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Muscle Plasma
20
0~15
Fig.1. TerltvdereofdltoofttlsldanofclrdinmTotal
solids
5mTotal
solids
Fig. 1. The relative degrees of dilution of total -solids and of chloride in muscle and blood
plasma resulting from absorption of water (6 p.c. of body weight) in cats (average of
ten experiments).
TABLE II. Dilution of plasma and muscle in cats following injection of water
(6 p.c. of body weight)
Muscle Plasma
Dilution of Dilution of Dilution of Dilution of
chloride total solids chloride total solids
Exp. p.c. p.c. p.c. p.c.
11 16 5.5 6 20
12 31 6 10 16
14 25 6 11 15
15 31 2 7-5 18
16 25 5 5 12
18 21 6-5 11 15
19 25 2 8 22
24 20 6-5 13 16-5
31 13-5 5 10 15
33 21 3-5 7-5 16
Average 23 ±3* 5 ±0.3* 9 ±0.6* 16-5 ±0.8*
* Standard deviation of mean, (1)') where r is the individual difference from
the mean and n is the number of observations.
experiments, though the changes were invariably in the same direction.
In Table II are shown the average changes in each experiment. The
UPTAKE OF WATER BY MUSCLE 469
actual experimental results from which the figures in Table II have
been calculated, and the range of variation amongst different muscles in
any animal, are illustrated in the following more detailed description of
a typical experiment.
Exp. 12. Cat, ?, 3-25 kg. weight.
205 c.c. water injected: 15 c.c. unabsorbed at end.
Pla8ma: Total solids initially ... 8-1 p.c.
,, finaly ... 7-1, 6-95 p.c.
=16 pc. dilution.
Cl' initially ... ... 408 mg./100 g.
finally ... ... ... 371, 371 mg./100 g.
= 10 p.c. dilution.
Muscle: Total solids initially ... 25-3 p.c.
,, finally
=6 p.c. dilution.
... 23-9 p.c.
Cl' in mg./100 g.
Initially Finally
Gastrocnemius 59.5 43-5
Semimembranosus 58 49
Semitendinosus 67-5 52
Gradilis 69 50
Average 63-5 48.5
=31 p.c. dilution.
Other factors affecting the concentration of chloride in muscle
An attempt was made to assess the relative importance of possible
factors responsible for the variation shown in different experiments.
Undoubtedlythe state of nutrition of the animalwould affect the numerical
result, for although the water injected was in all cases 6 p.c. of the body
weight, a greater degree of dilution would be expected in very fat
animals since the water-containing tissues form a smaller proportion of
the total weight. Three other factors were investigated experimentally:
haemorrhage, change in body temperature, and peritoneal effusion
resulting from handling of the intestines. Small variations in these
three factors probably occur in all experiments and their possible
influence on the chloride and water content of the muscle and plasma
was magnified by inducing extreme variations in each. The changes
produced in the two tissues are shown in Table III.
Under the influence of ansesthetic alone, there was a slight loss of
water and chloride from the muscles, but the composition of the blood
was unaffected. A rise of 30 C. in body temperature resulted in dilution
of the plasma solids, though not at the expense of muscles, while shivering
(mainly of the trunk and forelimbs) resulted in concentration of plasma
solids, a result in accord with the findings of Calvin et al. [1933]. The
30-2
470 M. G. EGGLETON
TABLE III. The effect of various treatments on the chloride and
water content of muscle and plasma
Muscle Plasma
Changes (p.c.) Changes (p.c.)
produced in produced in
concentration of concentration of
Total Total
Exp. Treatment Chloride solids Chloride solids
6 Anuesthetic alone (nembutal) - 1 -0 5
40 ,, - 4 +1-5 +1 0
4 Severe haemorrhage -12 -2
21 ,, - 8 -05 -05 -1
25 Peritoneal effusion -10 +3 -3-5 3-5
36 ,, - 6 +2 0 -1-5
38 ,, - 9.5 +2 0 -3
23 Temperature increase - 2 0 +2 -4.5
37 Shivering - 2 +2 -1 +5
+ Denotes concentration, - denotes dilution.
results with peritoneal effusion are interesting in showing clearly the
regulating relationship, in respect of water and chloride, of muscle on the
plasma. The peritoneal exudate is roughly a protein-free plasma filtrate
(though it contains a small amount of protein) and the loss of both water
and salt from the plasma is made good by the muscles. The effects
produced by excessive haemorrhage are less easy to explain, but again it
is seen that appreciable dilution of plasma salt has been prevented by
loss of salt from the muscles. In the water-injection experiments,
haemorrhage (apart from the 10-15 c.c. taken in blood sampling) was
usually negligible, peritoneal effusion slight and temperature change
within less than 10 C., but their combined effects may well account in
part for the individual variations.
In both the water-injection experiments and the control experiments
just quoted, it appears that the muscles respond to changes in the water
and electrolyte content of the blood plasma by suitable alteration in
their own composition. The question arises as to w.hether this behaviour
is regulated by means of some hormonal or nervous influence, or whether
it is inevitable from considerations of simple osmotic equilibria. Analysis
of the conditions in muscle, in so far as these are known, suggests that
the latter simpler hypothesis is sufficient to account for the observed
facts. The capillary walls are practically impermeable to protein, but
freely permeable to water and electrolytes. In respect of these, the plasma
can be considered as in equilibrium with the fluid of the tissue spaces.
This fluid, on the other hand, is separated from the muscle cells by
membranes impermeable to practically everything but water. In the
UPTAKE OF WATER BY MUSCLE 471
frog's isolated resting muscle, neither Na nor Cl [Fenn et al. 1934]
lactate [Ghaffar, 1935], creatine [Eggleton, 1930], phosphate [Eggle-
ton, 1933], glucose [Eggleton, 1935], nor carnosine [Eggleton &
Eggleton, 1933] can penetrate the cells, nor will the cell constituents
leak out at any appreciable rate. Given these conditions, and assuming
that there is no production of osmotically active substances within the
muscle cells, the following calculations can be made.
1 kg. cat contains 600 c.c. water, of which the muscles contain about half, 300 c.c.
Addition of 60 c.c. water to the animal must result in an overall dilution of 600 = 60
and any celUs permeable only to water muwt ab8orb water until their content8 are diluted in this
proportion.
Of the water in muscle . .. ... ... ... ... 300 c.c.
18 p.c. is in the "interspaces" (average figure derived from ratio
of chloride concentration in muscle to that in blood plasma) ... 54 c.c.
Hence the cells contain initially ... ... ... ... 246 c.c.
Since they must swell in the ratio 660/600, this 246 c.c. increases to 270 c.c. But direct
analysis of the whole muscle shows that it has increased in weight by only 5 p.c., i.e. the
water of the muscle (which constitutes 75 p.c. of its substance) has increased by
100 x5=6.7p.c.,
i.e. the initial 300 c.c. water has increased to 320 c.c.
Muscle water finally ... ... ... ... 320 c.c.
Muscle cell water finally ... ... 270 c.c.
Therefore, "interspace" water finally ... 50 c.c.
which is 15.6 p.c. of the whole muscle water, i.e. the " interspaces " must have been reduced
from 18 to 15*6 p.c. Experimentally, an average value of 1565 p.c. is found.
Furthermore, initially each 100 c.c. of muscle water contained 18 c.c. of chloride solution
equivalent in strength to that of plasma chloride and finally 15*6 c.c. of diluted chloride
solution, which is equivalent to 600 x 15-6 c.c. = 14-2 c.c. of original solution, i.e. the muscle
chloride has been diluted by 22 p.c.; the average figure found experimentally is 23 p.c.
If this calculation be applied to each experiment separately, then,
from the observed overall dilution of solids in the muscle and dilution of
chloride in the plasma (which represents the overall dilution of body
water), the expected dilution of chloride in the muscle can be calculated.
These figures are compared in Table IV with the observed results.
In eight of the ten experiments the agreement is surprisingly good.
Of the two exceptions, Exp. 11 was the first of its type to be performed
and therefore more liable to random errors in technique and Exp. 16
was made on an animal received straight from the dealer (i.e. its previous
food history was unknown) and noted as suffering from bad respiratory
infection. Normally the animals are kept in the laboratory for a few
days preceding the experiment. If errors in technique are not responsible
472 M. G. EGGLETON
TABLE TV. Comparison of the observed value of dilution of chloride in muscle with that
calculated from observed dilutions of chloride in plasma and solids in muscle. (For
calculation see text) Dilution of Cl' in muscle
C
A B Proportion I
Dilution of Dilution of of "inter- Calculated II
Cl' in solids in spaces"* from A, B Observed
plasma muscle initially and C value
Exp. p.c. p.c. p.c. p.c. p.c.
11 6 5*5 18-5 0 16
12 10 6 15-5 18 22
14 11 6 15-5 22 25
15 7*5 2 21*5 23 31
16 5 5 18 0 24
18 11 6*5 16 21 21
19 8 2 19 27 25
24 13 6-5 15*5 29 21
31 10 5 16 23 13-5
33 7.5 3.5 16.5 19 21
tAverage 23 ±1-7t 22.5 ±2-81
* "Interspaces" measured as the ratio of chloride concentration in muscle to that in
plasma.
t Excluding Exps. 11 and 16. t Standard deviation of mean.
for either or both of these anomalous results, then in these animals the
muscle cells must have absorbed more than the expected amount of
water; either they were permeable to substances other than water, or,
for some reason, an increase in osmotically active substances occurred
inside them during the course of the experiment. The general trend of
the results, 'however, suggests that simple osmotic forces are sufficient
to explain the behaviour of muscle under these conditions.
It will be noted that the calculated value of dilution of chloride in
muscle is derived only from the observed dilution of chloride in the
plasma and of solids in the muscle, and the initial size of the chloride-
containing "interspaces". The amount of water entering the system is
immaterial to the calculation, also the total dilution of solids in the
plasma. Their relatively great dilution appears to be due to the fact that
the muscles, owing to the inelasticity of the fascia surrounding them,
cannot swell to any considerable extent. When the cells swell, therefore,
pressure increases in the "interspaces", resulting in lessened filtration
from the capillaries and/or greater outflow of lymph, with consequent
reduction in "interspace" fluid. As a result of this indistensibility of
muscle, the plasma and other tissues more distensible than muscle have
to carry more than their share of the additional fluid.
In the case of peritoneal effusion, the muscle loses water and chloride
in approximately the proportion present in the "interspace" fluid. If
the supporting structures of the muscle are normally in a slight state of
UPTAKE OF WATER BY MUSCLE 473
tension, the decreased filtration pressure in the capillaries, due to loss of
saline in the peritoneal effusion, would satisfactorily account for the
small decrease in "interspace" fluid observed.
Changes in blood plasma during water absorption
It is now generally accepted that absorption of water is preceded by
passage of chloride into the gut. In some early experiments on the
effect of water-drinking in man, Priestley [1916] found a 5 p.c. dilution
of the chloride in plasma and attributed it to withdrawal of chloride into
the gut prior to absorption of the water. Smirk [1933] postulated the
0 0.-
; 10
go
5_-- __
15
I , I
oatecron A15 30 45 60 75 90
min.
Fig. 2. Exp. 27. Cat 3-6 kg. Dilution of total sohids (o ------ o) and of chloride ( x x)
in blood plasma during and after injection of water (6 p.c. of body weight) into the
intestine of a cat.
same effect and, in rats which had received water, found 0 55 p.c. NaCl
present in the fluid in the gut during the height of absorption. In view
of the type of dilution observed when equilibrium is established under
the conditions of the experiments reported here, it seemed of interest to
analyse the type of dilution occurring in the earlier stages. If chloride
was passed into the gut prior to absorption of the water, the dilution of
chloride in the plasma should precede the dilution of total solids, although
ultimately the positions are reversed. Such indeed was found to be the
case. In Fig. 2 are shown the degree of dilution of chloride and of total
solids in plasma following injection of water into the intestine. Four
such experiments were performed and although the shape of each curve
474 M. G. EGGLETON
varied, the relative positions of the two were consistently the same as
that shown in the figure.
The results are fully in accord with the suggestion that chloride
passes into the gut before absorption of the fluid can occur. The crossing
of the two curves would then coincide with the rapid absorption of a
dilute salt solution. Variation in the exact shape of the curves is probably
due to the rate at which chloride passes into the gut and the rapidity
with which the muscles react to changes in the plasma. Analysis of
muscles at an early stage (13-30 min.) shows that some water has already
0
455
NO
10
0 15 30 45 60 75 90
min.
Fig. 3. Exp. 28. Cat 4 kg. Dilution of total solids (o ------ o) and of chloride ( x x)
in blood plasma during intravenous injection of water. The injection was continuous
throughout the experiment, at the rate of ca. 3 0 c.c. per min. into the portal vein.
REFERENCES
Bayliss, L. E. & Fee, A. R. (1930). J. Physiol. 70, 60.
Calvin, D. B., Smith, A. H. & Mendel, L. B. (1933). Amer. J. Phy8iol. 105, 135.
Eggleton, P. (1930). J. Physiol. 70, 294.
Eggleton, M. G. (1933). Ibid. 79, 31.
Eggleton, M. G. (1935). Ibid. 84, 59 P.
Eggleton, M. G. & Eggleton, P. (1933). Quart. J. exp. Physiol. 23, 391.
Eggleton, M. G., Eggleton, P. & Hamilton, A. M. (1937). J. Physiol. 90, 167.
Eichelberger, L. & Hastings, A. B. (1937a). J. biol. Chem. 118, 197.
Eichelberger, L. & Hastings, A. B. (1937b). Ibid. 118, 205.
Fenn, W. 0. & Cobb, D. M. (1936). Amer. J. Physiol. 115, 345.
Fenn, W. O., Cobb, D. M. & Marsh, B. S. (1934). Ibid. 110, 261.
Ghaffar, A. (1935). Quart. J. exp. Physiol. 25, 229.
Hastings, A. B. & Eichelberger, L. (1937). J. biol. Chem. 117, 73.
Irving, L. & Manery, J. F. (1936). Biol. Rev. 11, 287.
Priestley, J. G. (1916). J. Physiol. 50, 304.
Rio ch, D. McK. (1930). Ibid. 70, 45.
Saiki, T. (1908). J. biol. Chem. 4, 483.
Smirk, F. H. (1932). J. Physiol. 75, 81.
Smirk, F. H. (1933). Ibid. 78, 127.
Winter, K. A. (1934). Z. ges. exp. Med. 94, 663.