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465

J. Physiol. (I937) 90, 465-477 6I2.744:6I2.014*46I*3

THE BEHAVIOUR OF MUSCLE FOLLOWING


INJECTION OF WATER INTO THE BODY
BY M. GRACE EGGLETON
From the Department of Physiology, Bedford College, London
(Received 16 June 1937)
ALL observers are now agreed that ingestion of a large volume of water
is followed by dilution of the blood, but opinion is divided as to the
nature and extent of this dilution. One definite type was first clearly
described by Bayliss & Fee [1930], working on decerebrate dogs,
though noted in passing by Rioch [1930] on unan.esthetized dogs
(diuresis inhibited by handling) and later by Smirk [1932] on un-
ansesthetized rabbits. In all these cases, the electrolytes of the plasma
were found to be considerably less diluted than the colloids, following
absorption of water from the gut. Moreover, the degree of dilution of
plasma colloids indicated that the blood was carrying more of the
additional water than would have been the case if this were equally
distributed throughout the water of the body.
These facts can best be explained on the assumption that some
tissues, instead of carrying their proportionate share of the additional
water, have added electrolytes to the circulating fluid. Recent work on
isolated muscle suggested that this tissue might act as a store-house for
the rapid removal or addition of electrolytes to the surrounding medium.
In the frog's isolated muscle, the muscle cells are completely impermeable
to chloride, but the relative proportion of "interspaces " (containing
chloride) to cells is found to vary with the composition of the surrounding
medium [Eggleton et al. 1937]. Muscles placed in hypotonic Ringer's
solution swell and those in hypertonic solution shrink, but the proportion
of muscle occupied by chloride is less than normal in the former case
and greater in the latter case. It appears that the muscle cells, which
absorb water when the surrounding medium is hypotonic, press upon the
"interspaces ", thus diminishing their volume. If muscle in situ were to
behave in a similar fashion, it might account quantitatively for the
results obtained by Bayliss & Fee, since muscle tissue forms so large
466 M. G. EGGLETON
a proportion of the total body weight. Experiments were made, there-
fore, in which the chloride concentration in blood plasma and in muscle
were estimated, together with the total solid content of both tissues,
before and after the absorption of a large volume of water from the
intestine.
EXPERIMENTAL
The experiments were performed on cats which had received their
last meal 18 hours previously but had free access to water. Anesthesia
was induced by nembutal (0-65-0.7 c.c. per kg.), given intraperitoneally.
Total solids of both plasma and muscle were determined by drying to
constant weight at 100-110° C. (24 hours). Chloride in the trichloracetic
acid filtrate of either tissue was determined by electrometric titration, in
the manner described by Eggleton et al. [1937].
Preliminary experiments showed that pairs of muscles contained
closely similar concentrations of chloride, so that one could be used as
a control on the other, and that the variation in water content of the
mixed samples was small. Different pairs of muscles varied in their
chloride content to some extent, the gastrocnemii always containing the
lowest and the sartorii the highest concentration, as is the case in the
frog. There was also a considerable variation in the ratio of chloride
concentration in muscle to that in plasma among different animals, a
difference probably referable in part to variation in age [Irving &
Manery, 1936]. The figures given in Table I indicate the degree of
random error involved in the whole technique and are representative of
the average results obtained on normal muscles. The question as to
TABLE I. The chloride and water content of muscles and blood plasma in a cat
under nembutal ansesthesia
A
Cl' concentration
in muscle B
mg./100 g. Cl' concentration
___ _A in blood plasma Ratio
Muscle Left leg Right leg mg./lOO g. A/B
Gastrocnemius 59 60 407 0-145
410 0-148
Semitendinosus 78 72 - 0*191
0-177
Gracilis 81 79 0 199
0*194
Rectus femoris 96 95 0*234
0-232
Water content (p.c.)
76*3 76-0 92-0
92*0
UPTAKE OF WATER BY MUSCLE 467
whether the observed ratio of muscle to plasma chloride is the same as
that obtaining in the unansesthetized state has not been seriously in-
vestigated; a few experiments on decerebrate cats gave similar values.
In an experiment the following technique was adopted. Half to
one hour after induction of anwesthesia, one blood sample (5-6 c.c.) was
obtained from a carotid artery which had been tied off, and samples of
muscles from one leg; the muscles were freed from surrounding fascia,
entering vessels clamped, ligatures applied to the muscles and the inter-
vening part removed. One portion was taken for estimation of total
solids into a weighing pot of known weight and another (1-3 g.), for
chloride estimation, weighed and dropped into 4 p.c. trichloracetic acid.
Overnight soaking in this fluid resulted in the muscle being more easily
disin'tegrated by grinding the following day.
The water (6 p.c. of the body weight) was then injected. In most
cases the injection (lasting 15-20 min.) was made into a loop of small
intestine, brought to the surface through a small abdominal opening,
the technique used by Bayliss & Fee. A blood sample, the remaining
muscle samples, and a further blood sample were removed 1'5-2-5 hours
after the end of the injection. The intestine was then removed and
opened to ensure that no water had been left unabsorbed. In some
experiments the bladder was washed out by means of a cannula placed
through the upper part of the urethra above the pubis, but only a few
cubic centimetres of urine were ever excreted during the experiments.
In others the ureters were tied as they entered the bladder. In one or
two cases the injection was made directly into the portal vein at a steady
rate of about 1*5-3-0 c.c. per min.; the second blood and muscle samples
were then removed 20-40 min. after the end of the injection, which had
occupied 1-1.5 hours.
Blood samples were usually oxalated (with a known amount) and the
tubes tightly stoppered with cotton-wool until after centrifuging.
Standing for 3 hours under these conditions led to an increase of approxi-
mately 1 p.c. in the chloride content of the plasma, presumably due to a
chloride shift under the lower CO2 pressure. Collection of the blood
under paraffin gave results identical with those obtained on freshly
collected blood treated with oxalate.
RESULTS
The results obtained were essentially similar to those found by
Bayliss & Fee in decerebrate dogs. -In every case, following absorption
of water, the total solids of the plasma were diluted much more than was
PH. XC. 30
468 M. G. EGGLETON
its chloride. Analysis of muscle showed that in this tissue the reverse
process had occurred. The total solids were always diluted much less
than the chloride. The average results of all experiments (10) are shown
diagrammatically in Fig. 1. There was considerable variation in different

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.

been absorbed and the volume of "interspaces" slightly reduced. In


the experiment quoted in Fig. 2, muscles were not analysed, but analogy
with other experiments suggests that in the first 15 min., during the
injection, chloride passed rapidly into the gut and the resulting hypo-
tonicity of the plasma led to uptake of water by the muscle cells (leading
to actual concentration of the plasma solids), with consequent reduction
of "interspace" fluid. This phase probably outlasted the injection by
5-10 min. and was followed by a steady rapid absorption of the weak
salt solution with subsequent further uptake of water by the muscle
cells and decrease of "interspace" fluid.
This interpretation of the results is strengthened by the fact that
when the water is injected intravenously the two curves do not cross.
UPTAKE OF WATER BY MUSCLE 475
The initial stages of such an experiment are shown in Fig. 3; unfor-
tunately, increasing haemolysis completely masked the dilution of plasma
solids in the latter part of the experiment. It was present in small
degree even in the earlier samples and the dilution of total solids was
therefore even greater than shown in the figure.
DIscusSION
It has been shown that the observed changes in water and chloride
content of muscle and blood plasma following injection of water into
cats can be satisfactorily explained from considerations of simple osmotic
equilibria, if it be assumed that the chloride-containing part of the
muscle (one-fifth of the whole) is separated from the remaining chloride-
free part by membranes completely impermeable to chloride. In the
frog's isolated resting muscle, direct proof of this assumption has been
obtained [Eggleton et al. 1937], but even here there is no proof as yet
of the quantitative identity of these "interspaces" with the anatomical
intercellular spaces. An attempt to count the latter directly by histo-
logical means has met with little success; both slow and rapid fixation
led to obvious shrinkage of the cells. In the muscle in situ (whether frog
or mammal) there is no proof that the cells are always or completely
impermeable to chloride.
So long as experimental results can be explained on the simplest
hypothesis (osmotic), the assumption that chloride is present only in
the "interspaces" of muscle seems justified. Increase in "interspace"
fluid as a result of injection of normal saline solution [Hastings &
Eichelberger, 1937] can be so explained, as a result of increase in
capillary filtration pressure, since the surrounding structures allow a
certain degree of swelling of the muscle and in this case there is no need,
on osmotic considerations, for swelling of the cells. Similarly, on in-
jection of hypertonic salt solutions [Eichelberger & Hastings, 1937b,
and experiments done in the present series], the resulting shrinkage of
muscle cells, with consequent reduced pressure in the tissue spaces, and
the increased capillary pressure due to increased volume of plasma, will
lead to an increase in "interspace" volume. The results of Fenn &
Cobb [1936], however, showing an increase in "interspace" volume
following stimulation of muscles in the rat, cannot be readily explained
in terms of this hypothesis, for contraction is accompanied by an increase
of osmotically active substances in the cells and should lead to swelling,
with a consequent decrease in intercellular volume. It is possible that
stimulation may render the muscle cells partially permeable to chloride.
476 M. G. EGGLETON
The ratio of chloride concentration in muscle to that in plasma,
which has been taken as an approximate measure of "interspaces ",1 is
reasonably constant in mammals. Winter [1934] gives values ranging
from 14-5 to 15*5 p.c. for a series of mammals, with 12-5 p.c. in the cat;
Saiki [1908], found a value of 13 p.c. in the pig; and in this series on the
cat an average value of 18 p.c. was found in the normal and 15-5 p.c.
following injection of water. In some other experiments on rats, values
ranging from 14 to 17 p.c. were obtained. It was mentioned earlier that,
of the leg muscles analysed, the gastrocnemius contained always the
lowest proportion of "interspaces" and the sartorius the highest. The
latter muscle appears to possess this property in common with other thin
flat muscles, for the rectus abdominis of the dog [Eichelberger &
Hastings, 1937 a] gave comparable values of 20-25 p.c. It seems likely
that the differences noted among various species by different observers
are due to the particular muscle analysed and to the manner in which
the animal is killed, in view of the effect of haemorrhage noted earlier,
rather than that there is any one value peculiar to each species.
SUMMARY
1. The observation of Bayliss & Fee, that absorption of water
from the gut results in a greater dilution of plasma colloids than of
electrolytes, has been corroborated.. In cats, under nembutal anaesthesia,
absorption of water (6 p.c. of body weight) resulted, on the average, in a
9 p.c. dilution of plasma chloride and a 16-5 p.c. dilution of total solids.
2. In the accompanying dilution of muscle constituents, the reverse
effect was observed: the chloride was diluted 23 p.c. and the total solids
only 5 p.c.
3. These results are explicable on considerations of simple osmotic
equilibria if it be assumed that (a) the chloride of muscle is contained
only in the "interspaces " and (b) the membranes of the muscle cells are
impermeable to those solutes which are responsible for maintaining the
osmotic pressure. The carriage of excess fluid by the blood plasma can
then be attributed to the fact that muscle as a whole is not a freely
distensible tissue.
4. In the early stages of such experiments, the plasma chloride is
diluted to a greater extent than the total solids, and only later is the
1 Eichelberger & Hastings [1937a] have recently devised a more accurate method
of calculation of the size of "interspaces " (based on the relative chloride concentrations in
serum and muscle) by correcting for the Gibbs-Donnan effect of a higher protein con-
centration in serum than in "interspace" fluid and on the higher water content of the
latter.
UPTAKE OF WATER BY MUSCLE 477
reverse condition attained. This effect is not observed when the water is
injected intravenously and provides additional evidence of the passage of
chloride into the gut before absorption of the water can occur.
I am greatly indebted to Dr L. E. Bayliss for helpful suggestions in the interpretation
of the results and also for constructing the electrical unit of the apparatus used in measure-
ment of chloride.

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.

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