Сколько кислорода переносит 1 г гемоглобина

22

Ζ

.

22.1.

,
.
.
,
. ,
,
, .
.

.
,
.
64 500, 16 000 [2, 7, 18].

.
. ,
.
,
, (.
22.1). ,
.
,
, .
.  O2
, .

, ()
(O2). ,
,
, ; .
, ,
.

,
.
(. 22.2). (),
.
,
.
,
.

. 22.1.

. 22.2. , ( )
( ).
( . 22.3)

.
, 10000 ,
. ,
140 .

.
[26, 27]. .
22.3 ,
. ,
.
. ,
, .
()

. 22.3. (
[26, 27])

, 141
, a,
( 146 )b.
(HbF) b g,
. HbF [2].

.
, ,
. ,
,
.

.

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

,
,
( ),
( )
577 541 [II].

, ,
.
.

555 .

.
.

, (
) .
.
(I0) (I)
. I/I0 ,
(I0 I)/I0 .

.

. 22.4. (bO2) (b).
,

. . 22.4
. ,
,
.
, , .
, ,
.

,
, ,
b bO2
.
,
. ,
,
(. ).
, , ,
,
, . . 600, 577, 470 .
. (. . 22.4).

.
()
( . 22.4 ):

E=lgI/I, (1)

I ,
I . ,

:

E=lgIo/I= e× C ×d,
(2)

d , e (,
). ,
.
.

;

.
158 / (15,8 /) 140 / (14
/) . ,
.

(. 22.5).

. 200 /,
(. 22.5). 115 /,
, .

, , .
,
.

. ,
, 130 / 120
/ .

.
, :

1)  O2 (1
1,36  O2);

2) (
0,34%);

3) (
);

4) ().

,

. 22.5.
(♂), (♀) .
,
; μ (),
(, ;
,
)

, .

.
.
,
, .

.
, (K3[Fe(CN)6]),
(KCN) (N3).
, HbCN (
), .
546
. e d,
, [ (2)],
. , ,
.

[32].

.

,

, . ()
( mean corpuscular hemoglobin,
MCH). ,
.
.

. 1
158 b 5,1 106
(1 = 106 ). :

= 158
/5 ,1·1012=
31 1012 = 31

:

= 140
/4,6·1012=
31 1012 = 31

( ;
= 29 ( ).

(2636 ) . ,
, .
. ,
,
,
. (, )
,
.
.
( ),

( ).

.

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

( mean corpuscular hemoglobin concentration, MCHC) ().
.
:

, = 5· 1061,
[] = 150 /, = 0,45, : = 30 ,
=/, =0,09106 = 90 () = 90 3

22.2.

() .
.
.

( ,
),
, .

.
, , .
( ),
1 , 1 (1 = 760 . . =
101 ).
:

[] = a/760 Pr (3)

760,

, r
.

, . . 22.1

. [ (3)],
,
. , ( O2 = 95 . . CO2 = 40 . .)
O2 0,003  O2 1 , O20,026 O2, 1
.  O2 ,  O2,
O2, 9 . ,
O2 20 , O2.

22.1.
a ( ·
1· 1)
 O2, O2 N2

 O2  O2 
,
.
,
. ,  O2 O2 
.

.
. ,  O2
, ,
(. 22.3). ,
:

+4O2
↔(O2)4. (4)

, 1 4
 O2. 1
22,4 , 64 500 4 22,4  O2,
1 1,39  O2.
(1,341,36  O2
1 ). ,
[25]. , , invivo 1 Hb 1,34 2 (
).

. 1 Hb (=
16 100 Hb) 1  O2 (= 22,4 O2).
, .

, ,
, : [O2]max =
(1,34  O2 1 )·(150 Hb 1 ) =
0,2  O2 1 .
,
, (PO2 > 300 . .);
(4) .
 O2,
.

.
(4) . ,

 O2 ;
, ,  O2.
(SO2 ) :

SO2=[O2]/
[Hb] + [O2]·
100% (5)

, SO2 =0%;
, SO2 = 100%.  O2.

. . 22.6, S .
(.
). , ,
2, ..
 O2,
50%. ( 7,4 t=37C) 50
26 . . (3,46 ) [9, 29].

.

S
.
 O2,

. 22.6. (Hb)
(Mb) 7,4 t 37

[11]. ,
,
(Mb), [1].

, 1:4.
,
O2:

Mb + O2 ↔ bO2. (6)

. 22.6. ,
S bO2
 O2,
. ,
 O2 ,
. ,

, .

,

,
. ,
,
S bO2 [11,14].

.
.
 O2
(PO2)
. O2
95 . ,. (12,6 ). . 22.6 ,
97%. (
)  O2
, ,
,
. , O2
60 . . (8,0 )
90%. ,

,
.

.
PO2 .
PO2
40 . . (5,3 ), 73% .

5 . . (0,7 ),
7%;  O2
.

 O2
.

(SO2 ). SO2 , ,
,  O2 (  O2
1 ):

[O2]= 1,34·[Hb]·SO2·105 (7)

SO2 , a [Hb]
.
, , (SO2 = 97%)
0,20, (SO2 = 73%) 0,15.
, (O2
) 0,05 (. 22.2). ,
25% .
,
(. . 23.2), ,
. 22.2, ,
.
0,1.

,

b2
,
[2, 12, 14], ,

S . , ,
 O2  ,
.

.
( )
.
, (. 22.7, ).

22.2.

. 22.7.
[9, 29]. . . . ( ). .
O2. . 2,3 (2,3)
. , ( )
( ),
,
.

PO2.
+ (
. 22.7, , +
). , . . ,
. . 22.7,
, +

. ,
.
(. . 22.7, ) .
O2 (O2 ): O2
, . . 22.7,
O2· ,
O2


bO2.
,
, O2
. ,
[14].

.
,
(
). , .
O2 O2,
.
. 22.7, ,
bO2.
( ; pO2 = 40 . ., CO2 = 46 . .),
, ( ;PO2 = 95 . ., CO2 = 40 . .),
. ,
,
. ,
. O2
.
O2,
.
( . 22.7, ).

,
2
. ,
.

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

,
.
2,32,3 (. 22.7, )
[16. 22].
. :
()
, , ()
. ,
.

. ,
, .

.
, , in vivo
( , ).

.
, , [7, 21].

. 22.8

. 22.8.
[O2] (PO2)
.
O2 (
) ( ), ‘
( ), ‘ ( )

.
.
(120 180 / ),
, O2 .

() ‘ ‘ (). ,
O2 , .
, PO2= 25 . . (3,3 ) O2
0,08, 0,11.

.
O2 ,
( . 22.8 ). .

( , )
, .
:

+ ↔
b. (8)

,
.
,
b , O2 [3].
O2
30 ppm ( ),
FCO2 3·105 (0,003 .%).
,
, b 5%
. O2,
b bO2
5%, 1 : 350. ,
b 350
, O2.

. ,
.
; ,
.
, O2.
b 1 %
; 20%. ,
, ,
3·104.
.

, .
b, (
, ) [3]
.
O2 .

,
,
,
.
, O2 .
,
, .

22.3. 2

O2

(O2, )

. ,
, .
O2 .
, , O2,

.

O2 [15]. O2 ,
, 40 . . (5,3 ).
, , O2 ,
.
O2
.
, O2
(. 22.9). O2
, :

O2 + 2
↔ 23+ + H+
(9)

;
10 . .
[6, 24].
, O2,
, .

. 22.9.
, ()
()

3
,
. 3
,
. 3
, .
,
, 3 1.
(
).

O2 3«, +.
, , ,
. , ,
. ,
, ,
+.

O2
.
():

HbNH2 + O2 ↔ HbNHCOOH + +)

, O2, ( ).

. 22.9. ,

O2 .
, O2 ;
, .

O2 . ,
, ; 40 .. ,
,
46 .. 1 1,8
O2. 12%
(bO2),
11% , 27%
, 50%
3 . ;
.

O2 (
O2, .
O2
( , ),
O2 .
O2;
, (saturatio),
. . (.:
, 1988).
,
, O2 )

O2 .

O2
O2 , . O2
. CO2 O2 .
O2 ,
. . 22.11
.
,
, ,
+ . ,
,
, O2. ,
, , O2
[6, 20]. O2

.

O2
. bO2
, O2 .
O2 O2 ,
.
O2 ,
( O2/ /).

O2 (. 22.10)
.
(. 22.16).

. O2
,
O2.
O2 .

( . 22.10).
,
O2 .
, O2 .

.

.
22.10. O2
. ( O2),
( ) ( ),

, ,
, O2 .

(. 22.10). , O2 ,
O2. , ,
O2 ,
.

22.4. ,

. ,
, (
), , (
).
.

↔
H+ + .
(11)

( ) .
,
(

).
.
, .
, l, (11)
. ,
.
(. 22.11).

.
.
.
+:

p=lg[+]. (12)

, 7 ( ),
H+ [+] 107
/. .

, , .
,
, . ,
,
,
H+, . .

.
22.11. .
‘ ‘.
+

. ,
.
,
.
( ͖)
.
, +.
.
,
.
.
͖ ,
.
.

.

.
( 37 ) 7,37 7,43,
7.40. ,
( , ,
). . ,
7,27,3, . .
. , .

,
.
,
, , .

,
. (. .
) .
, .

.
,
+
.

[H+] ·[]/[HA]= K (13)

,
, , ; ‘ ,
( ‘ ,
, ). H+ ,

. ,
,
H+, .
, ,
H+.
[H+].
+ +
.

(13) :

lg[H+]= IgK‘lg[]/ []’
(14)

= ‘ + lg[]/[] (15)

,
, .
‘, ‘,,
(‘ = IgK’). (15)
:

= ‘ +
lg a/1a (16)

a = []/
[]+[]

a,
, []
([] + []). ,
. 22.11, a .
,
, ‘ + 2.
.

,
, , H+
, , .

= ‘ (. 20.11).
,
‘ .

.
.
, O2,
:

O2
+ 2 ↔ 2↔ H+ + 3 (17)

:

= ‘ +
lg [3]/[2]
(18)

[O2] O2 (PCO2)

= ‘ +
lg [3]/0,03·P2](19)

0,03 ·
1 ..1,
, [3] /, a PCO2 ..

‘ 6,1.
, ‘ (7,4)
.
,
.
, O2 (40
. .), 3
(24 /). , O2 ,
.
,
, O2 ( , )
.

.
, ,
24 ,
42. ‘
(6,8) ,
.

.
.
,
‘ .

, , .

(
, ), .
,
,
.

, ,
O2.
[3],
O2,
[
(19)]. O2;
O3- ,
,
O2 (.
22.12). ,
,
,
O2.

.
22.12. O2
. ,
. O2.
,

. 22.13.

( 2,3) ( [19] ).
Pi- ;
. HbO2 Hb .
b
bO2, 1
b
0,45 H+

. . 22.13,
( )
. ,

, .
, , +
. ,
,
+ O2.
. , O2
.

.
;
().
, ,
( buffer bases, BB) [10].

. 22.14
, ( ).
. ,
, ( ),
( 1),
, S042
( ). ,
3
, . 1/3
.

48 /. ,
O2. .
, , O2 . H+
3 . +
, .
,
(. 22.15).

.
22.14. , .
(
); , .
, , CI; Kt+

.
22.15. CO2.
( 48
/)

,
, . , CO2 40 50
.., 0,06, 0,1.
, H+ 105 /.
, . 22.15.

PCO2,
,
.
(48 /) (
baseexcess, BE).
, BE .
BE
, .
, .

.
O2 ,
. 230
O2/, 15 .
.
H+.
,
.
. ,
,
H+ (),
O2, 3 + +→ 2
→ 2 + O2, ,
. ,
; O2 H+
,
.

.
. ,
. 4060
+, .
,
H+ .
. ,
H+ ,
.

H+
,
. H+ 4 NH3,
.
3
.

.
,
( ,
) .
, ,
.
( < 7,37) , (
> 7,43) .
.
(
2 ,
).
()

.
(, ) ;
, l ( )
. .
, ,
, .

. ,
,
2(PCO2)
(BE).
PCO2
(BE = 0). ,
PCO2 ,
BE .
( ),
( );

.
22.16. O2
(BE).
()

O2 .
O2
BE =

( )
( BE).

O2 , [3].
O2
(BE) (. 22.16).
( )
O2,
. ,

O2.

. 22.17.
, .
O2. , BE CO2
. ,
, .
(
).
, , BE .
,

.
22.17.
. BE, PCO2.
;
;

( ) BE = 0 /, CO2 = 60 .., ; BE = 15
/, CO2 =40 ..,
(, ) .

.

, . .
,
( ).
:

1.
. ,
,
.

2.
, H+
3 .

. 22.17.
( 1).

, . ,
O2 .
( 1).
CO2 ,
.
, CO2 , ,
()
.
( 2)
PCO2, ( 2).

, . ( ), , ,
, O2 .
.
,
( 36). , (
4), O2,

( 4).

.
.
,
,
.
[4, 5, 8, 10].
.

1. . ,
+ ( 7,377,43)
.
,
.

2. O2. O2
(3545 ..)
.

3. (BE). BE

. ( 2,5 +2,5 /)

.

4. .

.
,
(O2 = = 40 . .)
37 . 24 /.
, .

. 22.3
,
.

.

,
O2 [10].

22.3.
. ;
(↑ ;
↓)

,
O2. .
O2
(. 22.18). ,
( ),
.

.
22.18. O2
[10].
, ,
O2 . ,
, , .
O2.
BE
. 1
( ): O2 44 . ., 7,37, BE = 0 /;
. 2 ( ): O2= 32 . ., 7,28, BE = 11 /;

, ().
O2 .
. , . 22.18
,
(BE =11 /),
O2 (CO2= 32 ..;
).

O2
. , CO2 , BE
( ),
[28]. .
22.19 BE CO2.
. CO2 ,

.
22.19. BE
C2
. ,
.
; BE .
. ( ): C2=32
. ., 7,28; BE = 11
/. [29]

.
BE . , , .
22.19, CO2 =32 .. 7,28; BE =11
/. ,
( 2 . 22.18).

22.5.

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E., Brunori M. Hemoglobin and myoglobin in their reactions with ligands.
Amsterdam. North Holland, 1971.

2. Baumann
R., Barlels H., Bauer C. Blood oxygen transport. In: Fahri L. E., Tenney
S. M. (eds.). Handbook of Physiology, Sect. 3: The Respiratory System, Vol.
IV. Bethesda. Amer. Physiol. Soc., 1987.

3. Coburn
R.F., Forman H.J. Carbon monoxide toxicity. In: Fahri L. E., Tenney S.
M. (eds.). Handbook of Physiolofy, Sect. 3: The Respiratory System, Vol.
IV. Bethesda. Amer Physiol. Soc., 1987.

4. Hills
A. G. Acidbase balance: chemistry, physiology, pathophysiology. Baltimore.
Wiiliams and Wilkens, 1973.

5. Kildberg
P. Clinical acidbase physiology. Baltimore. Williams and Wilkens, 1968.

6. Klocke
R.A. Carbon dioxide transport. In: Fahri L.E., Tenney S. M. (eds.).
Handbook of Physiology, Sect. 3. The Respiratory System, Vol. IV. Bethesda.
Amer. Physiol. Soc., 1987.

7. Lungo
L. 0. Respiratory gas exchange in the placenta. In: Fahri L. E., Tenney
S. M. (eds.). Handbook of Physiology, Sect. 3. The Respiratory System, Vol.
IV. Betehesda, Amer Physiol. Soc., 1987,

8. Masoro
E. J., Siegel P. D. Acidbase regulation. Its physiology and pathophysiology.
PhiladelphiaLondonToronto. Saunders, 1971.

9. Severinghaus
J. W. Blood gas concentrations. In: Handbook of Physiology, Respiration II.
Washington, Amer. Physiol. Soc., 1965.

10. SiggaardAndersen
0. The acidbase status of the blood. Copenhagen. Munksgaard, 1974.

11. Weissbluth
M. Hemoglobin: Cooperativity and electronic properties. BerlinHeidelberg
New York. Springer, 1974.

12. Wood
S. C., Lenfant C. Phylogeny of the gasexchange system: red cell function.
In: Fahri L. E., Tenney S. M. (eds.). Handbook of Physiology, Sect. 3.
The Respiratory System, Vol. IV, Bethesda. Amer Physiol. Soc., 1987.

13. Adair
G.S. The hemoglobin system. VI. The oxygen dissociation curve of
hemoglobin. J. Biol. Chem., 63, 529 (1925).

14. Bauer
C. On the respiratory function of haemoglobin. Rev. Physiol. Biochem.
Pharmacol., 70, 1 (1974).

15. Bauer
C., Gros G., Bartels H. (eds.). Biophysics and physiology of carbon
dioxide. BerlinHeidelbergNew York. Springer, 1980.

16. Benesch
R. E., Benesch R. Yu, C.I. The oxygenation of hemoglobin in the presence of
2,3diphosphoglycerate. Effect of temperature, pH, ionic strenght and
hemoglobin concentration. Biochemistry, 8, 2567 (1969).

17. Braunilzer
G. The molecular weight of human haemoglobin. Bibl. heamat. (Basel), 18, 59
(1964).

18. Braunilzer
G., Hilse ., RudloffV., Hilschman N. The hemoglobins. Adv.
Protein. Chem., 19, 1 (1964).

19. Brodda
. Zur Theorie des SaureBasenHaushaltes von
menschlichem Blut. Akadem. Wiss. Lit. Mainz; Wesbaden. Steiner, 1975.

20. Christiansen
J., Douglas C.G., Haldane J.S. The absorption and dissociation of carbon
dioxide by human blood. J. PhysioL, XLVIII, 244 (1914).

21. Fischer
W.M., Vogel H.R., Thews G. O2 and CO2, exchange in
the human placenta. In: Lubbers D.W., LuftU.C., Thews G.. Witzler E.
Oxygen transport in blood and tissue. Stuttgart. Thieme, 1968.

22. Kilmartin
J. V., RossiBernardi L. Interactions of hemoglobin with hydrogen ions,
carbon dioxide, and organic phosphates. Physiol. Rev., 53, 836 (1973).

23. King
E. J., Gilchrist M. Determination of haemoglobin by a cyanhaematin method.
Lancet. II, 201 (1947).

24. Maren
.. Carbonic anhydrase: Chemistry, physiology, and
inhibition. Physiol. Rev., 47, 595 (1967).

25. MerletBenichou
E., Sinet M., Blayo M. C., Gaudebout C. Oxydencombining capacity in dog.
In vitro and in vivo determination. Respir. Physiol., 21, 87 (1974).

26. Perutz
M. F. The hemoglobin molecule. Proc. Roy. Soc., B, 173, 113 (1969).

27. Perutz
M.F. Stereochemistry of cooperative effects in haemoglobin. Nature, 228,
726 (1970).

28. Thews
G. Bin Nomogramm fur die O2Abhangigkeit des SaureBasenStatus
im menschlichen Blut. Pflugers Arch. ges. Physiol., 296, 212 (1967).

29. Thews
G. (ed.). Nomogramme zum SaureBasenStatus des Blutes und zum
Atemgastransport. BerlinHeidelbergNew York. Springer, 1971.