1

X. .

2008

1.
1.1.


: ,
-, .
: (4.184 j.g-1.oC-1)
.
: (44 kj.mol-1)
.


.

.

.


.

1.2.


:
, -


,

3





.







-


.

.
(
)
1
.
.
(
)
.

1.1.

1.3.
j ,
:
j= j* + RTlnaj + vjP + mjgh

(1)

(j*: , RTlnaj: , vjP:

, mjgh: )

( w),
(i) , (ii)
, (iii)
, (iv) :
w = s + p+ m+ g

(2)

(s: , p: , m: g:
)

(3) vant Hoff: s
.

s = RTcs

(3)

(s: , MPa (MegaPascal)

-1
atm (10 atm = 1.013 MPa), c: (mol L ), R:
-1
-1
o (0,082 L atm mol ) T: 273 +
C)

w :
w =RT/Vwln(RH)

(4)

w = 4.623bars.deg-1X2.303TXlog(RH)

(5)

(R , Vw RH ).

1.4.
: ,
:
:

Dn/Dt = -Dax(Dc/l)

s = RTcs

(6)

(3)

VFR=(r /8)(p/)
4

(7)

.
Fick :
Dn/Dt = -Dax(Dc/l)

(6)

Dn/Dt:
(mol.m-2.sec-1), Da
mol
1 mol Dc/l

vant Hoff (3)
(osmotic potential) s ,

6
(osmotic pressure)

p,

, :
s = RTcs

(3)

p = RTcs

(8)

:
s = p

(9)

Osmotic potential = osmotic pressure

1.5.

.
,
..

:
(p:
)
(
). :
= -


( )
( ).
w, = s + p

(10)

(w, : s : p : )

7


.

.

1.2.
. () . () 0.1 . (C)
w .

w . (D)
w=-0.244 MPa
s=-0.732 MPa.
. w=-0.244
MPa s=-0.244 MPa
.
.

(.. , ,
) Poiseuille ( m-3/sec-1):

VFR=(r4/8)(p/)

(7)

(VFR (Volume flow rate): , r , n

p/ , )

. r2
m.sec-1.
q = (r2/8n) (p/)

(11)

5.
(aquaporins) .

.


.

1.3.

,
. 4

6 .
aquaporins

9
(
).

1.4. H

6
.

2. -

(transpiration)

.
H -
(transpiration stream) :

.

.

2.1.

.
100 H2O
.
.
:

10

2. 2.



(
H2O )

2.3.

.

.
= - cRT (10 20 atm 1-2 MPa )
wair=4,623 atm.deg-1.2,303T. ln(RH)
=25 oC RH=80/100 wair=307 atm 30 MPa

2.4.

(
)

11
2.1. H

-- MPa 20
50% ( -0.5 -95 MPa)

12
2.2. H --
20 50%

2.5.

.

2.3. H

:




m (
)


( 3 Pa).


.

13

2.4.

2.6.

:
( )
( )

2.5. H




.

14
2.7.

( ).

2.6:

(,
)

.

2.8.

CO2.

15

2.7. .

2.9.

Fick
Dn/Dt = -Dax(Dc/l)
Dn/Dt:
(mol.m-2.sec-1),
Da
Dc/l

16
=Da x(e-e)/l
e:
2

:
= (e-e)/R
R



.
.

.

.

.

.
2.8:

17

2.10.

Phragmitis communis
2.8: H


Phragmitis communis

2.11.
(
)

18

:


/ ,




...

2.12.
:



(, )
( )
:

2.13. :

K, , ,

19
Ca
(, )

2.14. -


.


.

.
.

2.15.



( )

20

3. , ,
(Development, differentiation, Growth)
3.1.
: +
:

21

:
.
, ,
(morphogenesis):
(, , ,
, ,
, ,
,
(embryogeny):

:
,


(senescence):

(abscission):

(dormancy):

3.2.

22

E 3.1. :

( ).

(
).

H ( ) :

-



Aridopsis.

23

E 3.2.

E 3.3. Arabidpsis

:
:
H
T

24
T

E 3.4.

:
,
,
.
.

25
(.. , ). 5
.

E 3.5.

3.4.

:

26


, , (
)






.

.
,
( 2 )
.
: , ( 6).
3.6.

Chlamydomonas

.


,




.

,
. 7
8
.

27

3.7. :

.

3.8. Theobroma cacao

.

3.5.

28















Arabidopsis
.

3.9. Arabidopsis
: GURKE (
), MONOPTEROS (
) GNOM (
).

4.
(Plant Growth Regulators)

29
:
( )

(, , )

4.1.
():

( ) (
M)

,
.

. :


,

,
.
,
.
.

,

.
(promoters) # (inhibitors) :
(
) ..: , ,
, .
, ,
.. .
(active) # (inactive) :
(
). .

(..

30
(stimulus):

.
(signal): To ( )
.
(Signal transduction): To


(receptor): (
),
(messenger):
.
(sensitivity): -
,
.
, ,
.




, :

. :
in vitro , , ,
, ,
..
:
, , ,
..

4.2.

(,
)

31
( ,
, )

4.3.

1. (auxins)
2. (gibberellins GAs)
3. (cytokinin K )
4. (ethylene Eth)
5. (abscisic acid ABA)
6. (bassinosteroids BRs)
7.

4.4.
:

//

(/)

/




,
: , ,
.

32

E 1.
) (: auxin, GA: gibberellins, K: cytokinins, ABA: abscisic acid)

1.

33

1.1.

(Darwin 1880,
Boysen-Jensen 1913, Paal 1919, Went 1926). Went

.

1.2.

,

.

E 1.1. .
,

1.3.

1.3.1.

1).

, , , , ,
, ,

.

.

34
,
.

E 1.2.

1.3.2.

.
:

35
1.
() (-).
2. pH -
.
3. , pH
(-).
4. ,

.

E 1.3.

1.3.3.
,
.
,
.

36

E 1.4. ()
Zea mays
Vicia faba ()

1.4. -

1.5. (/)

, .

, .
,
, .

1.6.
:

: ,
in
vitro
:


:

37
: ,
, in vitro

(/):
()

:

:
:
.

:

:
,

:
, :
Bromeliaceae ()
: (
)

D .

E 1.5.

38

in vitro .
()
Escalonia,
in vitro .
E 1.6.

()

Escalonia

39

E 1.7. .
.

( )
,

( ) .


.

.
.

.
.


.

40

E 1.8. Phaseolus vulgaris

.
. ():
. ():
. (C):
( agar )



.

+
,
.


.
.

41

E 1.9. . ():
( ) ():

+-
+-.

.

.

:
+-:
() (1 auxin binding protein)
.
- 1 +-
( 1).
Ca
pH.
+-:
(

2)

+-

( 3).

42
(Rouph ER), ( 4)
( 5,6)

.

E 1.10. . ():
( ) ():

T

: ,
, , , .
:

43
I

2. (Gibberellic Acids: GAs)

2.1. GAs
1926 Kurosawa

Gas

Gibberella fujikuroi,

( foolish seedlings Bakanae

disease). To 1956
.

ent-gibberellane. : 19 20
. GA3, ,
,

GA1.

.
GA1.

E 2.1. ent-gibberellane,
GA1, GA3 GA12

44

2.2. GAs
2.2.1. GAs

.
( 1). GAs ,
.

E 2.2.

(, ,

:
1.

(GGPP) ent- ent (CPP).

45
2.

ent- GA12 GA53.

3.
GA12 GA53

46
2.2.2. GAs
,
.

( 1).
2.2.3. - Gs

Gs ( 2.3),
.

2.3.

GAs (/)

GAs,
.

.

Gas ( 2.4).

E 2.4. GAs

47

2.4.

Gas

:

.

E 2.5. Gs
. 4 GAs
( )
:

. GAs 1618
.
( 2.6).

48

E 2.6. G3 GAs -1618

.

. GAs
-1618. -1618 G3.

(/ ):


( 2.7).

E 2.7. G3

.
,
G3,
G3

49
(
)
:
.
( 2.8).

E 2.8.

.

( -). GAs .

-
.
(, ).
( 2.9).

50

E 2.9. GAs
:

(.. ).
.
.

51

E 2.10.
.
:
.

2.5.

GAs

:
()

52

3.
3.1.

DNA 1955 Miller
. 1963 Letham
. 3.1
t RNA .

E 3.1. tRNA

3.2.

3.2

E 3.2.

53

3.3.

3.3.1.

isopentenyl pyrophosphate ( 2.2 ).
.

,
.

E 3.3.

54
,
, ( 1).
3.3.2.

(E 1).
,
. 3.4

.



.

E 3.4.




,

3.3.3.


.
( 3.5).

55

E 3.5.

3.4.

A -

(,
), () .

.

3.5. (/)
Gs,
.
, .

3.6.

:
: (
) . 3.6.

56
Agrobacterium tumefacients.
.

E 3.6.

Agrobacterium
tumefacients.

in vitro :
in vitro .

. 3.7.

.
( ).

( ).

57

E 3.7.
.
( ).

( ).
:


.
:
.

/ . ,

.
:

. 3.8.
.

.

58
.
( ) .

E 3.8.

.




.

.
(
)

:
. 3.9
.

( )
.

59
E 3.9.

( )

3.7.

( 3.10):
1.

( Arabidopsis

).
2.
.
3.
.
4.
.
5.
.
6.

60

E 3.10

3.8.

in vitro .
: ,
.

.

61

4.

4.1.

. 1901

. 1910

Penicillium. 1959 (Burg and
Thimann).

4.2.

4.1 ,
.

E 4.1.

4.3.

4.3.1.
To

stress.
.

Yang

( the Yang cycle ethylene biosynthesis).

5- adenosyl-methionine.

62
ACC
ACC (1-minoCyclopropane-1-Carboxylic acid). To ACC
ACC . 4.2
,
ACC
ACC .

E 4.2. ,
ACC ACC

4.3.2.

. 1-aminocyclopropane-1carboxylic acid (ACC), ,

63

.
4.3.1.

CO2 ( 4.3.).

E 4.3.

4.4.

(/)

.
.

CC- ACC- ( 4.2).
stresses , , ,
.

4.5.
:



( )
4.4.
4 :

0.1, 1.0 10 l/l .

64

E 4.4. 4 .

0.1, 1.0 10 l/l
.
:
, ,
avogado, , . ( 4.5)

CO2 (

) .

.
E 4.5.

CO2
(
)

65

:

(abscission).

.
(abscission zone)
( 4.6).

E 4.6.

( ):

. 4.7
() ().
E 4.7.


()

()

stresses

: 4.2 stresses

66
ACC ACC
.

: 4.8
2
()

10 ppm .
.
E 4.8.

2

()

10
ppm
.

:

.
.

4.6.

67


.
.


.

4.7.

.
,
.
.
(. )
,

4.9.

ACC
() ().

E 4.9.
ACC
() ().

68

5. M ()
5.1.
1961 Addicott
(abscissin). 1965 Wareing

(dormin). 1965
(abscisin=dormin).

.


.

5.2.
5.1
5.2.

E 5.1. H

69

E 5.2. H

5.3.
5.3.1.
,
stress ( 1). ,

Gas BRs.
isopentenyl pyrophosphate ( 5.3).
.

70

E 5.3.

(, ,

5.3.2.

( 1).

.

5.3.3. -

( 5.4).

71

E 5.4. -
5.4.

(/)

.
stresses , , ,
.
.

5.5.
stresses
stress. stress

72

.
: stresses

.
.
/ :

stress .

5 stress .

. 5.5.
Commelna communis: 50 mM Cl.
10 ..

E 5.5. H Commelna communis:

50 mM Cl.
10

stress:

.

.

.
: 5.6

.

73
, 6
.

E 5.6. H

.
,
6

-
: 2.9

.

:

:
etula sp
:

5.6.

5.7
:
1. .
2.
Ca2+ .

74
3.
Ca2+ .
4.
Ca2+ .
5.
+.
6. Ca2+ Cl-

7.
Ca2+
pH.

8.
+.
9. + Cl-

E 5.7. H

.
+ Cl-
(R:,
ROS
=

75
cADPR:, G-protein:
, PLC: C

5.7.

.
.

76

6.
6.1.

(Rs)

Rs

To 1970 o Mitchell
Brassica napus Alnus glutinosa
: ,
.
( ),
( ). 1979
Brassica

napus

(Brassinolide). 6.1

E 6.1. , (C27)

77
1982
(Castanea crenata).

6.2.

BRs .
40 ( C26, C28 C29).
.
.
, ,
. .
(
1000 ng/kg ).

E 6.2.

6.3. BRs

78

6.3.1. BRs
BRs . 6.3.
BRs,
.

E 6.3.
6.2.1.

M BRs

79

BRs
.

6.4

E 6.4. O
.

6.3.

(/)

Rs
( GAs) (
) ACC (ACC (1-aminocyclopropane-1-carboxylic acid).
GAs
(uniconazole, paclobutrazol, brassinazole)
Rs.

6.4.

BRs

80

+




Arabidopsis



stresses
6.5. Arabidopsis 16
(wild type)
.
.

E 6.5. Arabidopsis 16 (wild type)

.

6.6. ACC ( )
Arabidopsis.

81
, CC ( ),
CC+ .
CC .
E 6.6.

6.5.

BRs

6.7

ACC
(
)

Arabidopsis.

,
CC
(

),

CC+
.

CC

.
.
.
.

82

E 6.7. .
.

.

BRs
Rs ,

stresses. Rs 10-3-10-6 ppm.

7.
7.1.

(JASMONIC ACID JA)

H JA

To (jasmonic acid JA) 71

tryodiploidi theobromae. 80 JA

. To
cis- (
Jasminum grandiflorum).

7.2.

JA

7.1
.

83

E 7.1. .

7.3.

JA

JA .

. 7.3 JA
.

E 7.2.

84
7.4.

JA

JA
( 7.3).

E 7.3. JA

7.5.

JA

JA ,
.
.
JA :

jasmonate-induced-proteins

ACC- (
)
,

85

8.
8.1.

50. .

(), (), (),
).

mol.

8.2.

8.1

( ).

E 8.1.
8.3.

86


.

8.4.

9.

(Salicylic Acid)

To
19

,
.
9.1 SA (

SA).

SA

E 9.1.
9.1.

SA

87



Araceae

10.
:

,
. .
:

tany online. http://www.biology uni-hamburg.de/b-online/e31/31.htm

88
uchanan, B., Gruissem, W., Jones, R. Eds. (2000). Biochemistry and Molecular
Biology of Plants. American Society of Plant Physiologists. ISBN 0-94308839-9
Davies, P. J. (1995). Plant Hormones: Physiology, Biochemistry and Molecular
Biology. P. J. Davies ed. Kluwer, Dordrecht, Netherlands
. (1998). . .
Moore, C.T. (1989). Biochemistry and Physiology of Plant Hormones. 2nd ed.
Springer Verlag. ISBN 0-387-96984-5
Shrivastava, L.M. (2002). Plant Growth and Development. Hormones and
environment. Academic Press. ISBN 0-12-660570-X
Taiz, L., Zeiger, E. (1998). Plant Physiology. 2nd ed. Sinauer Associates Inc.
Publishers. Sunderland. ISBN 0-87893-831-1
Taiz, L., Zeiger, E. (2002). Plant Physiology. 3nd ed. Sinauer Associates Inc.
Publishers. Sunderland. ISBN 0-87893-823-0

89