1

1999

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

1.
2. -
2.1.
2.2.
2.3.
2.3.1.
2.3.2.
2.3.3.
2.3.4.
2.3.5.
2.3.6.
2.4.
3.
3.1. ;
3.2.
3.3.
3.4.
3.5.
3.6.
3.6.1.
3.6.2.
3.6.3. ,
3.7.
3.7.1.
3.7.2.
3.7.3.
3.7.4.
3.7.5.
4.

5.
5.1.

5.2.
5.3.
5.4.
5.5.
6.

.

.
.

. , . . -.

, ,
.

1.

, .

.
( ) (
) .
E
.
1975, ..

( ). 1975
.
80
,
.
, " " ,
.

, 4
:

(. 1) -

,
.

(.

1)

" "
,
, ,
, ,
.

H (. 1)

. ' ,

, , ,
.

, ,
,
.

. 1.
(
)
(
) (: Amoros & Petts 1993).

, ,
( ,
, , )

, , )
.
, ,
.
,
,
. ,

" ".

2. -
2.1.

(.. ). ,
,

.
(drainage network)
.
()
(drainage basin) (. 2).
,
.

1:x.xxx.xxx

. 2.

,
( ) .

,

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

.
( ) (
)
. ,
,
.

,
, .
,
. ,
.

(. 3).


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

.

10

.


.
, ( )
(, )
.

. 3. (: Wetzel 1991 1993, ).

1 .
,
, .

,
.
.
Horton (1945) Strahler ((1957)
():

()
1

11

. , ,
.
... (. 4). Horton

Horton.
(Horton 1945)

. 4.
Horton-Strahler
(: Wetzel 1991).

. (
) (
).
:
1- :
.
,
-
- ,
. ,
,
.
.

.
.
2- (infiltration capacity):

.

12

( )
(, )

.
: - , -
( ), -
(), - (,
, ), - , -

- .
3- :
.
4- ( )
.
,
.

.

2.2.

.
().

.
,
.
,
.
,
.

13

. 5. . . .
. (: 1969).

(Davis, 1899).
V (
). U
( ). ,
( ).
(. 5).
,
.

() , .
, . ,
,

14

( ).
.

2
.
Davis


.

.
.
,
. ,
.
(.. )
( , , ).
3.

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

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

.
3
.

15

.
.
.

.

. ,
.

.

, .
(
) .

.

.

,
.
.
.

2.3.

2.3.1.

.
(.. ).
-
- .

16

. ,
.
(. 6).

.

1/3 -. ,
(current velocity), (water flow,
discharge),


.

).

. 6. .

.
(: Amoros &
Petts 1993).

,
. ,

( ).


.
.
(.. )

. ,
.

17

2.3.2.

,

.

.
.

,
,
.


.

.

,
.
.
.
. ,
(...)
500 000 000 , .

. ,
(, )
,
. ,

, .

18

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

,
.

.
. ,

.

( ),
.
() .


. :
1-

2-
3- ,
.
.

, ,
.

,
,
()
(alluvial fans) .

19

.
.

, .
, .

()
.

, ,
(floodplain)
.
: - (channel deposits) -
(overbank deposits)
. ,
, ,
.

( )

(natural levees).


.
,
. (backswamp
deposits), (braided stream
deposits)
.

2.3.3.

20

. , ,
.
.
.
, .

.
.

. 7. (:
Nachtnebel . 1991).

() .

2.3.4.

: ,
(. 7).

21

.
,
,
. .
,
' .

(. 8).

22

1.

2.

3.

. 8.
(: Ward . 1994).

: ,
(. 9).

23

:

.



.
,

.
:

().

( )

.

. 9.
(: Ward .
1994).

24

.

.

2.3.5.

. 10.
(: Ward . 1994).

(pools) (riffles) (. 10).

( ).

. 11.

(:
1986).

25

,
().
,
( ) (. 11).

.
,
,
(. 12, . 13). ,
() (oxbow

. 12. (: 1969).

lakes) (. 14).
, , .


.




.

. 13.

1969).

(:

26

( )

. 14. , ,
( ) (: 1986).

, .
, (Fisk
1944) : - (point bar deposits) -
(abandoned channel fillings) -
(natural levees deposits).


.
, .

2.3.6.

.
,
. ,
. , ,

27

,
.

.
.
,
.

:
.

(, ),

.

2.4.

, . ,

. 4
1% 30%
.
()

. ,
.
, 5
(.. ).

.
4
5

.
.

28

.
.

.
,

( ).

()

. , ,
(..
).

. ,
.
, .

29

3.
3.1. ;

(boundary) ,


.
( , ...).

, (lotic
ecosystems). (holistic concept)
() .
, .
,
.
, .
.
,

. ,

.
(.. ).

.
,
,
.
(. 15).

. ,

30

.
(
),
( )
( ).


.
.
,
. ,
.
.
(.. )
.
,
(self-regulating system)
.

. ,

.

3.2.

.

.
,
. ,
,
, , ,

31

, .
,
.
.
,
.
.
, ,

(, ).

,
, ,
, .
,
.
,

.
, , ,
, , (. 16) .
: , , 6 7.
(, , , , )
, ,
.

, .
.
,

. ,
, , .
7
,
, .

32

, . ,
, ,
(dead zones) .
1,7 /
.
, Gammarus Asellus aquaticus,
,
. ,

. ' ,
(.. )
.

/ .

(. 17):
-
- ( )
- (Simuliidae, )
-

- ,
(.. , Gammaridae)
.
,
,
.
, (drift).

.

.

33

, .

.

(behavioral drift).

. ,

.
,
. (, . .
) . '
. .
.
(, . . )
, ,
. ,
2 ,

. 2 ,
.

.
,
, , ,
, (. 18, 1).

1. .

:
, (pools)
, (riffles)
(gravel bars)

34

(banksides)
(side arm)
(backwater)
( )

(
)


/

(fen/swamp)
(marsh)

(Leusiscus cephalus)
(Salmo trutta).
.
Riparia riparia Alcedo atthis.
,
,
.
, , ,
. ,
.
.

.

19

.
.
,
, . T

35

' . .

.
.
(.. )
.

.
.
:
,
, , , ,
, ...

3.3.

:
- , 1. 2 2.

.
-. , 1 3. 2 50 2.
- , 3 10. 50 300 2.
- , >10. , 50
500 2 300 500 2.

,
pH.

(trophogenic) (tropholytic)
( ) .
, ,

36

( )
(),
.

( ).
. ,
( , ..)

.

.
,
- .
.

20
. (.
20):
- o (Salmo trutta, S. gairdneri),
- -
,
. 100C.
,
( ) .
- (Leuciscus cephalus) (Phoxinus
phoxinus).
.
.

150C.
.
- (Barbus barbus) (Rutilus
rutilus)

37

.

. 150C
.
: (Scardinius erythrophthalmus),
(Perca fluviatilis) (Esox lucius).
- (Abramis brama)
( ).
.
200C
. , .
: , , , , (Anguilla
anguilla), (Tinca tinca ) (Cyprinus carpio). ,
, ,
.
,
. .
, . ,
. ,
.
(, Krenon)
(, rhithron)
(, potamon).

. .
.
,
.
,
.
.
, , . -
- .

38

, ,
.
(, , )

200 C,
.
. ,
.

. ,
: Hildenbrandia Lemanea
Vaucheria Cladophora
.
Myriophyllum ( ) Ranunculus (. 21,
). .
.


. .

.
(, , )
200 C,

. .
Elodea ( )
Sparganium ( ).
, 2
. ,
. : ,
, , , .
(),
.

39

,
.
,
,
.

.
.
3.4.

, , .

, , ,
.

.

.
(River Continuum Concept, Vannote ., 1980)
. ,
,


.

:
(- ), (-
) (> ) (. 22).
() ()
.
,
,
. (:) < 1

40

(. 22). ,
.
. ,
: >1.
.

() . ,

. , :
<1.


.
,
(), ,

.


-
-
.

.
,

.

.
4 :
- , () (shredders)
(>1mm) .

41

- ()
(collectors). (50 m - 1mm) (0.5 50 m) .
.
- ()
() (scrapers, grazers).
,
( )
() .
,
. ,

.
- (predators),
, .
,
,
.

. .

.
,

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

, .
, ,
.

42

3.5.

.


,
. ,
,

.
,

. ,
.
,

. ,
,

. ,

. ,
, .
,
(boundaries)
(interfaces)
, , (hyporheic) 8.

,
(. 23). ,

43

/ .
,

. 24
.
,
.

:
,
.
, ,

,


.
3.6.

,
, .

.
, .

, ,
. ,
.
8

44

,
,
, ,

, .
20


,
.

.


. ,
.
.

3.6.1.

( , ,
) ,
,
.

.
(

)
.

45

,
,
.
.
. ,

, .
, (Lemanea,
Enteromorpha) (Cladophora),
(Oedogonium, Ulothrix,
Stigeoclonium), ,
(Chara, Nitella).
,
,
.
, Cladophora ,
4 .
.
,



.

, .
,
() ,

.
, ,
2 m s-1

46

, . ,
,
( Cocconeis Chamaesiphon),
( Achnanthes, Cymbella,
Gomphonema, . 25). (Hildenbrandia,
Lithoderma), o ,
(Phormidium Rivularia)
.
, ,
(.. Gongrosira) (.. Schizothrix).

,
. ,
(Cocconeis, Navicula Gomphonema spp.
, Achnanthes, Meridion ..
),
(.. Closterium)
(.. Aulacoseira, Oncobyrsa)
( Cryptomonas)
Daphnia.
,

.

,
,
,
, ,
.
,

47

,
. .
.

(Oscillatoria, Phormidium), (Microcoleus, Mastigocladus),
(Hydrodictyon), (Nitzschia, Caloneis, Surirella
Cymatopleura). ,
.
,
.

(suspended algae)
,

,
. ,
- -
.



, . ,

.
,

.

,


. Centrales
(Cyclotella Stephanodiscus) Pennales

48

(Nitzschia, Asterionella) Chlorella,

Ankistrodesmus, Scenedesmus Pediastrum (Trachelomonas,
Phacus). Dinobryon,
Cryptomonas, Microcystis Oscillatoria,

.

,


. (
, ,
)
.

.

3.6.2.

, ,

, ,

. ,
,
. ,
,
,
. ,

.
-

49

-
.

, ,

,
, 9 (bedrock)
, ,
,
.
,
( , )
.

1- -
pH.
,
,

,
.
,
.
,
, ,
, .
(acidification)
,
Eunotia Pinnularia.
(.. , ,)

50

, ,
(
Navicula, Caloneis, Gyrosigma).

. ,
.

2- - ,

. ,

.
,
(gradient). ,
,
,
.

.
( ,
) 6,7 x 1012 m3 ( ).
,
,

,
. ,
.

3- - ,
2 m s-1,
.

51

0 2 m s-1,
,
.

.

.
,
.

.

( < 0.1 mm).

.
,
. ,
-
-
.

4- -
,
.
,
.

.

5- - , ,
.
,

52

. (resuspension)
. ,
- -
.
,
,
.
,

(horizontal stress, shear) ,

.

o,
,
. ,
.
, ,
,
.

6- - ,

.
, .
, 0.1 kg m-3
.
,
, .
,
,
.

53

,
. ,
,
,
(photoinhibition) .

, .

,
, .
,
.
,

.

1- - ,
,
. ,

Ulothrix

Cladophora

. Hildenbrandia Batrachospermum

.

.

54

2- -
,
.
-
-
. ,
.
,
,
.
,

. ,
.
,

.

3- - , ,
. (grazing)


.
(filter feeders)
Simuliidae () ,
.

.
,
. ,
(
) .

55

, ,


.

.
,
. ,
,


,
.
,
, ,
.

,

(. 26). ,

( .).

3.6.3. ,

,
.

, ,
,

56

, . ,

.
.


- -
(residence time)
. ,

, , ,

,
.
(
)
.
27
.
<104
,
( )

.
105 ,
.
,
(R-
Grime 1979) ,
(C-),
.
,

57

( ) .
,
- -
Microcystis , . ,
, S
(S- ), Grime.
,
,
.
,

.

, .


,
.


. ,
,
.

Margalef (1960).
,
,
,
(Intermediate
Disturbance Hypothesis) Connell (1978). ,

58

Odum (1969). ,
.

.
(, ),
.
, Ceratoneis
Hydrurus foetidus. Achnanthes Meridion
,
Synedra Aulacoseira, ,
,
.
(Amblystegium eurhynchioides)
,
.

, Meridion,
Lemanea Ulothrix Nostoc.
,

.

,
.
. ,

, .
, Margalef (1960), Aulacoseira Navicula - Rhoicosphenia - Cymbella - Diploneis
. Phormidium Lyngbya
(), Amphipleura
Chaetophora, Chara Zygnema

59

. Fontinalis Fissidens .
,
, Eunotia, Fragilaria,
Melosira, Nitzschia Pinnularia. Oedogonium Tribonema
,
Callitriche Ranunculus. (
) , Cocconeis Melosira - Navicula - Surirella Cladophora,
Stigeoclonium , , Enteromorpha. ,
,
. ,
,
,
,
. ,
.

Stephanodiscus - Nitzschia - Asterionella ( )
Chlorella - Ankistrodesmus - Scenedesmus - (
),
Dinobryon - Sphaerocystis,
Oscillatoria agardhii - Oscillatoria redekei (
) Microcystis,
( ,
) .
/
,
( ),
() ()
.
,

. ,

60

.
,

,
.

.


.
3.7.

3.7.1.

10
.

.
,

.
(hydraulic
stream ecology) Statzner . (1988),


.

(site-specific
hydraulic conditions).
10

, ...

61

, ( ),
( )
( ),

.

,

.
,


.

.
,

.
, .


. ,


.

, ,

.

,
, , .

62

3.7.2.

. ,
, .
-
,

(reservoirs), .

. ,

.
, .
.
,
. , ,
,
.
, 11
.

.
,
.


.

. , ,

63

-.

. ,

,
.

.

.
,

. ,
,
,

.

3.7.3.

,
,
.
,
/ .
,
. ,

, ( ),
.
11

/ .

64

,
.
,
()

. ,
,
.
,
. ,
,
25
.
,

(. 28).

. ,


,
, , .

, ,
.
,

.
,
, (

65

) ,
(. 29 & . 30).

,

.
Chironomidae
,
. ,
Chironomidae
.
, .

- ,


.

.
.
,
.

.
, .
(catastrophic drift)
,

.
,
.

66

.

.
- - ,

.

.
.
.

.

,
.

.

,
.

3.7.4.

. , ,

.
,


.

67

.


.

,
.

,
.

, ,
.
,
,
.

,
.

.
75%.

.
. ,
,
.
,
,
. ,

, .

68

3.7.5.

,
, .

.

(. 31).

,
,


.

, / ,
,
- (poolriffle sequences) .



.

69

4.

4.1.

, , .

, 60
. ,
.
, ,
, 1/4 . ,
14, 29 48 ,
. 31
50.000 . ,
2/3
(
),
.
, .


.
, ,
,
,
20 . ,


IUCN ( ).

.

70

4.2. ()

(regulation)

. ,
.
.
, ,
.
20 4.000
, 15 ,
60.
' .
'
.
,
, ,
.
60, ,
.

. ,
.
,
,
.

4.3.

, ,
,

71

.
.
,
, , .
20 ,

.
, (..
, , )
.
80 90,
,
.
223 1977-82 1988-90
25% 91 ,
18.
40%
.

,
.
.

.

4.4. -

, (
)
, ,
,
. .
,

72

,
.
.

,
.

, .
,
,
.

.
221
80 90,
(61%). 1/3
25%. ,
1/2 25%. ,
,
,
.
,
.
,
,
. 230
1977-82 1988-90
2/3 .
,

.

73

.
.

4.5.

(acidification)

. 20

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

. pH 5.0
. 25
.
,
,
.
. , 21

. , .
.
1980 1990
27%. (48%)

74

(18%).
5% ,
.
.
, 70-80%
,
.
, , ,

. , ,
2-3
.

4.6.

19
20 ,
.
. ,
.

,

(monitoring) ,

(, -
).
,

. ,
.
,
.

75

,
.
70,

(
). 90

.
,

, . ,
-
-

. ,

,
.

4.7.

()
60 90,
(
, ) .
(.. DDT, PCBs) ,

.
,
.

.

76

. ,
,
.


.

4.8.

(,
),
.

, . ,

.
20 200
. ,
,
. ,
(.. ) .

.
,
.

4.9.

, -, ,
(salinisation) , (
)

77

.
, , .

4.10.

, ,
. ,


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

.
.


.
.
,
.

78

5.

. , .
,

.
. ,

.

5.1.

,
.

. ,
.
.
( ), ,

.
.
,

. ,
,
(..

79

).

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

.

5.2.

. ,
,
.
, 10
,
.
, ,

.
,
.
, ,

(, , , ),

.

80

.
,
.

5.3.

,
.
,
.
,
.

.

(. 32).
,
( ).
.
. ,
, .

. ,
.
, () ,
.
, : -
, -

. ,

81

.

.
(
).
,

.

5.4.

.

.

. , ,

.
. ,
.

. Cleve

.
20

.
1970-1990
. Coste (1974)
"" ,
" " (Biotic index),

82

. , Descy (1979) (Id

index)
" "
.
Decsy & Coste (1991).

(, , ..).

. ,
.

.

: , ,

,
,

- ,
.

.

, ,
.

.

83

.
.
...,
1940 R. Patrick Diatometer
. ,
70
.
( 3) ,
.

...

,

(ecoregions).
( 1000
) , .


(
, , ,
, .)
.
,
,
.

,
.

84

5.5.

.

( 3 & . 33). Coste & Leynaud (1974)
. 55 4
(),
7 ,
(), -
.
1-10, .

1984.
(EEC index, Descy & Coste
1991). , 233
( 3).

(relative
abundance), (pollution sensitivity)
(indicator value). O
Descy (1979). Zelinka & Marvan (1961)
. ,
106 .
1 5 :
ID = >4.5

ID = 4.5-4

ID = 4-3.5

ID = 3.5-3

ID = 3-2

ID = 2-1

85

Coste
IPS (CEMAGREF, 1982). IPS 263
1-20
. 1982
.
IDG (Coste & Ayphassoro, 1991) 174

. Sladeek
(1986) .
Leclercq & Maquet (1987) Sladeek,
210 . IBD (Lenoir & Coste, 1996)
, IPS,
. 209 .
1332 ( 1000
14 ) .
7 . Schiefele (Schiefele
& Schreiner 1991, ) 7 386 ,

7 . ,
Watanabe (1990)

(BOD5) .
1 ( ) 4 ( ).

86

6.

,
( )

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

,
1.200 (.. .. ), 300
(.. ).
.
,
.
, .
(), ,
. ,
, .
,

.
.
(.. , , , )
. ,

, .
,

87

.
4270 . 34
, RAMSAR .
, ,
. (..
) . ,
, , .
,
.
,
, .
,
(
, ),

,
. ,
,
, ,
.
, ,

: ,

.

(57%) ((16%), , ,
, (Skoulikidis, 1992).

(
CaCO3). ,
( 2).

88

2.
(: 1998, Skoulikidis 1993).

Ca++>Na+>Mg++>K+
Ca++>Mg++>Na+>K+
HCO3->SO4->ClHCO3->Cl->SO4-

, , , , ,
, , , ,

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

89

.. (. .), 1990 - .
, WWF, IUCN, ,
.
., 1986 - , .
.., 1986 - , University Studio Press, .
., 1998 - , , ,
1, ...., .
., 1995 - (, , ). : .
(.) ,
, WWF, , .
- ., 1993 - , .
., 1969 - , .

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