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8/11/2019 The Effect of High Salt Intake on NaKATPase Activity of Tissues in the Rat_Clin Exp Hyperten 1994
1/14
CLIN. AND EXPER. HYPERTENSION,
16 3),
327-340 1994)
THE EFFECT OF HI GH SALT I NTAKE
ON
NA, K+- ATPASE ACTI VI TY OF TI SSUES I N THE RAT
Paul Wai Chi ng Li , C S Ho and R Swamnat han*
Depart ment of Chemcal Pat hol ogy
Pr i nce
of
Wal es, Hospi t al
Shat i n, NT, Hong
Kong
and
UMDS, Guy s Hospi t al
St Thomas Str eet
LONDON
*Depart ment
of
Cl i ni cal Bi ochemst ry
Key Wor ds
:
Sodi um Na, K+- ATPase, sodi umt ranspor t
i nhi bi t or
ABSTRACT
1. The ef f ect of chroni c f eedi ng
of
hi gh sal t di et
on Na, K+- ATPase act i vi t y of hear t , l i ver ,
skel et al muscl e, ki dney and aor t a was st udi ed i n
t he r at .
2. Gr oups of r at s were ei t her gi ven t ap wat er or 18
g/ L sal i ne t o dr i nk. Af t er
7
days, mont hs
or
12 mont hs, t he cont rol group and sal t l oaded
~
* Present address and cor respondence t o:
Prof essor R Swamnat han
Depar t ment of Cl i ni cal Bi ochemst ry
Guy s Hospi t al
St Thomas Str eet
LONDON SE 9RT
Fax: 071 955
478
327
Copyright
1994 by
Marcel
Dekker, Inc
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DIETARY SODIUM AND Na+, K+-ATPase ACTIVITY
329
sodi umpump ( 6- 16) . Upregul at i on of sodi umt ranspor t
has been demonst rat ed i n several cel l t ypes 17-191,
i n
exper i ment al ani mal s ( 12- 16) and i n man
( 9- 11) .
The ef f ects of hi gh ci rcul at i ng l evel s of endogenous
sodi um t ranspor t i nhi bi t or on t i ssues ot her t han
ci rcul at i ng cel l s (20 ,211 i s not f ul l y document ed.
I n
ani mal model s, Na, K+- ATPase act i vi t y has been
repor t ed t o be i ncreased af t er hi gh sal t f eedi ng i n
t he renal t i ssue ( 22) , but not i n the hear t ( 23) .
I ncreased sodi um pump act i vi t y ( as measured by
rubi di um upt ake) i n vascul ar smoot h muscl e has been
shown af t er 4 weeks of hi gh sal t by Vasdev et a1 ( 24)
but not by Bradl augh et a1
25).
I n vi ew of t hese di screpant r esul t s, we examned
the ef f ect of chroni c f eedi ng of hi gh sal t di et on Na+,
K+
ATPase act i vi t y of di f f erent t i ssues i n the r at . ,
MATERI ALS AND METHODS
Adul t mal e Spr ague-Daw ey rats wei ghi ng about
250 g were used. The rat s had f ree access t o st andard
l aborat ory Pur i na rat chow ( composi t i on by wei ght :
prot ei n 23 f at 4. 5 mneral 2.5 , f i bre 6.0 and
ash
8 ) .
Unl ess ot herw se i ndi cat ed, t hey were
al l owed t o dr i nk tap wat er ad l i bi t um
Rat s were di vi ded randomy i nto t reat ment and
cont rol gr oups. Cont rol group was gi ven t ap wat er t o
dr i nk and exper i ment al groups was gi ven 18 g/ L sal i ne
t o dr i nk. Af t er 7 days ( n=12 i n each group)
,
3 mont hs
( n=14 i n each group) or 12 mont hs ( n= l i n each gr oup)
of t reat ment rat s were sacr i f i ced by cervi cal
di sl ocat i on and the hear t , l i ver , ki dney, skel et al
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330 LI, HO, AND SWAMINATHAN
muscl e ( hi nd- l i mb) and aor t a were qui ckl y exci sed and
kept i n col d hi st i di ne buf f er ( 33 mmol / L hi st i di ne,
2. 2
mmol / L EDTA, and
280
mmol / L sucr ose, pH 7. 2) . The
prot ocol f or the preparat i on of t i ssue homogenat e and
the assay met hods have been descr i bed i n detai l
el sewhere 16). Br i ef l y, the t i ssues were cut i nt o
f i ne pi eces and homogeni sed i n 10 vol of hi st i di ne
buf f er. The homogenat e was cent r i f uged and t he
supernat ant st ored at - 7OOC unt i l use.
Pr i or to assay, t he sampl es were treat ed w t h
sodi umdeoxychol at e ( DOC) i n or der t o sol ubi l i ze t he
membr anes 16)
Measurement
of
Na. K+- ATPase act i vi t v
To
reduce nonspeci f i c Mg++- ATPase act i vi t y,
homogenat es of hear t , l i ver , skel et al muscl e and renal
medul l a were prei ncubat ed w t h 5 mmol / L sodi um azi de
( f i nal concent rat i on) f or 30 mnutes at 37OC. The Na,
K+- ATPase act i vi t y of homogenat es of l i ver , hear t
,
skel etal muscl e and renal medul l a was det ermned by
t he coupl ed- enzyme assay. As the act i vi t y of vascul ar
Na, K+- ATPase act i vi t y i s l ow compared t o t he other
t i ssues K+- i mul at ed hydr ol ysi s of 3- 0-
met hyl f l uorescei n phosphat e ( 3- 0- MFP) ( 26) was used.
Al l measurement s were done i n dupl i cate.
Cowl ed enzvme act i vi t v
100
ul of t he enzyme preparat i on was pr ei ncubated
w t h and w t hout 100 ul of ouabai n sol ut i on ( f i nal
concent rat i on
5
mmol / L) f or
30
mnutes at 37OC. The
Na, K+- ATPase enzyme act i vi t y was measured by t he
decrease i n absorbence of NADH at
340
nm on a
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DIETARY SODIUM AND Na +, K+-ATPase ACTIVITY
33
1
cent r i f ugal anal yser ( Cobas Bi o, Hof f man La Roche,
Basl e, Sw t zer l and) as descri bed previ ousl y 16, 2 7 ) .
The Na+, K+- ATPase act i vi t y was cal cul at ed as the
di f f erence bet ween the ATPase act i vi t y i n the presence
and absence of
5
mmol / L ouabai n ( f i nal concent r at i on)
and was expressed at U/ mg pr otei n.
Pot assi um st i mul at ed 3- 0- MFPase assav
Homogenat e
of
aor t a was i ncubat ed w t h
20
mmol/l
of 3- 0- MFP i n Tr i s HC1. Hydrol ysi s of 3- 0- MFP bef or e
and af t er addi t i on of KC1 ( f i nal concent r at i on
10
mmol / L) was recorded. Based on t he sl opes
of
change
of f l uorescence, t he
K+
dependent phosphat ase act i vi t y
was cal cul at ed and expressed as umol of 3- 0- MFm n- mg- 1
prot ei n 16).
Protei n measurement
The prot ei n concent rat i on i n t he t i ssue
homogenat e was det ermned by the method of Lowry et a1
( 28) adapt ed to cent r i f ugal anal yser .
Resul t s are expressed as mean SEM The data
were subj ect ed t o two way anal ysi s of var i ance and
where t he nul l hypot hesi s was rej ect ed, Schef f e' s
compari sons were per f ormed; a p val ue ~ 0 0 5 was
consi dered si gni f i cant .
RESULTS
The ur i nary sodi umexcret i on i n the sal i ne gr oup
at 12 months was 27. 2 + 2. 4) mmo1/ 24 hour compared t o
1.6
+ 0. 1) mmo1/ 24 hour i n the cont rol gr oup. Fi gure
1 shows Na+, K+- ATPase act i vi t y i n t he var i ous t i ssues.
The Na, K+- ATPase act i vi t y of renal medul l a of t he
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332 LI, HO, AN D SWAMINATHAN
..
Z a
- 0 14
2 5
I
0 12
1
DAYS 3 MONTHS 12 MONTHS
SKELETAL
MUSCLE
0 06 7 1
LIVER
T
T
-
=
-
0 09
E
-:
me
0 08
a m
s
- 0 07
0 08
7
DAYS
3 MONTHS 12 MONTHS
KIDNEY
150 *+
.-
-
2 - 0 05 125
.c
d
= e
. 0 04
- E
= e
1 00
0 02 05 0
1 DAYS
3
MONT S
12 MONTHS
1 DAYS 3 MONTHS 12 MONTHS
1
DAYS 3
MONTHS
12
MONTHS
Fi gur e 1. Na, K+- ATPase act i vi t y of hear t , l i ver ,
skel et al muscl e, ki dney and aor t a i n sal t
l oaded ( hat ched col umns) and t hei r
r espect i ve cont r ol s ( sol i d col umns) .
Resul t s are mean & SEM.
*pcO.O5,
**p
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DIETARY SODIUM
AND Nat
,
Kt
ATPase ACTIVITY
333
sal t - l oaded group was hi gher at
7
days,
3
mont hs and
12
mont hs compared t o t hei r r espect i ve cont rol groups.
Na, K+- ATPaseof hear t , l i ver and skel et al muscl e
of the sal t l oaded group was not di f f erent f rom
cont rol gr oup af t er
7
days of t reat ment . The hepat i c
enzyme act i vi t y af t er 3 mont hs and 12 mont hs was
si gni f i cant l y hi gher . Enzyme act i vi t y of al l t i ssues
af t er 12 mont hs of sal t l oadi ng was si gni f i cant l y
hi gher t han cor respondi ng cont rol group.
Sal t l oadi ng f or 7 days or 3 mont hs had no ef f ect
on aor t i c Na, K*- ATPase act i vi t i es measured as
3- 0-
MFPase act i vi t y. A si gni f i cant i ncrease i n 3- 0- MFPase
act i vi t y of aor t a was f ound when r at s were chr oni cal l y
f ed on hi gh sal t i ntake f or
12
mont hs.
DI SCUSSI ON
We have used t he 3- 0- methyl f l uorescei n
phosphatase assay f or the measurement of Na+/ K+- ATPase
i n t he aor t a, as t he vascul ar aor t a has a hi gh
pr opor t i on of Na+/ K+i ndependent ATPase act i vi t y 29-
30)
and methods whi ch depend on i nhi bi t i on by ouabai n
are not sensi t i ve enough. 3- 0- met hyl f l uor escei n
phosphat ase assay has been used by ot hers f or t he
measurement of Na+/ K+- ATPaseact i vi t y i n hear t muscl e
and skel et al muscl e and found to cor rel at e wel l w t h
ouabai n bi ndi ng si t es 31-33).
Our resul t s show that i n al l t i ssues examned Na,
K+- ATPase act i vi t y af t er 12 mont hs of sal t l oadi ng was
hi gher t han that of the cont rol group. I n t he renal
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334 LI, HO, AND SWAMINATHAN
medul l a, i ncreased act i vi t y of Na, K+- ATPase was seen
as ear l y as 7 days. Swann ( 23) f ound no di f f erence i n
the Na+, K+- ATPase act i vi t y
of
the heart i n r at s
recei vi ng hi gh sal t di et f or 4 weeks. I t was al so
repor t ed that sodi um ef f l ux f rom aor t a was not
di f f erent i n r at s gi ven hi gh sal t di et f or one mont h
( 25) .
There i s overwhel mng evi dence t hat hi gh sal t
i nt ake causes t he rel ease of an endogenous Na,
K+-
ATPase i nhi bi t or ( 1, 2, 34- 38) . The presence
of
endogenous i nhi bi t or i n the ci rcul at i on i s expected t o
cause a reduct i on i n the act i vi t y of t he enzyme at
l east i n the short t erm Al t hough t he l evel of
ci rcul at i ng i nhi bi t or was not measured i n t hi s st udy,
i n other st udi es we have shown presence
of
i ncreased
sodi um t ranspor t i nhi bi t or act i vi t y dur i ng sal t
l oadi ng ( 39) as have ot hers ( 34- 38) . Thus
i t
i s
surpr i si ng t hat the act i vi t y of t he Na, K+- ATPase was
not di f f erent at 7 days i n the sal t l oaded group. One
possi bl e expl anat i on f or the apparent l ack of
i nhi bi t i on coul d be due t o di ssoci at i on of t he
i nhi bi t or dur i ng the preparat i on of the t i ssue
homogenat e ( 23, 25, 40) Another possi bl e r eason f or
the apparent l ack of i nhi bi t i on coul d be that t he
l evel s of the ci rcul at i ng i nhi bi t or are t oo l ow t o
show
a
si gni f i cant ef f ect .
Na, K+- ATPase act i vi t y of al l t he t i ssues was
i ncr eased at one year . I ncrease i n Na,
K+
ATPase
of
hear t and ki dney i n rat s gi ven hi gh sal t di et f or
4
weeks has been document ed previ ousl y ( 22, 231.
As
far
as we are aware thi s
i s the f i rst repor t
on
t he ef f ect
of
sal t l oadi ng on Na, K+- ATPase act i vi t y of l i ver ,
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DIETARY SODIUM AND
Na+,
K+-ATPase ACTIVITY
335
skel et al muscl e and aor ta. The i ncreased act i vi t y of
t he enzyme i s probabl y due t o upregul at i on of t he
enzyme. We post ul at e that sal t l oadi ng r el eases a
sodi um t ranspor t i nhi bi t or and as a resul t of t he
ef f ect of t hi s subst ance, new Na, K+- ATPase uni t s are
synt hesi sed t o br i ng the i nt racel l ul ar sodi um and
number of act i ve uni t s t o normal . Dur i ng t he
preparat i on of the t i ssues f or t he measurement
of
enzyme act i vi t y the i nhi bi t or i s washed away and the
measured act i vi t y i s t heref ore hi gher . The r esul t s
observed coul d al so be due t o changes i n ot her f act or s
regul at i ng Na, K+- ATPase ( 3) . I t has al so been
suggest ed that t he upregul at i on of Na, K+- ATPase i s
secondar y t o i ncreased i nf l ux of sodi um i nduced by
sal t l oadi ng ( 1 9 , 2 5 ) .
The di f f erence i n the t i me requi red f or t hi s
upr egul at i on var i es f rom days t o 12 mont hs. Thi s
we suggest i s due to the di f f erence i n t he t ur nover
rat e of t he enzyme i n t hese t i ssues. Ti ssues w t h a
rapi d turnover t i me such as ki dney show an
upr egul at i on w t hi n 7 days whereas aor t a t akes one
year . We have previ ousl y shown si ml ar di f f er ence i n
t he rat e of upregul at i on of Na, K+- ATPase among
t i ssues i n rat s gi ven di goxi n
16).
As Na+ K+- ATPase i n di f f erent t i ssues have
di f f er ent subuni t s ( 41) , t he di f f erence i n t he t i me
requi red f or t he change seen here may al so be due t o
t he expressi on of di f f erent i sof orms.
We concl ude that l ong termsal t
l oadi ng causes an
i ncrease i n t he act i vi t y of Na+, K+ATPase of hear t ,
l i ver , muscl e and aor t a probabl y due
t o upregul at i on.
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336
LI,
HO, AND SWAMINATHAN
ACKNOWLEDGEMENTS
Thi s st udy was par t l y suppor t ed by a grant f rom
t he Croucher Foundat i on (HK) . We grat ef ul l y
acknow edge t he hel p of Mrs Carol i ne Morgan i n t ypi ng
t he
1.
2.
3.
4.
5.
6.
7.
8.
manuscr i pt .
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Submitted: 5/04/93
Accepted: 10/22/93
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