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Body weight Its relation to tissue composition segment distribution and motor function II.

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Body Weight: Its Relation to Tissue Composition,
Segment Distribution, and Motor Function
11. DEVELOPMENT OF MACACA MULATTA 1
THEODORE I GRAND
Oregon Regonal Przmate Research Center, Beaverton, Oregon 97005
KEY WORDS Anatomy . Development
composition Limb segments . Macaca
. Musculature . Tissue
ABSTRACT
The relative composition of skin, muscle, and bone and their distribution patterns throughout the body are given for a series of Macaca mulatta
from 141 days conceptual age through adulthood. In terms of percent of total
body weight, the musculature of these animals doubles during the first postnatal
year whereas bone and skin decrease. Regionally, the muscles of the thighs, back
extensors, truncal-forelimb and upper arms increase most markedly. The thighs
double and the upper arms increase whereas the trunk, hands, feet, and tail decrease. The biomechanical implications of these changes for motor development
are discussed.
Body size is a complex variable. The component tissues vary considerably among the
adults of many species (Grand, '77). As a result, the notion of size is frequently oversimplified. Weight increases in domestic animals and man from birth to maturity are
accompanied by proportional changes in tissues and segments and by a major shift in
motor repertoire. In order to examine these
phenomena, I applied a method of dissection
similar to that used for interspecific comparisons to a series of Macaca mulatta. The
baseline for adult Macaca has been established from data in the preceding paper and
specimens of this species were available
throughout all stages of the life cycle. The
same questions could be raised as in the study
of adults: differences in tissue composition,
limb segment weight, tissue composition of
the segments and motor pattern differences of
fetus and adult.
Not surprisingly, this approach reconsiders
many data on body composition gathered by
physiologists (Keys and Brozek, '53; Beatty e t
al., '67; Young, '701, veterinary anatomists
(Palsson, '55; Fowler, '68; Butterfield and
Berg, '66; Butterfield and Johnson, '68),
human anatomists (Wilmer, '40), and other
physical anthropologists (Brozek, '55, '61, '63;
Brozek and Keys, '50; Parizkova, '68; Trotter
AM. J. PHYS. ANTHROP., 47: 241-248.
and Hixon, '74; Trotter et al., '75). The study
has blindspots because of its functionallocomotor orientation. Graphic and functional
extremes are emphasized. The usual statistical measures of growth and careful group
selection could not be applied; curves of
growth were not matched. Nevertheless, prenatal rhesus macaques appear to be a homogeneous population in their structural characteristics and resemble prenatal macaques of
other species (Grand, unpublished data).
METHODOLOGY
In table lA, the conceptual age, body
weight, sex and anatomical procedure used
are listed for the prenatal specimens (from
141 to 155 days). Age at birth averages 166167 days (conceptual age). Postnatal animals
are included in table 1B. Nine adult animals
were dissected for limb segment analysis
only; the specimens are not included in table
1, but the data are given in table 3.
The dissection procedure was reported in
the earlier paper of this study (Grand, '77).
Greater fluid loss occurred in the fetal forms
than in postnatal and adult specimens, but
this could not be corrected.
lPublication No. 827 of t h e Oregun Regiunal Primate Research
Cenler, supported in part by Grants RR-00163 from the National
Institutes of Health. USPHS
241
242
THEODORE 1. GRAND
TABLE 1
Spcimens of Macuca mulutta
A. Specimens from 146 to 155 duys (conceptual age:
B. Specunens to adulthood
A
CONCEPTUAL
IDENT. No.
PRENATAL
BPDSTNATAL
SEX
AGE
BDDY WEIGHT
(KO)
PRESERVATION
7030
t
141 ~ A Y I
,340
FRESH
6830
6117
6127
F
F
146
DAYS
I
FRESH
149
DAYS
N
1%DAYS
6963
7014
7018
6739
7045
6676
R
150
DKiS
)I
152
DAYS
A
F
153
DAYS
M
1% DRYS
155 DAYS
502
,328
372
.467
.447
,w 7
,2675
,371
,966
153 DAYS
M
w54
F
6393
1
6341;
5053
5545
M
fl
r:
AD I
1376
M
AD,
=Muscle
I B o n e
13 MONTHS
1.7
1.8
FRESH
FROZEh
I
YEARS
YEARS
AD.
FRESH
FRESH
FRESH
FRESH
FROZEk
FRESH
1.946
FRESH
2,500
2.700
5.500
9,500
11.75
FRESH
FRESH
FRESh
FRESH
FRESH
=Skin
Prenatal Averages
Skin: 15.2 To i 1.86
Muscle: 24.6X f 2.66
Bone: 19.9 f 0.69
7030
6830
61 27
6963
7014
7018
6739
7045
6676
I
Postnatal Average
Skin: 12.6% i3.45
Muscle 43.4 t 4 0
Bone: 14 4 X 2.9
6454
6393
6344
5053
5545
*
I378
,2004
4835
'4993
I0
20
Ja
4.0
Body Weight
io
$0
Fig. 1 Bar graph-tissue composition of whole body.
RESULTS
A. Tissue composition in relation to
body weight
The weights of skin, muscle, and bone and
their percentage relations to body weight are
given for each specimen in table 2. The bar
70
so%
*, specimens reported in Grand ('771
mauh (fie. 1) represents numerically the bilateral cinfigura-tion for each animal. Calculation of mean and standard deviation shows
that the promrtions of fetal tissue vary less
and differ- significantly from adult tissue proportions.
The skin, which in prenatal animals repreV
I
243
ANALYSIS OF BODY WEIGHT: DEVELOPMENT OF MACACA
TABLE 2
%sue composition ofwhole body. Su: mdiulduals of the bottom row are postnatal animals
#7030
Tlrruf Y ~ , l G f i l %
16.4
55.7
S%W
TBW
5.7
imt
8.1
HIND
TRUHn
2;
BOO" WT.
(KG)
BliCX
"T*EI
WT.(GN)
i. Taw
%7
9.8
447
10.2
54,2
20.7
i6Y.5
748.2
19.4
58.6
73.5
WT.IGM) 2 TBk
220
1 1 8 s ~h'r.lGM)
~
21.3
69.6'
7R
o e
2::
193.7
266,5
68.8
101.9
?:
14.7
7.7
2a1
.267
447
116963
P6i27
TISSUE
4.G
8.9
187
117.0
166.8
u1.u
52.7
FORE
HIND
116830
risPUE
9.5
44.3
20.7
23.4
---
---
466
371
194.3
587.9
138 2
5 5
10.4
m4
737.1
793.2
2z,7
313.2
1OOR ci
:2411.6
219
463.6
u87 7
263
Bow
26,3
14,3
175.0
FORE
HINO
TRUNI
BODY WT,
40.2
152
25
(KG)
*, bilateral weight;
56.1
---
li,
38.0
56,5
244,o
2.7
77.5
160
104.5
---
55
lOk,k
123.8
548.0
95
165.0
198.9
10s
1175
low eshmate. -, weight not taken.
sents about 15%TBW, drops to 12.6%TBW in
the postnatal series. The postnatal decrease is
even greater (to 11.0%TBW) if one unusually
fat animal (#4835) is removed.
The prenatal musculature yields a mean of
about 24.5% TBW; that of the postnatal animals is over 43.0%TBW. The regional changes
are discussed below.
The mean for the prenatal skeleton is
almost 20% TBW, with narrow variability; in
the postnatal animals, this drops below 14.5%
TBW. Bone weight is more variable after
birth, a fact which concurs with data obtained
from other species (author's unpublished observations).
B. Limb segments in relation to body weight
In the prenatal specimens (table 3A1, the
hindlimb segments constitute 15.8% of body
weight, the forelimbs 11%TBW. The tail averages 1.1%
TBW and the trunk and head the remaining 72.1%TBW.
The postnatal forms (table 3B) were divided
into two groups: (1)animals weighing up to 3
kg, (2) animals weighing over 5 kg. The relative weights of segments were averaged for
each of the groups. The lighter, younger animals diverge from the prenatal series. This
shift is continued and "completed developmentally" in the fully adult animals. In the
heaviest animals, the hindlimbs average 23%
TBW, the forelimbs 12.6% TBW. The tail
drops in relative weight, so that the head and
trunk now constitute 63.7%TBW.
A detailed analysis of two representative
animals in figure 2 illustrates the complexity
of weight distribution and of regional tissue
changes. The profiles are similar to those
obtained for other prenatal and postnatal
specimens.
1. Prenatal M mulatta
In a 150-day fetus (#6127) (fig. ZA), each
hindlimb represents about 8% TBW, each
244
THEODORE 1. GRAND
TABLE 3
Maraca mulatta-segmentation analysis and averages
A. Prenalal
B. Postnatal-up to three kg
C. Postnatal-over five kg
A
C!%
SiDXl
SEGMENT WT,
IGRI
SEGniui
6117
S E ~ E NW T
i,
Wi,
(LM
!?I)
P
L
5,7
11.1
12.3
...
...
.~.
1.9
10.J
25.5
25.6
18.2
31.0
54.6
nn~,,.
FORE*PII
UWER A R V
FOOT
CALF
r ~:GH
TAIL
l c ~ vWT, (KG)
10.0
24.0
27.1
18.2
30.4
53.5
11.1
‘I , ’:
6.5
‘I , 3
6.8
7.1
7.3
6.’:
7.3
12.0”
6.1
-----
3,4
3.0
8.7
8.2
6.9
8.6
L.6
15,/
32
4.b
2.9
L.li
25,?
67 R
86 6
7l.i
K 6 2
SiGMEVT
41.4
93,l
3 2 ,0
26.6
91.3
:ae.o
553.0
52.7
...
7.5
8.1
nr
L
50.2
253 2
391.9
382,t
471
257 6
YY.5
K76
S ~ i n ~ tUT,
lr
(GY:
AVERAGE
1 T?il
6.0
10.7
10.4
9.5
]‘.I
19.7
5 4
24.2
61.1
85.0
44.1
86.1
194,7
2..4
46.5
190.3
282.4
81.9
212.0
b83.2
59.3
8.27
57.2
223 8
267.7
86.0
232.5
619.5
87.0
9.5
~1378
(69)
886.5
43,3
141 ?
715 r
97 1
240 2
569 3
214,6
R
51.0
262.4
261.0
45 1
6 ‘ 4
199, L
7 c
6.13
23 7
37.5
52.5
161.3
52.0
235.0
87.3
193.F
483 7
73.0
24,s
67 1
83.:
183.b
19.4
:&l;
50
14 2
3 4
5.3
60.2
YilL5
S~GnrhlrU T ,
5.2
8..
8.9
7,:
5.3
20.2
131,u
175,b
552.3
5.8
4.8
14.0
33.0
139,m
410.8
26.11
pjp9
SEGMENTW T ,
(LM)
59.0
70.2
34.6
70.2
16.7
28.0
53.9
128.3
212. 0
66.5
55
L
18.9
55.2
68.’:
34.4
68 3
163.2 1’19.2
17.0
15,s
41.7
53.5
34.7
125.4
185.9
65.3
164, :
Q
Sl27
SLBIIENT
Wi,
!GM)
918 3
88.7
11.35
SElElENT
ki.
lBW
:GK
69.6
0.6
304.7
24
33
1.1
27
7.7
0.7
333 3
124.5
323 1
915 1
78.5
I:, 75
, weight not taken, *, low estimate
forelimb about 6% TBW. Almost 72% TBW is
concentrated in head and trunk. The skeleton
is 11%TBW; the viscera, heart, and brain,
40%; the truncal muscular components, 15%;
and the back extensor muscles 2.6%.
Bone increases and muscle decreases distally in both hindlimb and forelimb. In the
hindlimb, bone is over 18%of thigh weight,
and 43%of foot weight. Muscle is 48%of thigh
weight, but 14.5%in the foot (table 4).
2. Postnatal M mulatta
In the adult (#5053) in figure 2B, the hindlimbs constitute almost 25% TBW; the thigh
“increases” to 8%TBW and the foot “drops” t o
1.2% TBW. The tail “drops” to 0.5% TBW.
Muscle increases considerably in each segment, but both bone and skin are relatively
reduced. The musculature in the thigh constitutes 80% of its weight. Muscle increases in
the calf and foot (table 4). A similar shift occurs in the forelimb. Two changes should be
noted: first, the upper arm segment increases
in percent of total body weight, whereas the
hand decreases; second, the muscular proportions in each segment rise while bone and skin
are usually lowered (table 4).
The adult trunk and the viscera relatively
245
ANALYSIS OF BODY WEIGHT: DEVELOPMENT OF MACACA
X TBW
X TBW
2 2 0 Slun
37
I
Muscle
0Bane
23
23
19
13
A
6'27
B
5053
I
58
L
l50doys
TAIL HINDLIMBS
051 2 4 8
I
TRUNK
62 I
FORE-
100% TBW
Adult
Fig. 2
Segment figures-segments and tissue composition
A. #6127-150 days (conceptual age).
B. #5053--adult male
decrease by 10% TBW (fig. 31, and the skin
and truncal skeleton decrease somewhat.
Each group of muscles (particularly the truncal-forelimb and back extensors) increases;
the back extensors, for example, constitute 5%
TBW. Thus, in adulthood the entire trunk is
reorganized in relative weight, and musculature is the dominant component.
C. Segment variability
The variability of the prenatal segments
and tissue components is somewhat narrow
(fig. 3). Hand, foot, and thigh are significantly
different from those of the postnatal sample.
As expected, the segments of younger animals
(1-3kg) are intermediate between those of the
prenatal and adult forms; with increased body
weight, the foot and hand decrease and the
thigh doubles in percent TBW. The growth
curves are clearest here. The calf increases
and the tail decreases in percent of weight,
but both segments are more variable than
those previously mentioned.
TABLE 4
T u su e
composition
of
segments of #6127 and #5053
_._
..3
'l.4
1.6
159
53.7
21.9
1.3
14.9
552
21.8
13.7
22.2
10.4
70.7
17.7
31.2
16.7
41.7
12.C
Y.1
13s
34.6
282
37.5
45.5
103
80.6
'1.8
1.1
1 5
0,8
7 0
3.81
65
2.5
I.!
'1.0
23
2.c
1.0
3,o
1
L1.l
13i 5
21, 7
8R.6
21.9
481
18.2
565,'
lZ4
465
26.7
112 0
29.0
14.5
43.5
122
74.0
13.2
41 1
8.9
I! 8
9.6
68.3
21.6
55 4
10 j
19 1
25 3
29.9
29.2
38.7
246
THEODORE I. GRAND
90:
. ... .. .
.
..
50
..
. .
40
1
u
-
.. .. .
:I
22
L
2oL2
I
S
3
4
5
6
7’
B’
9
’ 10
!
I1
’
I2
’
Ekdy Weight [kgs)
1
2
3
4
5
8
7
8
9
1
0
112
Body Weight ( k g s l
Fig 3 Segment variation compared w i t h body weight
DISCUSSION
At any stage of life, bone, muscle, and skin
together constitute 60 to 70% of body weight;
and from birth to maturity, these tissues are
responsible for the motor repertoire. (Paradoxically, the remaining 30 to 40% [viscera,
heart, brain, larger glands1 have received far
greater attention. See Spector, ’56 for a
review.) The prenatal form undergoes dramatic segmental redistribution of weight as
well as alteration of tissue and organ ratios to
achieve adult form and composition.
To examine the fate of the “locomotor tissues and segments,” I framed certain questions that had been applied to the adults of
various species: How does the composition of
these tissues change from prenatal through
postnatal development? How are the tissues
distributed to the limb segments and how does
the distribution change? How are these
changes related to locomotor demands?
The tissues in the prenatal macaque are
almost equal in weight (skin 15%,bone 20%
muscle 24%TBW), but during the first year
they begin to diverge considerably (skin 12%,
bone 15%, muscle 43%). Although time of
maturity varied, Palsson (’55) and Fowler
(’68) found these shifts in various birds and
mammals, and Wilmer (‘40) found a similar
shift in man. In rhesus macaques, the decrease in the relative weight of skin can be explained in two ways: (1)the skin is a protective covering which does not need to keep pace
with an increasingly independent motor life;
(2) volume and weight increase by the cube
whereas surface area and skin covering increase only by a square (Thompson, ’69).
Muscular development is most important
because the motor patterns of early postnatal
life change with locomotor independence. The
survival mode of the newborn is clinging to
the mot her; therefore, truncal coordination
and fore- and hindlimb propulsive thrust are
minimized. The hands and feet are heavy; the
thigh, upper arm, and back are low in muscle.
ANALYSIS OF BODY WEIGHT: DEVELOPMENT OF MACACA
Increasing maturity, however, is directly associated with muscle and with whole body activity. Muscle develops in thigh, back extensor and upper arm. As a result, the thigh and
upper arm segments increase relatively, the
hands and feet decrease.
The tissue shift brings about two other phenomena. First, the hands and feet become relatively lighter, forearms and calves become
slightly heavier, thighs and upper arm segments much heavier. The centers of mass of
fore- and hindlimbs are elevated. Recovery
motions may require less effort in the older
animals. The limbs, however, increase in
absolute weight. These weight increases go
directly into limb musculature, and the trunk
becomes relatively lighter.
A second developmental process is change
of density. Although the water content of the
body drops, this is complicated by tissue and
organ changes. The viscera, less dense than
the musculoskeletal apparatus, become lighter. The limb bones become more dense and
muscle volume increases. The precision of
other methods (ash weight or calcium content
of bone, dry weight of muscle) is required to
trace these changes (Mitchell e t al., '45;
Rathbun and Pace, '45; Morales et al., '45;
Pace and Rathbun, '45; Brody, '45). An apparent difference between the results of this
study and those of Trotter and Hexon ('74)
and Trotter et al. ('75) is traced to procedure.
This study demonstrates that the wet weight
of bone drops in relative % TBW; the other
studies show that the dry, fat-free skeleton increases in % TBW up t o six years. I am
describing living tissues and live weight
whereas they are characterizing changes in
body chemistry and tissue density.
Nevertheless, the functional framework
presented here is another approach to body
decomposition or somatolysis (Brozek, '63),
which reinforces earlier findings that tissue
features distinguish newborn from adult, and
that the components of body weight must be
separated a t different stages of development.
Further, it demonstrates that selection operates upon the components of size and upon
survival skills of each stage of life: the motor
requirements of the clinging newborn contrast to those of the independent juvenile and
adult, in whom climbing skill, acceleration,
and endurance are enhanced by truncal coordination and integration of forelimb and
hindlimb motions. The ratios of skin, muscle,
and bone and their distribution throughout
247
the body illustrate how the adult form is the
resultant of many selective forces.
ACKNOWLEDGMENTS
This study has extended over several years
and many people have cooperated with me:
Alan Rogers of the University of Oregon Medical School and Paul Miller of the Oregon
Regional Primate Research Center have obtained my nonprimate specimens (mixed
breed dogs, racing greyhounds, cats, jack rabbits); Nicholas Smythe (Smithsonian Tropical
Research Institute), Gary Dawson, and John
Eisenberg (National Zoological Park) obtained my South American specimens;
Margaret Rarss, Adrienne Zihlman, Patricia
Ladd, Robert Kuehn, and William Montagna
have reread critically several drafts of each
section of the paper; Rose Mary Bocek and
Clarissa Beatty have read over and commented upon this paper; discussions with
William Hunter and Charles Wood have been
important. The ORPRC staff was helpful
throughout research and preparation, particularly Steve Estvanik and Ingrid Palm for
mathematical and statistical evaluation, and
Joel Ito and Kathleen Kerr for the evolution
of the tables and figures. Sheila Alderman,
Joyce Hodges, Barbara Renstrom, and Karen
Anderson brought order t o the remaining
chaos with their typing and retyping of successive stages of the entire manuscript and its
tables.
LITERATURE CITED
Beatty, C. H.. G. M. Basinger and R. M. Bocek 1967
Differentiation of red and white fibers in muscle from
fetal. neonatal, and infant rhesus monkeys. J .
Histochem. Cytochern., 15: 93-103.
Brody, S. 1945 Bioenergetics and Growth. Rcinhold
Publishing Company. New York, 1023 pp.
Brozek, J. 1955 Role of anthropology in the study of
body composition: toward a synthesis of methods. In:
Dynamic Anthropometry. Roy Waldo Miner, ed. Ann.
N. Y. Acad. Sci., 63: 491-504.
1961 Body composition. Science, 134: 920-930.
1963 Quantitative description of body composition: physical anthropology's "fourth' dimension. Current Anthropology, 4: 3-39.
Brozek, J., and A. Keys 1950 Limitations of the "normal"
body weight a s acriterion of normality. Science, 112; 788.
Butterfield, R. M., and R. T. Berg 1966 A classification of'
bovine muscles based on their relative growth patterns.
Res. Vet. Sci., 7: 326-332.
Butterfield, R. M., and E. R. Johnson 1968 The effect of
growth rate of muscle in cattle on conformation a s
influenced by muscle-weight distribution. In: Growth
and Development of Mammals. G. A. Lodge and G. E.
Lamming, eds. Butterworths, London, pp. 212-223.
Fowler, V. R. 1968 Body development and some problems of its evaluation. In: Growth and Development of
Mammals. G. A. Lodge and G. E. Lamming, eds. Butterworths, London, pp. 195-211.
248
THEODORE I. GRAND
Grand, T. I. 1977 Body weight: its relation to tissue
composition, segment distribution, and motor function. I.
Interspecific comparisons. Am. J. Phys. Anthrop., 47:211240.
Keys, A,, and J. Brozek 1953 Body fat in adult man.
Physiol. Rev.. 33: 245-325.
Mitchell, H. H.. T. S. Hamilton. F. R. Steggerda and 11. W.
Bean 1945 The chemical composition of the adult human
body and its bearing on the biochemistry of growth. J .
Bid. Chem., 158: 625-637.
Morales, M. F., E. N. Rathbun. R. E. Smith and N. Pace
1Y45 Studies on body composition 11. Theoretical considerations regarding the major body tissue components.
with suggestions for application to man. J. Biol. (:hem..
158: 677.684.
Pace. N.. and E. N. Rathbun 1945 Studies on body cumpusition. 111. The body water and chemically combined
nitrogen content in relation to fat content. J. Biul.
Chem., 158: 685-691.
Palsson. H. 1955 Conformation and body composition. In:
Progress in the Physiology of Farm Animals. Vol. 2. John
Hamnrond. ed. Butterworths, London. pp. 430-542.
Parizkovd, J . 1968 Body composition and physical fitness.
Current Anthropology, 9: 273-287.
Rathbun, E. N.. and N. Pace 1945 Studies on body composition. I. The determination of total body fat by means of
the body specific gravity. J. Biol. Chem.. 158: 667-676.
Spector. €1. S., ed. 1956 Handbook of Biological Data. W. B.
Saunders and Co.. Philadelphia.
Thompson, L). W. 1969 On Growth and Form. (Abridged by
John Tyler Bonner). Cambridge Univ. Press, Cambridge,
346 pp.
Trotter, M., and B. B. Hixon 1974 Sequential changes in
weight, density and percentage ash weight of human
skeletons from a n early fetal period through old age.
Anat. Rec., 179: 1-18.
Trotter, M., B. B. Hixun and S. S. Deaton 1975 Sequential
changes in weight of the skeleton and in length of long
limb bones of Macaca mulatta. Am. J. Phys. Anthrop., 43:
79-94.
Wilmer. H. A. 1940 Changes in structural components
of human body from six lunar months to maturity. Proc.
SOC.Exp. Biol. Med., 43: 545-547.
Young, V. R. 1970 The role of skeletal and cardiac muscle in the regulation of protein metabolism. In: Mammalian Protein Metabolism. Vol. IV. €1. V. Monro, ed.
Academic Press, New York, pp. 585-674.
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