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The relative weight of the testes in primates.

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THE RELATIVE V~EIGHTOF THE TESTES
I N PRIhIATES
$DOLF’II 11. SCIIULTZ
L a b o r a t o r y of Physical A n t h r o p o l o g y , Johns I I o p k i ~ i s Mrdicnl School,
Baltimore, X u r y l a n d
The relative size of the testes varies very strikingly among
different primates. I n order to gain some accurate information on this subject the author had collected some relevant
data at differelit occasioiis during the past few years. Last
year it became possible to increase this series of data to an
extent resulting in definite conclusioiis. Thj opportunity
arose during the Asiatic Primate Expedition of 1937 of which
the author was a member.l Fiftyfive of the adult male
primates, recorded in thc accompanying table, were weighed
and autopsied on this expedition. Observations on seventeen
other monkeys and apes (mostly from the Hoplziiis comparative-anatomical collection) hare been added, as also data on
three Negroes (unclaimed bodies of the anatomy department
of the Johns IIopkiiis University). Corresponding data from
the literature could be found only in a paper by Hrdli6ka ( ’as),
who has listed the weights of the bodies and of the testes in
seven adult howling monkeys from the field notes of the late
Xr. J. L. Baer. The preseiit paper records the relative weight
of the testes in a total of eighty-tvo simian primates. The
testes alone, without the epididymides, are considered herc.
The first twelve primates listed in the table are platyrrhines
o r New World monlreys. All the others are catarrliines.
This expedition was orgnihcrl by Mr. H. J. Coolidge, Jr. The (larnegie
Tnstitutioii o f Washiiigton generously eoiitributed a grant toward the author’s
expenses 011 this expedition.
387
Weights of body and of tca-tes n.ad percentage relatlon hetween. tlse t w o Wt?fghtS
PIL crglitu adult and two juvenile pitinates
smcim
Ocdiponiidas geoffr0.i
Aotus zonalis
Alouatta palliata
Alouatta palliata
Alouatta palliata I
Alouatta palliata
Alouatta palliata
Aloiiatta palliata *
Alouatta palliata
Alouatta palliata
Average of 8 Alounttn
Lagothrix lagothricha
Ateles geoffropi
Macaea irus
Macaca irus
Macaca irus
Macaca h i s
Macnca irus
Macaca inis
Macaca irus
Macaca irus
Macaca irus
Macaca irus
Macaca irus
Macaca irus
Average of 1 2 Macaca irus
Macaca philippinensis
Maeaca phiIippiiieiisis
Macaca radiata
Macaca nemestrinus
Macaca mulatta
Macaca mulatta
Papio sphinx
Pygathrix cristatus
Pygathrix cristatus
Pygathris cristatus
Pygathris cristatus
Pygathrix cristatus
Qgathrix cristatus
Pygatlirix cristatus
Pygathrix cristatus
Pygathrix cristatus
Pygathrix cristatus
Pygathrix cristatus
Pyga thrix crista tus
Pygathrix cristat us
Ppgathrix crista tus
Average of 14 Pygathrix cris.
;OXTDITIOI
w. p.
w. p.
w. p.
f.
m. f
IT.
.
w.f.
f.
w.f .
1'. f.
w. f .
M'.
e. p.
w.p.
w.f .
f.
m. f .
w. f .
TV. f .
R.
w. f .
w, f .
w.f .
w.f .
w.f.
w.f.
p-.f .
tv. p.
w. p.
c. f .
m. f .
c. f .
e. f .
c. p.
c. p.
nr. f.
w.f .
w. f.
IT. f.
w. f.
TV. f.
w.f .
w.f .
w. f .
w. f.
m. f.
w.f.
w. f .
388
BODY
WEIGHT
TESTES
WhlCHT
Izg.
gin.
0.37
1.02
- c
1.12
6.92
7.03
7.03
7.26
7.26
7.48
7.82
7.SB
3.33
7.94
1.8
1.2
24.8
30.1
21.3
24.8
26.6
19.5
24.8
28.3
25.U
11.2
13.4
0.316
0.118
0.321
0.435
0.303
0.352
0.366
0.268
0.331
0.362
0.313
0.215
0.169
3.63
3.85
3.85
4.30
4.30
4.54
4.54
4.76
4.98
4.98
5.44
5.44
4.55
4.31
4.53
8.40
9.98
9.98
10.88
31.98
19.6
18.9
17.8
27.9
36.0
37.8
2 i.4
43.8
26.9
36.4
36.8
39.7
0.540
0.491
0.463
0.648
0.837
0.833
0.604
0.920
0.540
0.731
0.676
0.7.30
0.668
0.800
0.794
0.686
0.668
0.742
0.717
0.278
4.77
4.98
5.67
5.90
5.90
6.13
6.35
6.58
6.58
6.58
7.03
7.48
7.48
7.94
6.98
2.8
0.6
5.0
6.6
4.8
5.4
6.3
7.0
5.5
7.8
5.9
6.7
6.1
7.0
5.5
30.8
34.5
36.0
57.6
66.7
74.0
78.0
88.9
'ERCENTAGE
0.059
0.012
0.088
0.1 12
0.081
0.088
o.nw
0.106
0.084
0.118
0.084
0.089
0.081
0.088
0.085
SFRCIRS
ONDITION
BODY
WEIGHT
TESTES
IVEIGIIT
kg,
ym.
EZOENTAGE
~
Pygathrix rubicundus
Pygatiirix rubicundu9
Pygathrix rubicundus
Py g athrix r ubicundus
Pj-gathrix rubicundus
Pygathrix rubicundus
Pygnthrix rubicundus
P ? gathrix rubieuiidus
Pygathrix rubicundus
Pygatlririx rubicuiidus
Pygatlirix rubieuiiclus
Pygathrix rubicuiidus
Average of 1 3 P:gatlirix rubie.
Nasalis larvatus
Nasalis l a n a t u s
Nasalis larvatus
Nasalis larvatus
Nasalis larvatus
Nasalis larvatus
Nasalis larvatus
Sasalis larvatus
Average of 8 Nasalis lnrvat.
HJ-loba tes
Hylobates
Hylohates
Hylobates
Hylobates
cinereus
cinereiis
chereus
cinereus
cinereus
Hylobates cinereus
Hylobates cinereus
Average u f 7 Hylobatcs ciner.
Orang utaii
Orang utaii
Orang-utan
Orang-litan
Average of 2 adult orang-utaii
Chimpanzee
Cliimpauzce
Cliimpanzee
Average of 3 chimpanzees
Negro ( 3 7 years)
Negro (41 years)
Negro (31 years)
Average o f 3 Negroes
w. f.
f.
m. f .
R. f .
w.f.
w.f.
w.f.
w. f .
w.f.
w. f.
I\*.f.
IV.
I\’.
f.
w. f.
1v. f .
IT. f .
w.f.
17. f .
m. f.
71’. f .
w.f .
1v. p.
m. f.
w.f.
n-.f.
m. f.
IT. f .
IT. f.
f.
m. f .
TT.
w.f.
c. p.
c . 13.
e . p.
e . p.
f.
f.
f.
5.45
5.6;
5.67
6.12
6.12
6.12
6.36
6.35
6.35
6.58
6.58
7.03
3.6
3.9
3.4
3.4
3.2
4.3
4.9
3.7
3.7
0.039
0.0.53
0.060
0.059
0.064
0.056
0.053
0.050
0.068
0.074
0.056
0.053
3.4
6.30
3.6
o.n.5.9
14.06
20.42
80.42
20.42
20.57
20.87
23.14
23.59
97.47
5.2
7.32
16.0
11.8
9.6
14.6
18.0
13.2
4.98
4.98
4.98
5.44
5.90
6.12
6.35
5.5d
29.95
34.50
72.18
77.10
2.9
3.0
6.1
6.1
5.4
5.2
74.64
85.3
40.20
43.77
49.05
119.5
108.0
129.0
0.037
(1.035
0.078
0.058
0.046
0.070
0.078
0.056
0.057
0.058
0.060
0.122
0.112
0.091
0.085
0.058
0.084
0.097
0.068
0.045
0.050
0.048
0.297
0.247
0.263
44.34
118.R
0.269
34.89
66.30
69.42
43.0
51.0
36.5
0.078
0.077
0.081
6.7.,511
50.d
0.07.9
After €Irdli?ka ( ’25).
a One testis undescended and vestigcal.
Juvenile ( C and N3 not yet eruptcd).
w.,wild; c., captive; f., fresh; p., preserved (embalincd).
389
3.2
3.0
11.3
3.73
4.6
29.0
23.4
32.4
,382
390
ADOLPH H. SCHULTZ
The macaques and baboons (Macaea and Papio ) represent
the subfamily ('ercopitliecinae and the langurs and proboscis
monkeys (Pygathrix and Nasalis) the subfamily Semiiopithecinae of the entire family of Old World monlieys. Last
in the table are the higher primates : Gibbon (Hylobates),
orang-ntan, chimpanzee, and Ilian. Tnfortunately it was impossible to obtain actual weights 011adult male gorillas. Based
upon palpation of the scrotiim in several preserved bodies of
adult gorillas, the author does not hesitate t o state that the
testes of this aiithropoid ape are comparatively small and
appear to have a relative weight considerably below that in
man.
The ratio of testicular. weight to body weight arnonnts to
more than 0.1% i n a11 the New Worlcl monkeys examined. I t
is smallest in the night monkey (,4otns) and the spider monkey
(Ateles) and largest in the marmoset (Oedipomidas) and the
howling monkey (Blouatta). One of the latter was collected
and embalmed by the writer in Nicaragua, the other sere11
come from Panama and were weighed fresh, yet their ratios
appear fully comparable.
The relative weight of the testes differs to a very marked
extent in the two subfamilies of Old World monkeys. I n the
five species of macaques the relative weight of the testes
fluctuates beween 0.46 and 0.92 and the baboon lias a corresponding valne of 0.28. I n langnrs, 011 the other hand, the
relative weight of the testes shows very much smaller averagcs,
i.e., 0.08 in one species and 0.06 in another, the latter average
occnrring also in the closely allied proboscis monkey. Thus
it is found that a rhesus monkey, weighing riot quite 10 Bg.,
has testes weighing 74 gm., whereas proboscis monkeys with
an average body weight of over 20 kg. have an average testiciilar weight of only 11.9 gm.
Among the higher primates the chimpanzee has undoubtedly
the proportionately heaviest testes. Adult orang-utans possess
a smaller relative testicular weight than even the langurs.
This relative weight averages about the same in gibbons as
in man. The value for the latter agrees very closely with that
ITEIGHT O F TESTES IK PBI&IATES
391
giwn by Vicrordt (1688, p. 17), who lists 0.08 as relative
weight of tlie testes in adnlt man (presumably n'hite). It
is rather surprising t o fiiid that, e.g., a bonnet macaque
weigliing only 8.4 kg. has slightly heavier testes (57.6 gm.)
than a Negro weighing 69.4 kg. whose testes tip the scale at
56.5 gm. I n other words, per kilogram of body weight the
former has 6.9 gm. of testicdar tissue, idicreas the latter
only 0.8 gm. or eight and a half times less.
It is difficult t o suggest any reasons f o r these midc discrepancies in thc rclative weight of the testes among different primates. Tliep are iiot connected with any seasonal
changes in the life of the monkeys; at least the author's field
notes and the distribution of the ages of all the animals collected give 110 illdieation of any breeding season among the
Old World primates. It w a s attempted t o compare the relative size of the testes with the physiological age (as indicated
by the state of attrition of the teeth) of tlie specimens in ilie
series of macaques and of langurs, but it was soon apparent
that there exists no correlation whatsoever between these two
factors among adult animals.2
Tt seems possible, finally, that testes of widely differing
relative size may contain the various tissues in very different
proportions. Some of the testes, therefore, were sectioned
and stained f o r microscopic study. The author is greatly
indebted to his colleague, Dr. I. Gersli, for help in this undertaking. Sample sections of the testes were drawn thirty
times eiilarged with the camera lucida on heavy paper of
uniform thickness. The parts representing ( A ) the tubules
or the sex-cell producing glandular tissue were carefully cut
out and these as well as bhe remaining parts, representing
(B) the reticular and collagenous connective tissue, interstitial glandular tissne, blood vessels and nerves, were weighed
separately with the following results :
" I t is interesting to note t h a t the two juveiiilc o r a n g u t a n s i n the table possess
relatively larger testes t h a n the two full grown orang utans. On the other hand,
small series of juvenile macaques and gibbons showed much smaller relative
testieular weights than the corresponding series of adults.
392
ADOLPH H. SCHRTJLTZ
A
PRIMATE
(TUBULBB)
g m.
Langur
Gibbon
Negro
Chimpnnzee
Baboon
Macaque
0.06
0.06
0.08
0.26
0.28
0.72
B
(FONNBCTlVE
TISSUS)
A+B
m.
m.
4.51
4.49
4.52
4.50
4.49
4.49
2.15
2.30
2.54
3.17
3.31
2.36
3.08
1.41
2.19
1.98
1.33
1.18
Contrary t o what might have been expected, it is foiind that
the primates with relatively large testes (chimpanzee, baboon
and macaque) have a considerably greater proportion of sexcell producing glandular tissue and a smaller proportion of
connective tissue than have the primates (langur, gibbon and
man) with comparatively small testes. On the assumption that
the sample sections which have been analyzed are representative of the entire testes, it can be calculated that per kilogram
of body weight the langur possesses 0.29 gm. of tubular tissue
(A) and 0.31 gm. of connective tissue (B), whereas the
macaque 4.9 gm. of tubular tissue and 2.3 gm. of connective
tissue, This means that per unit of body weight the macaque
has seventeen times more of the sex-cell prodncing glandular
tissue than the langur.
Two of the adult, wild-shot primates in the table are marked
as having one testis undescended and vestigeal. A few notes
on this rare condition of unilateral cryptorchism are added
here since it appears to have some connection with the size
of the testes. I n one of the adult proboscis monkeys only
the riglit testis, weighing 7.0 gm., had descended in the scrotum
and the left had been retained in the abdomen, weighing only
0.2 gm. This is unquestionably an abnormal condition and
identical with the finding in an adult gibbon (Hylobates
cinereus) from Borneo. I n this ape the right testis was i n the
scrotum and the left way up in the abdomen; the former
testis weighed 3.4 gm. and the latter only 0.3 gm. Among
the male gibbons collected by the author in Asia there are two
further cases of a very similar nature, thongh they appertain
to juveniles. I n a HyIobates cinereus, weighing 3.18 kg., in
WEIGHT O F TESTES I N P R I M A T E S
393
which the permanent canines and third molars had not yet
erupted only the left testis of normal size was in the scrotum
while the right, of excecdingly small size, was found in the
abdomen. Exactly the same condition occurred in a Hylobates
lar, wcighing 3.40 kg. and with all permanent teeth, except the
third molars. This unilateral cryptorchism was found in
three out of sixty-two wild gibbons (twelve juveniles and
fifty adults), or in 5% of the cases. The two cases in jux-eniles
are not caused merely by a slightly retarded descent of one
testis, as is evident from thc vestigeal nature o f the testis
in the abdomen and also from the following observations on
the normal time of descent of the testes in gibbons, based upon
the author's field notes of last year :
HYMRATXS
BODY WBIGIIT
DhNPlTTON
TESTES IN
kg.
lar
lar
lar
lar
cinereus
lar
cinereus
cinereus
lar
lar
ein wells
cinereus
lar
la r
lar
0.45
0.68
0.68
0.68
0.91
1.36
1.36
1.59
2.04
3.15
3.40
3.40
3.63
3.83
4.08
Inc~ompletcmilk dentition
Complete niilk dentition
Complete milk dentition
Complete milk dentition
Complete milk deiititioii
Tncomplete pennanrnt dentition
lncomplete permanent dentition
Tncomplete permanent dcntitioii
Incomplete permanent dentition
Incomplete permanent dentitioii
Ineorriplete perinanent dentition
Incomplete permanent dentition
Incomplete permanent dentition
Incomplete permanent dentition
Incomplete lxrmanent dentition
Scrotum
Abdomen
Scrotum
Scrotum
Scrotum
Abdomen
Scrotum
Scrotum
Rcrotuni
Scrotum
Srrotum
ScrotuKtl
Scrotum
Swot 11m
Scrotum
NOTE: Adult male Hylobates Par hare a n average body weight of 5.7 kg.
a i d adult male Hllobates cinerens one of 5.5 kg.
These two spccimens were living i n captivity when examined, all the other
specimens, exnmined fresh immediately after death.
gibbons listed are ~ i l shot
d
I n his paper on the genitalia of primates Wislocki ( ' 3 6 )
lias listed one juvenile Hylohates lar and two juvenile H.
cinereus in all of which the icstes had dcscended in the scrotum.
It seems safe to conclude, therefore, tliat at least in these two
species of gibbons the testes descend normally during in-
394
ADOLPH H. SCHULTZ
fantile life, ie., before any permanent tpeth make their ~ p pearanc c.
SUMMARY
The weight of the body and of the testes and the percentage
relation between the two is recorded for a total of eiglltly
adult and two juvenile simian primates. This relative weight
of the testes fluctuates between 0.1 and 0.4 among American
monkeys. I n monkeys of the Old World the testes are proportionately much heavier in the suhfamilj- Cercopithecinae
The relative testiealar
than in the subfamily Scinno~~itlrecinae.
weight varies among five species of macaques between 0.46
and 0.92, whereas in langurs it averages oiily 0.06 and 0.08 in
different series. Among higher. primates this ratio amounts
to 0.05 in orang-utans, 0.08 in gibbons and in man, and 0.27
i n chimpanzees. Such striking differences i n the relative
size of the testes are best demonstrated by the example of
a bonnet macaque, weighing 8 kg., which has slightly heavier
testes than a Negro weighing 69 kg.
Small testes, relative to hodj7 size, coiitaiii a proportionately
miwh smaller amount of tubules and a much larger amount
of conncctive tissue than relatively large testes. It is calculated that, with equal body weight, a macaque, e g . , has seventeen times more sex-cell producing glandular tissue than has
a langur.
In one proboscis monkey and in three gibbons there exists
unilateral crpptorchism, i.e., one testis was retained i n the
abdomen and is of diminutive size and the other had descended
in the scrotum and is of usual size. Normally the testes of
these gibbons descend during infantile life.
LITERATURE CITED
A. 1925 Weight of the brain and of the internal organs in American
monkeys. Am. J. Phys. Anthrop., vol. 8 , pp. 201-211.
VImomr, H. 1888 Anatomischr, physiologiaehe nnd pliyikalischc Daten uud
Tabcllen zum Gebraiiche fiir Mediziiier. Jeiia.
WISLOCKI,
G. B. 1936 The erterual genitalia of thc simian primates. Human
Biol., vol. 8, pp. 309-347.
HRDIJCK-4,
Thc only juvenile Hylobates 1ar viith both testes still in the abdomen has
merely its first pcrmaiirnt molars besides thc complete milk delitition and appears
to be a n exception, though Wislocki records undescended testes in three juvenile
gibbons of other species (H. concolor and 13. leucogenys).
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