CHANGES IN THE COMPOSITION OF THE URINE AFTER MUSCULAR EXERCISE.*

BY D. WRIGHT WILSON, W. L. LONG, H. C. THOMPSON, AND SYLVA THURLOW.

(From the Department of Physiological Chemistry, School of Medicine, Uni- versity of Pennsylvania, Philadelphia.)

(Received for publication, July 6, 1925.)

The effect of strenuous muscular exercise on the body has been studied extensively. The responses of many of the mechanisms which take part in attempting to furnish oxygen and fuel and to remove carbon dioxide and other waste products have been analyzed in considerable detail. Studies on the respiratory exchange, the blood, and the urine have contributed much toward furnishing us with a detailed knowledge concerning the extent to which the mechanisms for maintaining neutrality in the organism are brought into play.

The work of Barr and his collaborators (1923, a, b, and c) has strikingly demonstrated the relatively great variations in the re- action of the blood brought about by short periods of strenuous exercise. One would expect that such large variations in t,he blood would be accompanied by a response by the kidney resulting in similarly great changes in the composition of the urine. The data in the literature which permit such comparisons are, however, meager and sometimes inconclusive and contradictory. In most of the older work, long periods of exercise and long periods of urine collection were employed. We believe that our studies of short periods of strenuous exercise and short periods of urine collection have yielded- data of greater value for showing the immediate effects of the exercise and attempts at compensation on the part of the body.

* A preliminary announcement was made in the Proceedings of the Society for Experimental Biology and Medicine (Wilson, D. W., Long, W. L., Thompson, H. C., and Thurlow, S., Proc. Sot. Exp. Biol. and Med., 1923-24, xxi, 425).

755

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756 Urine after Muscular Exercise

EXPERIMENTAL.

Urine was collected from normal men at frequent intervals before and after short periods of strenuous exercise. In most of the experiments the urine was collected every 10 minutes. No difficulty was experienced in voiding at such frequent intervals and the data indicate that such short periods of collection are quite suitable for studies such as these and ought to be useful in other connections. It is impossible to state how much overlap- ping of periods occurred due to the incomplete removal of all urine from the urinary system but it obviously did not interfere with the main purpose of the experiment. Low urine volumes after exer- cise associated with an increased excretion of certain constituents, together with a moderately constant output of creatinine, demon- strate that the observed variations are not due to errors of col- lection. Our controls demonstrate that our experiments were of sufficiently brief duration not to be influenced appreciably by the diurnal variations which have been shown t,o occur by Simpson (1924) and others.

The urines were voided directly into measuring cylinders con- taining toluene to diminish the loss of COz and the consequent change in reaction. After recording the amount collected, the sample was diluted to constant volume with boiled neutral dis- tilled water and stirred carefully under toluene. The reaction was estimated at once calorimetrically. The acidity was deter- mined by titrating with phenolphthalein. In the early experi- ments, nearly all of a 5 or 10 minute specimen of urine was used, potassium oxalate added, and the titration made without dilu- tion. The titration varied from 1 to 5 cc. In the later experi- ments, one-fifth or one-tenth of a 10 minute specimen was used, diluted with neutral distilled water until there was no tendency for the calcium phosphate to precipitate (Fiske, 1921), and the titration carried out with a micro burette. The titrations varied from 0.2 to 1.0 cc. Ammonia was determined by the micro method of Folin and Bell (1917), creat,inine by Folin’s micro method, total nitrogen by Kjeldahl, and phosphates by Briggs’ (1922) modification of Bell and Doisy’s (1920) method. Chlorides’ were determined by a slight modification of Whitehorn’s (1920-

1 Miss Cecilia Riegel carried out the chloride analyses.

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Wilson, Long, Thompson, and Thurlow 757

21) method, using 1 cc. of urine. The quantity of urine passed in 10 minutes is sufficient for suitable analyses by these methods. Determinatiors for acidity, total nitrogen, and chlorides were made in duplicate. Calorimetric determinations for pH, ammonia, phosphate, and creatinine were not regularly made in duplicate though many duplicates were run.

All of the experiments were carried out in the early afternoon on days when no lunch had been taken and only the ordinary activity of a student had been undergone. After a suitable fore period, the urine was voided and exercise was begun at once by running up and down one flight of stairs as rapidly as possible, swinging the arms vigorously. After the exercise the subject, rested in a chair for 10 or 15 minutes and then usually resumed some laboratory work involving, but little activity.

In Tables I to XII are recorded the data of a number of experi- ments performed on two subjects. All data involving quantity output are esprcssed in terms of excretion per minute.

In the early experiments (Tables I, II, III, and IV) running up and down one flight of stairs was undertaken at a moderate rate of speed. The exercise extended over periods of from 1 to 3 minutes in which time from two to seven round trips were made. This amount of exercise was strenuous and led to hyperpnea, but later the rate of running was about doubled. The urine was collected in periods of 5 or 10 minutes and in one case 2 minute periods were made use of immediately after the esercise.

The data from these experiments demonstrate that after short periods of strenuous exercise there is a sharp rise in the excre- tion of acids and ammonia and a diminution in pH and urine volume. An attempt was made in these and in subsequent esperi- ments, frequently without success, to eliminate the change in urine volume after exercise by drinking water. The rise in the acidity factors reached a maximum in about 10 to I5 minutes after exercise and thereafter diminished, returning to normal in 30 to 45 minutes.

An examination of the results obtained in the 2 and 5 minute collections during and after exercise shows that there is a delay of several minutes before the changes described above begin to appear in the voided urine. During this interval the pH changes little or not at all, but the titratable acidity falls. The urine volume

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758 Urine after Muscular Exercise

TABLE I.

May 10, 1923. Strenous exercise 3 min., 4.18-4.21 p.m. 7 round trips. Subject W. Ingested 209 cc. water at 3.50 p.m. Figures represent excre- tion per minute:

Time.’ Period. VOlUW3.

p. m.

4.08 4.13 4.18

min.

5 5 5

5 5 5 5 5 5 5 5 5 5 5

cc.

0.9 0.7 1.2

4.23 4.28 4.33 4.38 4.43 4.48 4.53 4.58 5.03 5.08 5.13

0.8 1.0 2.0 1.6 1.0 0.8 0.8 0.8 0.9 0.9 1.0 5.2

PH -

6.2

- Acid.

cc. 0.1 N

0.34 0.31 0.34

0.20 0.60 0.90 0.73 0.60 0.35 0.42 0.43 0.46 0.30 0.37

* Column 1 records the end of the period of collection in all tables.

TABLE II.

May 12,1923. Strenuous exercise 2 min.. 12.07-12.09 n.m. 3 round trips. Subject W. Figures represent excretion per minute. -

Time.

a. m.

11.52 11.57 P. m.

12.02 12.07

12.12 12.17 12.22 12.27 12.32 12.37 12.42

Period.

min. 5 5

5 5

5 5 5 5 5 5 5

VOIUUICI.

cc.

1.3 1.3

1.3 1.2

0.7 0.9 0.8 0.5 0.4 0.4 0.8

-

_-

-

PB Acid.

6.6 6.9

cc. 0.1 N

0.21 0.16

6.6 6.6

6.3 6.0 5.0

5.0 5.2 5.4

0.20 0..17

0.13 0.49 0.49

0.34 0.29 0.44

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Wilson, Long, Thompson, and Thurlow 759

tends to be lower than that of the periods preceding or following. It is difficult to be certain in these very short periods of collection

TABLE III.

May 17, 1923. Strenuous exercise 1 min., 12.59-1.00 p.m. 3 round trips. Subject W. Ingested 200 cc. water at 12 noon, 50 cc. at 1.02, and 50 cc. at 1.40 p.m. Figures represent excretion per minute.

Time. Period.

p. 7%

12.38 12.48 12.58

1.03 1.13 1.23 1.33 1.43 1.53 2.03

min. 10 10 10

5 10 10 10 10 10 10

VOlUUl.3. PH Acid.

cc.

4.5 3.4 4.0

6.5 6.5 6.6

cc 0.1 N cc. 0.1 N

0.247 0.133 0.207 0.108 0.230 0.141

0.7 6.6 0.037 2.6 5.5 0.550 1.8 5.5 0.403 1.9 5.7 0.316 0.7 5.7 0.204 0.6 5.7 0.262 0.9 5.7 0.280

NHJ

0.407 0.315 0.243 0.154 0.168 0.208

TABLE IV.

May 23, 1923. Strenuous exercise 1 min., 11.00-11.01 a.m. 2 round trips. Subject W. Ingested 100 cc. water at beginning of experiment. Figures represent excretion per minute.

Time.

a. m.

10.40 10.50 11.00

11.02 11.04 11.14 11.24 11.34 11.44 11.54 12.04

T Period.

min.

10 10 10

2 2

10 10 10 10 10 10

cc. 1.0 1.1 2.1

7.0 7.2 7.0

2.1 7.0 1.5 6.3 4.2 5.5 3.6 6.0 5.6 6.8 2.3 6.9 1.2 7.1 0.9 7.2

PH Acid.

cc. o.! N ec. 0.1 N

0.096 0.078 0.088 0.069 0.160 0.106

0.479 0.213 0.242 0.152 0.128 0.112

0.279 0.121 0.102 0.087 0.066

that the low acidities and urine volumes are real on account of the uncertainty of complete emptying of the bladder. Some difficulty

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760 Urine after Muscular Exercise

was experienced in voiding during the first few minutes after exercise due to fatigue. It is thought, however, that the observa- tions are reasonably correct and may perhaps be explained by a sudden shutting down of activity on the part of the kidney due to diminished blood flow through it.

The pH of such urine specimens must also be influenced by the contamination of the specimen by the portions of fluid still remain- ing in the urinary tract. The fact that Barr and his collabora- tors have shown that there is an immediate and rapid change in the reaction of the blood in strenuous exercise suggests that the 2 or 3 minute delay in observing any change in pH of the urine represents the period of time necessary to flush out the urinary tract with the diminished flow of urine.

A more extended series of analyses on each urine specimen was planned and in order to have sufficient material for analysis it was decided t.o make collections in 10 minute periods. Control experiments were carried out in which collections were made under conditions similar to those upon which the exercise was super- imposed. Each experiment was started early in the afternoon, on a day when no lunch had been eaten, and the accustomed activities had occurred in the morning (except in one control experiment). During the experiments moderate activity in the laboratory was continued. Attempts were made to maintain the urine volumes constant by drinking water during the experiments.

An examination of the data obtained from four control experi- ments on two individuals shows that during the period of 2 hours or less used for the experiment no great variations occurred although both chlorides and total nitrogen diminished at a rnt.her uniform rate. In a single experiment the inorganic phosphat,e escretion was fairly constant.

While considering normal values it is of interest to examine the magnitude of certain of the data obtained in bhe experiments on the different subjects, making use of the normal values included in the fore pefiods of the exercise experiments also. The urines of Subject W. were almost always more alkaline than those of Subject L. The range was: Subject W., pH 6.2 to 7.4; Subject L., pH 5.4 to 6.2. The single exception occurred in a control experi- ment on Subject W., undertaken after an experiment involving strenuous exercise 3 hours previously. The urines with high pH

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Wilson, Long, Thompson, and Thurlow 761

were found to contain the minimum amounts of titratable acids and ammonia. In the one experiment which permitted the calcu- lation, the ammonia nitrogen was found to be only 1 per cent of the total nitrogen at a time when the pH ranged between 6.6 to 6.9. The analyses were carefully checked so that we are confident that the results are correct. Values of the percentage of ammonia

TABLE V.

May 21,1923. Control. Subject W. Ingested 100 cc. water at 11.55 a.m. Figures represent excretion per minute.

Time. Period. Volume. PH Acid. NHa Creatinine.

p.m. mm. cc. cc. 0.1 N cc. 0.1 N ml.

12.10 10 0.95 7.4 0.091 0.055 1.05 12.20 10 2.15 7.1 0.133 0.057 1.15 12.30 10 3.40 6.9 0.168 0.089 1.03 12.40 10 4.25 6.9 0.212 0.091 1.28 12.50 10 3.15 6.9 0.195 0.082 1.08

1.00 10 2.35 7.0 0.181 0.080 1.05

TABLE VI.

May 30, 1924. Control. Subject W. (After strenuous exercise in morn- ing.) Figures represent excretion per minute.

Time. Volume. pH

p. m. cc.

2.20 3.90 5.3 2.30 4.55 5.3 2.40 5.90 5.4 2.50 6.80 5.6 3.00 3.60 5.2 3.10 2.35 5.1 3.20 1.60 5.1 3.30 1.50 5.0 3.40 2.10 5.3 3.50 1.30 5.1

Acid. NH2

cc. O.lN EC. 0.1 N

0.323 0.252 0.294 0.237 0.346 0.254 0.384 0.231 0.323 0.206 0.364 0.242 0.399 0.241 0.399 0.251 0.388 0.249 0.413 0.222

Total N NHs-N Total N

mg. pm cent

9.56 3.69 10.10 3.29 10.95 3.25

8.48 3.82 7.74 3.72 8.10 4.19

7.68 4.56 7.71 4.34 7.71 4.02

Cl

cc. 0.1 N

1.48 1.23 0.93 0.83 0.88 0.70 0.85 0.75 0.73 0.73

nitrogen as low as these may be inferred from some of Simpson’s data which include figures for ammonia and urea. It is apparent that, for urine specimens collected during short periods, the nor- mal values for the proportion of ammonia nitrogen to total nitrogen will vary over a much wider range than those found to hold for 24 hour specimens.

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762 Urine after Muscular Exercise

The variations caused by short periods of strenuous exercise were clean-cut and striking. There was a diminution in the

TABLE VII.

Mar. 24, 1924. Control. Subject L. Ingested water every 20 min., 200 cc. from 1.30, 166 cc. from 3.05 p.m. Figures represent excretion per minute.

Time. VOIUIXN?. PH Acid. NHa Creatinine.

p. m.

2.35 2.45 2.55 3.05 3.15 3.25 3.35 3.45 3.55

7T9 5.8 8.5 5.8 9.8 5.8 9.0 5.8 7.9 5.8 8.3 5.8 7.2 5.8 7.5 5.8 7.6 5.8

cc. 0.1 N cc. 0.1 N mv.

0.649 0.239 1.19 0.675 0.287 1.14 0.530 0.279 1.29 0.526 0.387 1.12 0.490 0.327 0.96 0.439 0.263 1.16 0.477 0.364 1.03 0.447 0.269 0.96 0.435 0.302 1.14

-

TABLE VIII.

May 19, 1924. Control. Subject L. Water every 15 min., 100 cc. at 1.06 p.m. and thereafter until 2.00 p.m., then 50 cc. thereafter. Figures represent excretion per minute.

Time. Volume.

__- p. m. cc.

2.00 4.20 2.10 5.77 2.20 5.80 2.30 4.30 2.40 3.05 2.50 2.25 3.00 1.90 3.10 2.17 3.20 2.13 3.30 2.23 3.40 1.42 3.50 0.90

-

--

-

PH Acid. NH3

6.2 6.2 6.2 6.2 6.1 5.8 5.8 5.8 5.8 5.8 5.6 5.4

-- c. 0.1 N cc. 0.1 N

0.293 0.211 0.285 0.223 0.286 0.207 0.235 0.203 0.229 0.213 0.188 0.181 0.258 0.213 0.258 0.184 0.258 0.203 0.252 0.194 0.258 0.188 0.270 0.181

-

1

.-

-

MaI N. NHrN

PO, ~ Total N

w. %. 0.1 M per cent

10.05 0.123 2.95 8.06 0.157 3.87 8.24 0.154 3.52 7.74 0.134 3.67 7.59 0.136 3.94 6.47 0.129 3.93 7.38 0.151 4.04 7.87 0.144 3.28 7.54 0.136 3.76 7.32 0.138 3.71 6.13 4.29 6.02 0.112 4.21

-

.

-

Cl

:c. 0.1 N

1.31 1.00 0.83 0.66 0.80 0.61 0.63 0.54 0.34 0.37 0.37 0.32

volume of urine excreted which could not be overcome by drink- ing even relatively large amounts of water. In Experiment 12 (TabIe XII) 200 cc. of water were ingested every 30 minutes for a

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Wilson, Long, Thompson, and Thurlow 763

TABLE IX.

Apr. 15, 1924. Strenuous exercise 2 min., 2.05-2.07 p.m. Subject W. Ingested 100 cc. water at 1.50 and at 2.03 p.m. Figures represent excretion per minute.

Time. VI+ ume. PH Acid. NH1

p. m.

1.44 1.54 2.04

cc.

3.45 5.05 4.90

. - e

6.9 6.8 6.6

e. 0.1 N cc. 0.1 N

0.530 0.072 0.402 0.068

0.398 0.061

2.14 3.95 6.0 0.530 0.233 2.24 3.50 5.3 0.795 0.318 2.34 1.10 5.2 0.596 0.260 2.44 1.25 5.2 0.635 0.176 2.54 2.90 5.6 0.424 0.116 3.04 3.30 6.1 0.371 0.086 3.14 4.90 6.4 0.331 0.114 3.24 3.90 6.4 0.291 0.099 3.34 2.60 6.4 0.331 0.074

-

- -

r

-

-

btal N. 1

PO4 Cl

m7. pm cent cc.0.f M x. 0.1 N

9.68 1.04 0.293 1.52 9.66 1.00 0.245 1.43 8.37 1.03 0.248 1.22

7.46 4.33 7.48 5.95 6.56 4.97 7.86 3.13

10.30 1.57 8.16 1.48 9.86 1.61 8.40 1.64 8.38 1.24

0.328 0.467 0.336 0.304 0.235 0.177 0.199 0.152 0.147

-

0.43 0.22 0.45 0.80 1.04 1.07 1.54 1.21 1.43

-

Creati- nine.

ml.

1.19 1.00 1.03

1.06 1.10 1.03 1.08 1.03 0.99 1.07

1.06

TABLE X.

Apr. 7, 1924. Strenuous exercise 2 min., 3.X-3.18 p.m. Subject L. Water ingested every 15 min., 25 cc. until 3.00 p.m. then 40 cc. Figures represent excretion per minute.

Time. Vol- ume.

p. m. cc.

2.45 2.40 2.55 1.95 3.05 1.75 3.15 2.20

3.25 3.35 3.45 3.55 4.05 4.15 4.25 4.36 5.00

1.48 1.40 0.75 0.68 0.55 0.70 0.85 1.35 3.8

-

PH

5.4 5.4 5.4 5.4

5.2 5.0 5.1 5.1 5.1 5.1 5.1 5.2 5.3

:c. 0.1 N

0.397 0.478 0.494 0.476

-_ cc. 0.1 N

0.249 0.294 0.315 0.317

ml.

10.26 9.43 9.61

10.20

0.782 0.564 7.12 0.874 0.658 7.34 0.566 0.406 5.51 0.571 0.361 7.52 0.302 0.216 6.00 0.365 0.302 8.21 0.371 0.290 8.86 0.399 0.348 10.65

-

NH3

-

Ibtal N.

-

_ -

-

NHrN rota1 ii

PO4 Cl

per cm1 cc. 0.1 M :c. 0.1 A

3.48 0.146 1.21 4.38 0.161 1.23 4.59 0.159 1.13 4.35 0.183 1.19

11.08 0.239 12.55 0.382 10.33 0.201

6.73 0.176 5.33 0.090 5.15 0.094 4.59 0.058 4.57 0.048

0.52 0.23 0.38 0.44 0.51 0.75 0.73 0.85

-

i

-

Creati- nine.

mg.

1.05 1.08 1.12 1.14

1.11 1.21 1.08 1.17 0.95 1.22 1.19 1.36

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764 Urine after Muscular Exercise

TABLE XI.

Apr. 28, 1924. Strenuous exercise 1 min., 2.51-2.52 p.m. Subject L. 6 round trips. Water ingested every 15 min., 100 1.15-3.50 Figures cc., p.m. represent excretion per minute. -

Time. Vol- ume.

p. m. cc.

2.20 1.90 2.30 3.10 2.40 4.55 2.50 5.60

3.00 3.10 3.20 3.30 3.40 3.50 4.00 4.10 4.20 4.30

1.90 1.35 0.65 0.60 0.85 2.10 4.70 6.40 6.20

PH

5.6 5.6 5.6 5.8

5.6 5.2 5.2 5.2 5.3 5.6 5.8 5.6 5.8 5.8

-7

Acid NHs

-

‘1 LAal N. VHFN ‘otal N

cc.0.1 N :e. 0.1 N m3.

0.308 0.198 9.03 0.292 0.189 10.14 0.234 0.205 10.03 0.268 0.189 8.76

,er cent

3.08 2.61 2.87 3.01

0.319 0.293 6.16 6.65 0.505 0.384 6.79 7.92 0.345 0.253 6.01 5.91 0.250 0.189 7.03 3.75 0.239 0.190 8.78 3.03 0.213 0.210 10.36 2.84 0.223 0.144 10.47 1.92 0.238 0.181 6.57 3.85 0.231 0.181 7.43 3.41

T- - -

L

-

PO4 Cl Creati- nine.

-

x. 0.1 M c. 0.1 N

0.092 0.94 0.097 0.80 0.094 0.92 0.094 1.00

ml.

1.11 1.12 1.12 1.11

0.101 0.29 0.96 0.203 0.08 1.01 0.140 0.25 0.97 0.078 0.32 0.93 0.065 0.46 1.09 0.058 0.51 1.14 0.060 0.54 1.19 0.077 0.34 1.13 0.062 0.39 1.10

TABLE XII.

May 5, 1924. Strenuous exercise 1 min., 2X-2.51 p.m. 6 round trips. Subject L. Water ingested every 30 min., 200 co., l.O>3.05 p.m. Figures represent excretion per minute.

-

Time. 7 JOlUlll0. PII Acid. NH3 Total N NHS-N Total N

- -- P. m. cc.

2.10 6.80 2.20 7.70 2.30 7.20 2.40 7.00 2.50 6.28

5.8 5.8 5.8 5.8 5.8

x. 0.2 N cc. 0.1 N m7. per cent cc. 0.1 M

0.317 0.318 10.69 4.08 0.191 0.359 0.275 10.44 3.69 0.201 0.291 0.208 9.09 3.20 0.143 0.305 0.225 10.56 2.98 0.145 0.274 0.299 8.86 4.70 0.149

3.00 2.28 5.5 0.285 0.348 6.36 7.70 3.10 1.25 5.1 0.467 0.474 6.00 11.09 3.20 0.85 5.0 0.415 0.270 7.31 5.10 3.30 1.35 5.1 0.285 0.232 9.07 3.57 3.40 2.80 5.3 0.233 0.237 10.48 3.17 3.50 4.08 5.8 0.233 0.218 9.14 3.35 4.00 4.25 5.8 0.207 0.190 8.42 3.16 4.10 2.60 5.8 0.207 0.187 7.59 3.47

-

PO4

0.163 0.247 0.228 0.111 0.094 0.090 0.085

Cl

x. 0.1 N

1.75 1.70 1.25 1.23 1.14

0.36 0.20 0.44 0.51 0.74 0.63 0.63

.- ,

- - -

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Wilson, Long, Thompson, and Thurlow 765

period of 2 hours preceding and extending through the period of exercise. In spite of these large quantities of fluid the urine vol- ume fell to less than one-seventh of that of the fore period. The exercise was carried out for 1 minut,e only on a day when the tem- perature was 19.8%. There was no evidence of perspiration. This together with other experiments performed in cold wet

-i-- 6.41

----_ -- /Y/n

FIG. 1. Experiment 9 (see Table IX). Since the hydrogen ion concentration rises and falls with the other

acidity factors the pH of the urine is plotted inversely in Figs. 1 and 2 to show this relationship more clearly.

weather leads us to believe that the variations observed have little to do with excessive losses of fluid by way of the lungs and skin. The fact that Simpson has found increased urine volumes accom- panying a slight increase in body temperature cannot be used to explain these observations.

After exercise the urine quickly became more acid and there was a rise in excretion of titratable acids and ammonia. No

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766 Urine after Muscular Exercise

matter what the reaction of the urine during the fore period the acidity increased to pH 5.5 to 5.0. In one experiment not reported a pH of 4.8 was observed. The titratable acidity was nearly doubled and the ammonia increased 2 to 5 times. The total nitrogen fell so that the rise in the percentage of ammonia nitrogen to total nitrogen was even greater. The excretion of inorganic phosphates rose while chlorides diminished. There was no signifi- cant change in the excretion of creatinine. All these changes appeared in the urines collected during the first 10 minutes after exercise and reached maxima during the second or third 10 minute

0.2

I p . --. ’ -- /Y/i?* /O to JO &J ~3 60 70 go

Time a/ter exert/se

FIQ. 2. Experiment 10 (see Table X).

period. Some of these variations are shown in Figs. 1 and 2 which illustrate data from Experiments 9 and 10.

The return toward normal proceeded smoothly after the extreme variation was reached. A careful comparison of the data with the variations observed in the control experiments permits a close approximation concerning the time required for the effects of the exercise to disappear.

After I or 2 minutes of strenuous exercise all of the variations persisted for a relatively long time and disappeared only after 30

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Wilson, Long, Thompson, and Thurlow 767

to 90 minutes. The titratable acidity and ammonia usually returned to normal in 30 to 50 minutes, while the pH changed more slowly (50 to 90 minutes). The inorganic phosphate and total nitrogen values were back to normal in the minimum time and then swung beyond, the inorganic phosphate diminishing and the total nitrogen rising beyond the normal levels. Chlorides and urine volumes changed somewhat less rapidly.

DISCUSSION.

The variations observed are in some respects different from those found in studying long periods of strenuous exercise and suggest different explanations. In most of the older experiments the subject exercised as vigorously as possible for many hours. The most extreme variations were usually observed in subjects unac- customed to the strenuous exertion who, therefore, became extremely fatigued and sometimes recovered only after 2 or 3 days of rest. Forms of exercise such as mountain climbing, wood chopping, ditch digging, and bicycle riding for 50 to 80 miles over hills have been resorted to.

Such experiments are recorded in papers by Engelmann (1871), Dunlop, Paton, and coworkers (1897-98), Garratt (1898-99)) and Campbell and Webster (1922) where also may be found many other references to similar experiments. While the reports are not uniform the general conclusion can be drawn that during hours of severe exercise there is a loss of water and sodium chloride in the perspiration which may result in a diminished elimination of these materials for several days. As a result of the exercise the excretion of urea, phosphates, sulfates, titratable acidity, and ammonia rises. These experiments have led to the conclusion that extreme fatigue following long periods of strenuous exercise is associated with increased protein catabolism.

A few investigators have studied urines collected over suffi- ciently short periods so that their data and conclusions may be considered in connection with our studies. Their work may be conveniently discussed by topics.

Volume.-The short experiments of MacKeith and his collabora- tors (1923-24) permit conclusions to be drawn concerning the effect of exercise on the volume of urine. Diuresis was brought about by drinking tea just before the experiments. This diuresis

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768 Urine after Muscular Exercise

was quickly obliterated during rapid running, though the urine volume rose after the exercise was stopped. The loss of water by way of the skin and lungs could not account for the drop in urine volume and the authors suggest that dilatation of the cutaneous vessels and constriction of the splanchnic vessels may, together with other factors, cause the phenomenon. They found that the tense attitude assumed by a runner about to start a race caused a similar drop in urine volume.

Reaction.-Talbert (1919-20) and Endres (1922) report in- creased Cn of the urine after periods of 14 and 15 minutes of

exercise. Our experiments show more extreme variations than they report.

Titratable Acidity.-Titratable acidity was studied in detail by MacKeith and collaborators who found an increased excretion after exercise. Ryffel (1909-10) report,s two experiments showing the same. MacKeith and Ryffel record a few data on the excre- tion of ammonia which show an increased excretion after exercise.

Phosphates.-Embden and Grafe (1921) observed an increased excretion of inorganic phosphates by studying 2 hour urine col- lections after 2$ to 4 hours of strenuous exercise. Blatherwick, Bell, and Hill (1924) have recently observed that less phosphate was excreted by patients when quiet in bed than when normally active.

Chlorides.-Embden and Grafe state in their study on phosphate excretion that the chlorides diminished during the 24 hour period of work but give no figures. They mention that part of the diminution is due to the loss of chlorides in the sweat, but they state that, as higher values were obtained on the following day, they think there is some type of antagonism between phosphate and chloride. The experiments of Riickemann (1922-23) indi- cate that this may be true. The latter investigator found that the feeding of acid sodium phosphate caused a retention of chlorides. MacKeith and coworkers report an increased excre- tion of chlorides after exercise.

Total Nitrogelz.-Embden and Grafe state that no characteris- tic change was observed in the nitrogen excretion. MacKeith and coworkers report low values for total nitrogen during exercise with increases as the urine volumes rose after exercise.

Certain correlations may be made between t.he changes in the

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Wilson, Long, Thompson, and Thurlow 769

composition of the urine and changes occurring in the body. The diminished urine volume after exercise may be due in the main, as MacKeith and coworkers suggest, to diminished blood flow through the kidneys. Alterations in the water distribution between plasma, corpuscles, and tissue cells due to changes in the osmotic relations may also play a role. Scott, Hermann, and Snell (1917) have demonstrated that muscle contractions cause water to pass from the blood into the muscle, resulting in an increased water content of muscle and a more concentrated blood. Loss of water by way of the lungs and skin can hardly account for the variations observed in our short experiments.

The changes in the reaction of the blood after exercise observed by Barr, Himwich, and Green may easily account for the dimin- ished pH and increased ammonia and acid excretion in the urine. The acid and ammonia excretions return to normal in about the same interval of time required for the abnormal reaction of the blood to disappear. The diminution of pH of the urine and the increased excretion of ammonia indicate that the elimination of acid metabolic products was being accomplished with the mini- mum loss of base from the body. The effectiveness of these mechanisms, however, was not great. In Experiment 9 (Table IX) an increased excretion of about 0.8 mM of ammonia resulted from the exercise and about 0.9 m.M of base was made available by eliminating the phosphate in urine with a lower pH. Judging from similar experiments reported in the next paper on the same individual from 5 to 16 mM of lactic acid were excreted combined with base. The body, therefore, lost base.

The same mechanisms undoubtedly remain in force to bring about a retention of base after the large quantities of acid had been excreted. Such an adjustment probably accounts for the unusu- ally low pH values and high ammonia excretion observed in the control experiment of Subject W. (Table VI) carried out 3 hours after an experiment involving two periods of strenuous exercise in which 2.16 gm. of lactic acid were excreted with 24 mM of base.

Embden and Adler’s (1922) experiments on isolated muscles would indicate that the increased excretion of phosphate is due to the liberation of phosphoric acid in the active muscle and its diffusion into the blood stream and thence into the urine. The greatly diminished excretion during the later periods in the experi-

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770 Urine after Muscular Exercise

ments suggests that the recovery after muscular exercise involves the retention and utilization of phosphates, possibly for synthe- sizing the “lactacidogen” of muscle. Embden and Lange’s (1923) experiments also point to a diffusion of chloride into the active muscle. This observation, together with other shifts which prob- ably occur due to change in reaction of the body fluids, may account for the diminished chloride excretion after exercise. It is improbable that appreciable quantities of chlorides were lost by way of the skin. The retention of water by the body necessi- tating a retention of chlorides as well as urea should not be over- looked.

The relationship between the changes in the composition of the urine and the composition of the blood which may be inferred from the correlations mentioned above demonstrates how rapidly and efficiently the kidney responds to assist in the regulation of the neutrality and osmotic pressure in the body. A study of the variations of water, phosphate, and chloride concentrations in blood and tissues may throw further light on these relationships.

SUMMARY.

The effect of short periods of strenuous exercise on the compo- sition of the urine was studied. For this purpose, urine was col- lected in 10 minute periods. There were observed, following exer- cise, a decrease in urine volume, and an increase in hydrogen ion concentration? and acid and ammonia excretion. The elimination of phosphates rose while that of chlorides fell. Ea,ch of these changes reached its maximum within 20 to 30 minutes and was followed by normal values within 40 to 90 minutes after 1 or 2 minutes of exercise.

BIBLIOGRAPHY.

Barr, D. P., and Himwich, H. E., J. Biol. Chem., 1923, a, Iv, 525. Barr, D. P., and Himwich, H. E., J. Biol. Chem., 1923, b, Iv, 539. Barr, D. P., Himwich, H. E., and Green, R. P., J. Biol. Chem., 1923, c, Iv,

495. Bell, R. D., and Doisy, E. A., J. Biol. Chem., 1920, xliv, 55. Blatherwick, N. R., Bell, M., and Hill, E., J. BioZ. Chem., 1924, lxi, 241. Briggs, A. P., J. BioZ. Chem., 1922, liii, 13. Campbell, J. A., and Webster, T. A., Biochem. J., 1922, xvi, 106. Dunlop, J. C., Paton, D. N., Stockman, R., and Maccadam, I., J. Physiol.,

1897-98, xxii, 68.

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Embden, G., and Adler, E., 2. physiol. Chem., 1922, cxviii, 1. Embden, G., and Grafe, E., 2. physiol. Chem., 1921, cxiii, 108. Embden, G., and Lange, H., 2. physiol. Chem., 1923, cxxx, 350. Endres, G., Biochem. Z., 1922, cxxxii, 220. Engelmann, G. J., Arch. Anat. u. Physiol., 1871, 14. Fiske, C. H., J. Biol. Chem., 1921, xlix, 163. Folin, O., and Bell, R. D., J. BioZ. Chem., 1917, xxix, 329. Garratt, G. C., J. Physiol., 1898-99, xxiii, 150. MacKeith, N. W., Pembrey, M. S., Spurrell, W. R., Warner, E. C., and

Westlake, H. J. W. J., Proc. Roy. Sot. London, Series B, 1923-24, xcv, 413. Rockemann, W., Arch. Kinderh., 1922-23, lxxii, 161. Ryffel, J. H., J. Physiol., 190910, xxxix, p. xxix. Scott, F. H., Hermann, E. T., and Snell, A. M., Am. J. Physiol., 1917, xliv,

313. Simpson, G. E., J. BioZ. Chem., 1924, lix, 107. Talbert, G. A., Am. J. Physiol., 191920,1,579. Whitehorn, J. C., J. BioZ. Chem., 1920-21, xiv, 449.

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Thompson and Sylva ThurlowD. Wright Wilson, W. L. Long, H. C.

EXERCISETHE URINE AFTER MUSCULAR

CHANGES IN THE COMPOSITION OF

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