
Pasteur
2,

NADH/NAD+.

(
CN-), O2.

- -,
.
. ? ,

6-- .
,
.

- .
,
,
(
) ,
99 of 130
97

,
.
,
. , 6--
- .

. ,
ADH
. ,
NADH ( NAD+)
.
3.5

40%.
.

, 1660 kJ/mol
.

.
, , .
Araceae
( Arum maculatum.).
3.6

CN- ,
.
.
100 of 130
98

2 ,
2 ?
. 2 22
2, .
( ) .

(.. D-). H
-
.

,
( ,
), .
,
.
,
.
,
, .

,
.
, ,
2 . ,

- .
101 of 130
99
4.
4.1

.
.
4.2
1.

teflon . 0.8 V
,
:
: 2 + 2 2 + 4e- 4 : 4 g + Cl- 4 AgCl + 4eTo
.
3% C,
.

. 90%
10 .

,
0.8 V. , , ,
.
,
,
.
.
102 of 130
100
H

. .
UO22+, UO2(CO3)34- UO2(HPO4)22-.
<1 6 mg.kg-1.

.
Ca(UO2)2PO4.
H
,
.
pH, ,
(, , )
.
,

.

, .
.
6 mg.kg-1
.
4%, 0,5%.

103 of 130
101

.
. .

.

.

. .
100 g.kg-1

0,6 mg.kg-1. .

, ,
.
Fe3+ , Fe2+
. ,
. .
.
, Fe3+
Fe2+ ( .1).
Fe3+.
2., Fe3+
. Fe(III)
104 of 130
102

Fe3+
.

Fe(III)
.

- .
, -,
,
.
, ,
(NADP).

,
.

Fe 50 75 mg.kg-1 (ppm).
105 of 130
103

. , (
),
. ,
,
, .
.

.
,
.

.
106 of 130
104

- .
300-400 mg.kg-1 .
pH
pH (>7.0)
. - Fe(OH)3
pH HCO3-
.

. pH

pH .
.
P:Fe = 29:1 .

.
.1
107 of 130
105

,
.2.
, &

.

.

.
.
.
90
Sr
.
.

1 mg.kg-1
15,000 mg.kg-1.
.

108 of 130
106
.
4+
+3 +2.
.
,
.
1
mg.kg-1. 80
mg.kg-1,

.

. 200 mg.kg-1
, .
4,5 mg.kg-1
. .

,
.

0,1
40 mg.kg-1.
109 of 130
107

,

, .

0.05 35 mg.kg-1.
.

.
,
, pH,
,
.
.
. F o . .
30 mg.kg. (30-300 mg.kg-1)
, .

. ,
.
,
,
.
110 of 130
108

24-
.
100-1000
.
/
. AT
,
. pH
. HCO3-
2PO4- . 2PO4 , - ,
pH .

.

.
.
,
(
-) .

(-).
(85-90%)
.
0,2 0,5%.

ATP.
111 of 130
109
H
.
Cu2+ Cu.
O
, .
.

. ,
.

.

.
. -
.
, 2 22 2.
,
, .
.
, 3
2-
22. .
,
L--. -
.
2
20 mg.kg-1 (ppm). 3 7 ppm.
200 ppm
.
112 of 130
110

.
.
.
O
.
1-2 ng.kg-1. H 0,002
mg.L-1 Au(OH)4-. H
0,0005 0,125 mg.kg-1.

H
(+2 +6)
. Cr(III)
.
.

: 5-1000 mg.kg-1
: 65 mg.kg-1
10 g.L-1
0,2 g.L-1
CrO42-, Cr(OH)3
0,02-0,2 mg.kg-1
10 mg.kg-1
113 of 130
111

(VI) (Cr6+) ,
. Cr(VI) 100
Cr(III). pH.
pH 5,5 8,0.

,
.
. 5-15
mg.kg-1
150 mg.kg-1 .

,
.
,
(ZnCl)+ -n. Zn2+
.
95%

.
.

.
114 of 130
112

.
.
,
Fe3+ Cu2+.
Mn2+ Mg2+.
Zn
.
.
CO2
2 2CO3. E
,
pH +
CO2 1,5- .
,
.
. ,
, Zn2+
L- .
,
Zn Cu. ,
- 2-
,
2-.
.
115 of 130
113

n, RNA
RNA.
RNA.
grana . Zn
3-
D-
, L- D-,
c D-,
. Zn2+
, Mg2+, Mn2+, Cu2+ Ca2+.
-
.
.

15 50
mg.kg-1. E ,
.

, ,
. ,

. ,

,
.
116 of 130
114

Zn
,
Zn
200 mg.kg-1,

, .
, .
,
.
Zn Fe
.

1.1

,
117 of 130
115

, :
'
,
,
.

Jones et al. (1971).
,
.
1.2
,

, .
,
'.
.
: (1)
,
. (2)
, Mn,
118 of 130
116

(Jones, 1970). ,

,

. 1-1
,
. ,
,

'.

, ,
,
.
1.3

. ,
(
) :
2
).
119 of 130
117

,
,
. ,
,

.
1.4

,
.

,

(Jarrell and Beverly, 1981).

.
. ,

.
1.5 &

,
. ,

120 of 130
118

( ).

.

, 1-1.
1-1,
,
. ,
.

,
.

.

,
.

.
1.6
, '

.
15
.

121 of 130
119

.
1.7

.
, ..

.
1.8

.

,
.

.
12 ,
4
C. .

. ,
,
. ,
-,
.

122 of 130
120

4 C.
24 ,
.
1.9
Jarrell W. and Beverly R.B. (1981). The dilution effect in plant nutrition
studies. Advances in Agronomy 34: 197-224.
Jones, Jr., J.B. (1970). Distribution of fifteen elements in corn leaves.
Communications in Soil Science and Plant Analysis 1: 27-33.
Jones, Jr., J.B, Large R.L., Pfeiderer D.B. and Klosky H.S. (1971). How to
properly sample for a plant analysis. Soils and Crops 23: 15-18.
Mills H.A. and Jones Jr., J.B. (1996). Plant tissue sampling. IN: Plant Analysis
Handbook II, p.112-115, MicroMacro Publishing Inc., Athens, Georgia, USA.
123 of 130
121
1
.
:
1. ;
2. ;
3. ;
4.
.
5. ' ;
.
2

.
,

.
1. .
2.
,
.
3
11. .
124 of 130
122
1-1

. (: Mills & Jones, 1996)
1-1

(:
Jones,
1970)
(ear leaf)
1
2.93
3.20
3.65
2.75
1.95
0.22
0.22
0.23
0.25
0.18
1.22
1.26
1.19
1.44
1.23
Ca
0.48
0.75
0.58
0.48
0.35
Mg
0.39
0.40
0.41
0.45
0.40
mg /kg
B
11
25
14
Cu
12
10
10
Fe
96
110
102
75
57
Mn
73
124
79
62
49
Zn
22
30
22
22
18
125 of 130
123
2.1

. ,

.
: 1)

2)
.
, .
,
.
.

.
2.2
,
(Fe) (l).
(2%)
, (Sonneveld and van Dijk, 1982).

.
,
.
126 of 130
124

Fe
Al.
,
.

,
. ,

. ,
.
2.3
80 C .

,
, .

(freeze-drying).
2.4
,
,
.
, '
.
127 of 130
125
2.5

, ,
.
2.6
2.6.1
50 mg
. 2 mL 2SO4 10,
. 20
min 10 22 30%
.
5 20 min ( 1)
22 ( ). 20 min
.
22.
0.5 mL .
15 min .
(
2). ,

( 3).
10 mL. 3 ,
( 4)
( ,
5).
1. 22
. 10 0.5 mL .
128 of 130
126

22
.
2. ,
.
3.
, .
4. .
2.6.2
1 g
100 mL. 5 ml 3
2 mL ClO4 70% ( 5),

3-5 mL ( 6),
( 7 & 8).
10 mL (
, 4).
1. :
.
.
2.
.
3. ,
4 min 3-5 mL.
4.
,
,
.
129 of 130
127
2.7
1
(
1) , .
,
.
2

.
3

, ,
, , , .
4

.
130 of 130
128

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(Brunold and Suter 1984).

Gramineae, Leguninosae Cruciferae (Deloch

> Brassica nigra

> Sinapis alba

Anderson A.J. and Spenser D. (1950). Sulphur in nitrogen metabolism of

homogenates of cucurbit plants. Plant Cell Physiol 32, 733-736.

C6H12O6 + 6O2 + 6H2O

12H2O + 6CO2 (G0 = -2880 kJ/mol )

2C3H4O3 + 2ATP + 2NADH + 2H+

2C2H5OH + 2ATP + 2CO2

(G0 = -160 kJ/mol )

(G0 = -120 kJ/mol )

2CO2 + 8[H] + ATP +CoASH

(G0 -100 kJ/mol )

(G0 = -1150 kJ/mol )

(G0 = 440 kJ/mol )

3CO2 + 2H2O + 15ATP

(G0 = -710 kJ/mol )

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