You are on page 1of 9




(From the Division of Animal Nutrition, Urbana, Illinois, and the Department of
Anatomy, University of Illinois, Chicago, Illinois)

(Received for publication, January 19, 1953)

Published analytical data on the chemical composition of the adult, human

Downloaded from by guest on September 9, 2019

body are scant, and statistically inadequate. References to the earlier
literature may be found in a previous communication from this labora-
tory (1) and in a more recent contribution by Widdowson, McCance, and
Spray (2), which are the most complete reports available.
An important application of such data relates to the validity of met,-
abolic balance studies of nitrogen, calcium, iron, etc., as measures of the
daily net requirements of humans for these nutrients. If the results of
such studies, conducted at various stages of growth, are integrated, and
the total amount. of nutrients so calculated to be present in the adult are
in good agreement with analytical data obtained by direct chemical anal-
ysis of the adult,, then the use of metabolic balances for the above purpose
would appear to be a valid technique.

Methods and Materials

The subject was a white male 46 years of age, 53.8 kilos in weight, and
168.5 cm. tall. Other measurements taken were as follows: stem length
(top of head to perineum) by calipers 87.2 cm., shoulder (biacromial)
width 38.4 cm., width between iliac crests 29.3 cm., circumference of upper
chest 78.0 cm.
Death was due to a skull fracture as a result of a fall; a postmortem ex-
amination performed by one of the authors (A. R. C.) confirmed this diag-
nosis and revealed hypostatic congestion of the lungs. No other abnor-
malities were noted.
The use of Edwards’ nomogram (3) leads to the conclusion that this
specimen was 23 per cent below the average weight for his age and height.
His appearance was that of a moderately thin man in good physical con-
After admission of the patient to the hospital in an unconscious state,
medication consisted of penicillin and intravenous dextrose and saline; 4
* The data reported in this paper were secured in the course of an investigation
of the trace element content of human tissues, covered by the Atomic Energy Com-
mission contract No. AT(ll-l)-67, project 7, with the University of Illinois.

hours before death, 5 days after admission, 30 gm. of aminopyrine were

given for elevated temperature.
The body was preserved at 4-5“ for the 1st week after death, and by
freezing at -18” until dissection was started 1 month later. Dehydra-
tion during storage in the frozen state was limited to 0.03 per cent of the
body weight by wrapping the specimen in a plastic sheet.
Dissection of the cadaver to obtain the various organs and tissues de-
sired for study was made under the supervision of two of the authors (A.
R. C. and R. M. F.). Care was taken to minimize and to measure moisture
loss during all the steps of preparation of the samples. Since trace min-
eral elements were to be determined, particular precautions were taken to

Downloaded from by guest on September 9, 2019

prevent contamination. The dissection was performed on a stainless steel
table covered by a polyvinyl sheet, and all the instruments used were of
stainless steel. The organs and tissues, as soon as obtained, were placed
in bags of polyvinyl plastic. When dissection was completed and the sam-
ples were weighed, representative portions of the larger samples and all of
the smaller samples were placed in stainless steel containers with close
fitting covers, and all the samples were autoclaved at 15 pounds pressure
for 45 minutes. Following this, the containers were sealed and the samples
preserved by freezing until prepared for chemical analysis.
Soft tissue samples were prepared for analysis by first being diced in a
porcelain bowl with stainless steel knives. After thorough mixing, a por-
tion was removed for macronutrient analysis and was ground in a power
grinder. The remainder was reserved for subsequent analysis for trace
element content. Representative portions of the skeleton were commi-
nuted as finely as possible with an iron mortar and pestle. The entire left
tibia, ulna, and femur were so treated and analyzed separately.
Moisture was determined on all the samples by drying them to constant
weight in a vacuum oven at 70”. Ether extraction of the dried samples
was accomplished by the Soxhlet principle, a low boiling petroleum ether
being used for 48 hours. Total nitrogen was determined by the Kjeldahl-
Wilfarth-Gunning method with mercury as a catalyst; the samples were di-
gested for 4 hours after clearing. The factor 6.25 was used to convert ni-
trogen data to protein equivalent. Ash was determined with a muffle
furnace at 600”. Calcium and phosphorus were determined on the ash,
calcium in bones by permanganate titration of the oxalate (according to
the method of the Association of Official Agricultural Chemists), calcium in
soft tissues by calorimetry (4), and phosphorus in bone by the standard
phosphomolybdate titrimetric procedure and in soft tissues by the color-
imetric procedure of the Association of Official Agricultural Chemists.
The total weight of the individual tissues and organs was 1.40 kilos less
than the weight of the cadaver when dissection was started. This “loss”
is treated as water in adjusting all the results to the original fresh basis.


The analytical results obtained on the organs and tissues are shown in
Table I, as is the precentage contribution of each chemical subdivision to

Chemical Composition of Adult Human Body
The values are given in per cent.

water Ether
rxtract Ash Ca P

Downloaded from by guest on September 9, 2019

Skin. ................. 6.33 57.71 14.23 27.33 0.62 0.0034 0.070
Skeleton. ............. 17.58 28.17 15.04 19.71 26.62 10.68 4.61
Teeth ................. 0.08 5.00* 23.00* 67.95 25.05 12.09
Striated muscle. ...... 39.76 70.09 6.60 21.94 1.01 0.0066 0.156
Brain, spinal cord,
nerve trunks. ....... 2.99 75.09 12.35 11.50 1.37 0.0147 0.299
Liver ................. 2.34 71.58 3.11 22.24 1.35 0.0133 0.303
Heart. ................ 0.52 62.95 16.58 17.48 0.61 0.0058 0.144
Lung@. ............... 3.30 77.28 1.32 19.20 1.03 0.0090 0.132
Spleen*. .............. 0.11 78.69 1.19 17.81 1.13 0.0089 0.217
Kidneys .............. 0.51 70.58 7.18 19.28 0.87 0.0057 0.188
Pancreas*. ............ 0.14 73.08 13.08 12.69 0.93 0.0143 0.155
Alimentary tract ...... 1.86 77.40 9.17 12.77 0.53 0.0140 0.098
Adipose tissue. ....... 11.37 23.02 71.57 5.85 0.20 0.0078 0.031
Remaining tissue, solid 11.43 59.29 22.47 17.28 0.85 0.0257 0.088
“ “ liq-
uid ................. 0.59 81.45 2.55 13.58 0.82 0.0044 0.104
Bile, content of blad.
der and alimentary
tract ................ 0.99
Hair and nails. ....... 0.10
Average (weighted)
all tissues.. . . 98.91 55.74 19.66 18.82 5.49 1.928 0.936
Total composition
whole body weigh.
ing 53.80 kilos.. . . 100.00 55.13 19.44 18.62 5.43 1.907 0.925
Composition fat-
free body. . . 69.38 23.43 6.83 2.400 1.164

* Chemical composition assumed.

t Congested with blood.

the entire body. The last three horizontal rows of figures show the compo-
sition of the entire body on the various basesindicated. On the basis of
the tissuesanalyzed, or for which analyses were assumed,only 0.29 per cent
is unaccounted for.
Table II shows the macronutrient composition of the three “normal”

specimens for which these data are available. Owing to the variation in
fat content, the data are presented on the fresh and on the fat-free basis.
The specimen analyzed by Mitchell et al. was definitely of higher moisture
and lower fat content than were the other two. In spite of this, the sep-
arable fat in the present specimen amounted to only 11.37 per cent of the
body weight while constituting 13.63 per cent of the weight of the specimen
analyzed by Mitchell et al. The additional fat found in the skeleton
(25.04 per cent versus 17.18 per cent) and skeletal muscle (6.60 per cent
versus 3.35 per cent) of the present specimen largely accounts for the diff-
erence. Unfortunately, Widdowson et al. report no data on these tissues

Downloaded from by guest on September 9, 2019

Chemical Composition of Adult Human Body on Fresh and Fat-Free Bases As Reported
in Literature and from This Study
The values are given in per cent.

This study Mitchell el al. (1) Widdowson el OZ. (2)

Fresh Fat-free Fresh Fat-free Fresh Fat-free

Fat.. ..:. 19.44 12.51 23.6

Moisture. . . 55.13 69.38 67.85 77.55 56.0 73.2
Crude protein. 18.62 23.43 14.39 16.45 14.4 19.2
Ash...................... 5.43 6.83 4.84 5.53 6.0 7.6
Calcium, 1.907 2.40 1.596 1.82 2.0 2.48
Phosphorus. 0.925 1.16 0.771 0.88 1.04 1.29

Ca:P ratio.. 2.07:1 2.07:1 1.92:1

separately. It should also be noted that they determined the water con-
tent by difference, not by analysis.
The following data show the contribution, in per cent, of the striated
muscles to the total body composition; the figures in parentheses are those
obtained by Mitchell et al.: body weight 39.76 (31.56), water 50.54 (38.8),
fat 13.50 (8.1), protein 46.85 (34.6), ash 7.36 (5.6), calcium 0.14 (0.2), phos-
phorus 6.7 (4.5), dry matter 26.51 (19.2). It may be seen that, compared
to the data of Mitchell, the skeletal muscle of the present specimen con-
tributed more than its proportionate share of fat-soluble material to the
total body fat.
Table III shows the composition of the individual bones as well as of
the entire skeleton on various bases. The percentage contribution of the
skeleton to the total body composition, with comparative data from Mitchell
et al. in parentheses, is shown as follows: body weight 17.58 (14.84), water
8.98 (9.0), fat 22.64 (19.5), protein 18.62 (18.6), ash 87.27 (85.7), calcium

99.58 (99.0), phosphorus 88.65 (90.0), dry matter 28.15 (30.1). These data,
as well as those for the individual bones, are strikingly similar. The most
significant difference is in the calcium to phosphorus ratio of the rib; in

Chemical Composition of Bones
The values are given in per cent.

Downloaded from by guest on September 9, 2019

Moisture. ........ 11.92 15.76
Ether extract ..... 49.12 50.09 19.91 23.63
Crude protein. ... 15.56 17.67 35.40 22.25 26.41 34.58
Ash. ............. 30.47 34.59 69.30 42.22 50.12 65.63
Calcium .......... 11.44 12.99 26.03 37.56 15.04 17.85 23.37 35.61
Phosphorus. ...... 5.04 5.72 11.46 16.54 6.54 7.76 10.16 15.48
Ca:P ........... 2.27:1 2.30:1

10th rib Total skeleton

Moisture. ........ 26.32 28.17

Ether extract ..... 7.88 10.69 25.04 34.86
Crude protein. ... 29.00 39.36 44.07 19.71 27.45 42.14
Ash. ............. 34.28 46.53 52.10 26.62 37.06 56.89
Calcium .......... ‘12.68 17.21 19.27 36.99 10.68 14.86 22.81 40.09
Phosphorus. ...... 5.72 7.76 8.69 16.68 4.61 6.42 9.86 17.33
Ca:P ........... 2.22:1 2.31:1

Body water Fat Protein Ash Ca P Dry

weight matter

Contribution of
skeleton to total
composition of
body, 17.58 8.98 22.64 18.62 87.27 99.58 88.65 28.15
Contribution of
striated muscle
to composition
of body. 39.76 50.54 13.50 46.85 7.36 0.14 6.70 26.51

this investigation the tenth rib had a Ca:P ratio of 2.22’:1, while in that of
Mitchell et al. the ninth rib was calculated to have a Ca:P ratio of 2.35:1.
The ash of the latter bone contained 38.97 per cent of calcium and 16.60 per
cent of phosphorus, while in the specimen herein reported the figures were
36.99 and 16.68 per cent, respectively.

These data support the thesis that analysis of a single bone is not likely
to give a true estimate of the composition of the entire skeleton of an ani-
mal. This matter and a discussion of bone analyses performed by other in-
vestigators are treated more t.horoughly by Mitchell et al. (1) and by Follis
Other measurements made on the tibia, ulna, and rib, respectively, are
as follows: maximal length 38.5; 27.2, 24.1 cm.; minimal diameter 2.15,
1.15, 0.40 cm.; density by water displacement (determined on the fresh,
intact bone) 1.25, 1.30 (not determined on the rib).
It is worthy of note that the calcium to phosphorus ratio of the entire
body is identical with that obtained by Mitchell et al. and that Widdow-

Downloaded from by guest on September 9, 2019

son’s data also support the finding of a ratio approximating 2: 1. It would
appear that this ratio is more representative of that found in healthy adult
humans than is the ratio of 1.73 : 1 suggested by Shohl (6) as a result of
estimating that the human body contains 1.66 per cent calcium and 0.96
per cent phosphorus. The data reported herein and by Widdowson et al.
(2) indicate that total calcium may amount to approximately 2.0 per cent
of the body weight.
The higher water content of the specimen analyzed by Mitchell et al.
persists when the data are presented on the fat-free basis (Table II).
Analysis of individual tissues, expressed on the fat-free basis, shows that
skin, striated muscle, adipose tissue, and solid remaining tissues contribute
the major portion of the difference in moisture content. The skeleton and
nervous system do not differ materially in water content between the two
specimens. These data are shown in Table IV.
Considerable interest has been evidenced in recent years in determina-
tion of the total water and fat content of the living body by indirect meas-
urements. The validity of such methods has been well established. Kray-
bill et al. (7) compared antipyrine, specific gravity, and direct analysis
methods, employing cattle as experimental subjects; Messinger and Steele
(8) obtained favorable comparisons between the antipyrine and specific
gravity methods, employing humans as subjects of investigation; Rathbun
and Pace (9) compared specific gravity and direct analytical methods with
guinea pigs. Other investigators who have employed antipyrine (8, 10,
1 l), specific gravity (8, 12), or deuterium oxide (13) methods on normal
human males have found water content ranging from 43 to 73 per cent on
a total body basis, and 66 to 79 per cent on the fat-free basis.
A comparison of these data with those given in Table II shows that the
cadavers analyzed directly all have water content within the range of values
reported by indirect procedures. This would seem to contradict the state-
ment of Edelman et al. (13) that the specimen analyzed by Mitchell et al.
was “grossly edematous.”

The concept of constancy of composition of lean body tissues, well ex-

pressed by Murray (14), has been verified and the normal limits of varia-
tion have been more firmly established in relation to water content in
recent years. Further evidence is given by Pace and Rathbun (15), who
reported from their own work and that of others that, for the rat, guinea
pig, rabbit, cat, dog, and monkey, the mean water content of fat-free tissue
was 73.2 per cent with a range of 69.9 to 76.3. Also, Kraybill et al. (16)
found in cattle a mean value of 72.6 per cent water in lean body mass,
with a range of 69.3 to 75.7.

Downloaded from by guest on September 9, 2019

Water Content of Selected Tissue Expressed on Fat-Free Basis, As Found in This
Study and by Mitchell et Al.

Tissue This study Mitchell et al.

per cent per cent

Skin. ................. 67.29 74.34
Skeleton. ............. .. 37.58 38.41
Striated muscle ........ .. 75.04 82.28
Nervous system. ...... 85.67 83.98
Adipose tissue. ........ 80.94 87.02
Remaining solid tissue. 76.47 80.36


The chemical composition of the body of a normal adult human, 46

years of age, is reported. The whole body contained 19.44 per cent ether
extract, 55.13 per cent moisture, 18.62 per cent protein (N X 6.25), 5.43
per cent ash, 1.907 per cent calcium, and 0.925 per cent phosphorus. On
the fat-free basis these data are, respectively, 69.38, 23.43, 6.83, 2.40, and
1.16. Analyses are reported on thirteen separate tissues and organs and
are summed to give the above data. Comparisons are made with similar
data reported previously.
Evidence is presented that the normal adult human body contains ap-
proximately 1.8 to 2.5 per cent calcium on the fat-free basis and possesses
a Ca to P ratio of 2:l.
The water and fat contents of the adult human body as determined di-
rectly are in good agreement with the data obtained by indirect methods.


1. Mitchell, H. H., Hamilton, T. S., Steggerda, F. R., and Bean, H. W., J. Biol.
Chem., 168, 625 (1945).
2. Widdowson, E. M., McCance, R. A., and Spray, C. M., Clin. SC., 10, 113 (1951).
3. Edwards, T. I., Am. J. Hyg., 36, 307 (1942).

4. De Loureiro, J. A., and Janz, G. J., Biochem. J., 38, 16 (1944).

5. Follis, R. H., Jr., J. Biol. Chem., 194, 223 (1952).
6. Shohl, A. T., Mineral metabolism, New York (1939).
7. Kraybill, H. R., Hankins, 0. G., and Bitter, A. L., J. Appl. Physiol., 3,681 (1951).
8. Messinger, W. J., and Steele, J. M., Proc. Sot. Exp. Biol. and Med., 70,316 (1949).
9. Rathbun, E. N., and Pace, N., J. Biol. Chem., 168,667 (1945).
10. Steele, J. M., Berger, E. Y., Dunning, M. F., and Brodie, B. B., Am. J. Physiol.,
162, 313 (1950).
11. Osserman, E. F., Pitts, G. C., Welham, W. C., and Behnke, A. R., J. Appl.
Physiol., 2, 633 (1950).
12. Broiek, J., Federation Proc., 11, 784 (1952).
13. Edelman, I. S., Haley, H. B., Schloerb, P. R., Sheldon, D. B., Friis-Hansen,
B. J., Stoll, G., and Moore, F. D., Surg., Gynec. and Obst., 96, 1 (1952).

Downloaded from by guest on September 9, 2019

14. Murray, J. A., J. Agr. SC., 12, 103 (1922).
15. Pace, N., and Rathbun, E. N., J. Biol. Chem., 168,685 (1945).
16. Kraybill, H. R. Bitter, A. L., and Hankins, 0. G., J. Appl. Physiol., 4,575 (1952).
R. M. Forbes, A. R. Cooper and H. H. Mitchell
J. Biol. Chem. 1953, 203:359-366.

Access the most updated version of this article at

Downloaded from by guest on September 9, 2019

• When this article is cited
• When a correction for this article is posted

Click here to choose from all of JBC's e-mail


This article cites 0 references, 0 of which can be

accessed free at