AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 86:37-44 (19911

Serum and Saliva Sex Hormone Levels in !Kung San Men

KERRIN H. CHRISTIANSEN
Institute of H u m a n Biology, Unioersity of Hamburg, 0-2000 Hamburg
13, Federal Republic of Germany

KEY WORDS Northwestern Kalahari, Androgens, Estradiol, Nu-

trition, Alcohol

ABSTRACT Serum concentrations of testosterone (Tser), 5cu-dihydrotes-

tosterone (DHT), estradiol17p (E2), and free testosterone in saliva (Tsal)were
determined by means of the radioimmunoassay method in 114 !Kung San men
living in the Bushmanland district of Namibia. The healthy men (mean age
26.4 years) were asked about their dietary habits over the last two months and
their acute alcohol intake during the 24 hours preceding the blood and saliva
sampling. Although the sex hormone status of the !Kung lies within the range
of normal men reported for Caucasoid samples, both Tser and Tsal exhibit
relatively low concentrations in comparison to the great majority of published
mean values. On the other hand, comparatively high DHT levels point to a n
elevated 5cu-reduction of testosterone to DHT in our sample. Estradiol concen-
trations show no deviation from normal values reported elsewhere for healthy
young men. Different dietary habits of the !Kung lead to significant differences
in their sex hormone status: both levels of Tsal and the androgen ratio
TsalPTser decrease with increasing supplement of the traditional hunter-
gatherer diet with domestic and Western food products. The amount of alcohol
consumed during the day before the blood and saliva sampling shows a
significant effect on the DHT metabolism, and the shorter the time after
drinking, the greater decrease of DHT and DHT/E2 can be observed.

The !Kung o r Zhdoasi are members of the
largest remaining single division of the San.
Geographically, the !Kung extend from north-
western Botswana across the northeastern
part of Namibia into southern Angola. Nu-
merous aspects of their biology, ecology, and
culture have been extensively studied and
described in the anthropological literature
(e.g., Gusinde, 1966; Eibl-Eibesfeld, 1972;
Marshall, 1976; Lee and Devore, 1976; Nurse
and Jenkins, 1977; Tobias, 1978; Lee, 1979;
Howell, 1979, Nurse et al. 1985). Only very
limited information is available on the sex
hormone status of the !Kung. This is a signif-
icant gap in anthropological knowledge of
these people. Androgens and estrogens not
only play a role in genital dimorphism be-
tween the sexes and human sexual behavior
but also are a major determinant of extra-
genital sexual dimorphism and physical
growth (Christiansen and Winkler, 1990;
Winkler and Christiansen, 1991).
In 1967, Tobias published the first data
concerning hormone levels in !Kungmen and
reported on their levels of estradiol and es-
trone. His findings that the !Kung have ele-
vated urinary estradiol values could not,
however, be confirmed by other researchers
who investigated the blood levels of the “fe-
male” steroid hormone estradiol and of the
androgen testosterone (van der Walt e t al.,
1977,1978; Worthman and Konner, 1987).
The present study was conducted to fur-
ther elucidate the sex hormone balance in
!Kung men. Serum levels of testosterone and
estradiol were determined as well a s the
concentrations of 5a-dihydrotestosterone in
the serum and of free testosterone in the
saliva. In addition, some major factors (age,
dietary intake, and alcohol consumption)
known to influence the intraindividual vari-
ability of sex hormone concentrations in
Caucasoid populations were investigated.
Received J u n e 1,1990; accepted March 24,1991

@ 1991 WILEY-LISS, INC

38 K.H. CHRISTIANSEN
MATERIALS AND METHODS
Population
The field research was conducted in vari-
ous locations of the Namibian Bushman-
land. Here, the Kalahari is a semi-desert
area of sand-veld or woodland savanna with
a n average rainfall of 300 to 500 mm (van der
Merwe, 1983).Population density is very low
with fewer than 0.25 persons per km2 and
the Bushmanland is predominantly inhab-
ited by San (over 90% of the population). The
!Kung in the Namibian Bushmanland have
until recently been hunter-gatherers, and to
a certain extent they still are. The savanna
carries a wide range of grazing and browsing
species which are hunted by the !Kung men
and which provide about 30% of the caloric
content of their diet. San obtain excellent
protein supplement from their extensive use
of wild vegetation. More than 70% of their
diet consists of vegetable food like the nutri-
tious mongongo nut (or mangetti) and about
100 species of roots and bulbs (Lee and De-
Vore, 1976; Truswell and Hansen, 1976;
Singer, 1978; Campbell, 1983). Even sea-
sonal shortage of calories a t the end of the
dry season when food is scarce is not
matched by any qualitative deficiency of es-
sential nutrients in the San’s diet (McElroy
and Townsend, 1985). A few San engage in
some form of temporary or permanent occu-
pation giving them access to cash money and
thus to domestic food products such a s milk
and maize meal and to Western food such a s
refined sugar products (there is one store a t
Tsumkwe, the administrative center of
Bushmanland). In almost every camp there
are elders who receive a monthly pension
from the Namibian government and who
usually share the money with their family.
Moreover, in !Kung society every member of
a camp participates in evening meals around
a fire (Lee, 1978). The amount of purchased
food is thus more evenly divided among
!Kung camp members than is to be expected
from the percentage of San who actually
have cash money from paid labor a t their
disposal.
Sample
During the southern winter months (June
and July 1987), 114 healthy !Kung men of
the Namibian Bushmanland participated in
this study. The majority of the men lived in
a n area (25 different locations) within 70 km
around Tsumkwe where they settled in
small more or less permanent camps made of
7 to 15 grass huts each. The camps were
located near waterholes with distances be-
tween settlements from 5 to 30 km. Accord-
ing to their own accounts, the !Kung men,
between the ages of 18 and 38 (26.4 2 4.7
years), had no parents or grandparents of
other racial origin. This corresponds well
with our morphological observations and
with data from Nurse et al. (19851, who
found hardly any or no gene flow between
the neighboring Kavango peoples and the
San.
All subjects were carefully examined in
order to exclude individuals with acute or
chronic diseases whose symptoms or medical
treatment thereof could possibly influence
their sex hormone status.
Hormone assays
In order to avoid the influence of diurnal
endocrine fluctuations as far a s possible,
blood and saliva samples were collected in
the mornings within one to two hours after
rising (between 8:OO and 10:30 a.m.). The
blood samples were drawn through conven-
tional venipuncture into lithiumized vacu-
tainers and allowed to clot for one hour
before the serum was separated through cen-
trifugation. The subjects provided a saliva
sample after carefully rinsing their mouths
with cold water in order to prevent a contam-
ination of the salivary sex hormones by ex-
ogenous steroids from blood or food (Riad-
Fahmy et al., 1982; Ellison, 1988). Because
subjects with more serious oral bleeding
might continue to contaminate their saliva
while spitting into the collecting container,
all saliva samples were tested by using dip-
stick indicators designed to detect the pres-
ence of blood in urine (Hamo dip, Merckog-
nost, West Germany). Both serum and saliva
were immediately stored at -20°C in a n
electrical deep freezer at Tsumkwe. The
samples remained deep frozen in a portable,
battery-run freezer during the transporta-
tion from the field to the laboratory at the
Institute of Human Biology in Hamburg
where they were assayed four to eight
months later.
The following sex hormone concentrations
were determined by specific and sensitive
radioimmunoassays (RIA):
Androgens: total testosterone in the serum
(Tser)
5a-dihydrotestosterone in the
serum (DHT)
 SEX HORMONE LEVELS IN !KUNG MEN 39

free testosterone in the saliva maize, sorghum and cow’s milk; more regu-
(Tsal) lar consumption of domestic food as supple-
Estrogen: estradiol 17p in the serum (E2) ment of the traditional subsistence was clas-
sified as category I11 (cat 111);a mixed diet of
All samples were analyzed in duplicate in traditional and domestic food with more or
the same assay. less regular intake of Western food such as
Androgen concentrations in the serum refined sugar, tea, or dried milk was charac-
(Tser and DHT) and in the saliva (Tsal)were terized a s category IV (cat IV).
determined by means of commercial tritium- The subjects’ alcohol intake was assessed
marked RIA-kits (T/DHT-3H-Kit, Charcoal by asking them to remember and report the
Technique, Amersham Buchler, West Ger- amount of alcoholic beverages (in cups or
many). The standards were 12.5 to 400 pgl glasses) consumed during the 24 hours pre-
tube for T and 25 to 800 pgltube for DHT. The ceding the venipuncture as well as the num-
correlation coefficients of the logit-log trans-ber of hours elapsed since the last drink. Men
formation of the standard curves were who consumed alcohol had exclusively drunk
r = 0.996 for T and r = 0.995 for DHT. The self-brewed beer made of brown sugar, ber-
intra- and interassay coefficients of varia- ries or grain, and other ingredients such a s
tion were within the usual limits reported hops, honey, and spices (cf. Brooks et al.,
elsewhere (Tser: 6.6% and 11.4%; DHT: 1984), with low alcoholic content. Three cat-
10.0% and 12.9%; Tsal: 7.1% and 14.3%). egories of alcohol intake during the preced-
Blanks were indistinguishable from zero. ing 24 hours were observed: abstinent (no
Estradiol concentrations were analyzed alcohol), moderate consumption (maximum
with very sensitive iodine-marked RIA-kits 1.0 1 of self-brewed beer), and considerable
( lZ5I- Oestradiol -Extraction -Radioimmuno- alcohol consumption (more than 1.0 1of home-
assay-Kit from Baxter Merz + Dade AG, made beer).
Switzerland), especially suitable for low E2
levels as they occur in males. The standards Statistical analysis
ranged from 0.5 to 50 pgltube. The sensitiv- Means, standard deviations, skewness,
ity of the standard curve was 0.25 pg/ml. The and kurtosis were computed for the absolute
intra- and interassay coefficients of varia- hormone values and for the hormone ratios
tion (8.2% and 13.1%)were well within ac- indicated in Table 1. Checking the distribu-
cepted limits. Blanks were consistently in- tions for skewness and kurtosis (Table 1)
distinguishable from zero. Intralaboratory reveals that only Tsal, TserE2, and DHTE2
quality of the E2-RIAs was tested with im- have a t least a normal kurtosis (K4). There-
munoassay control serum (Baxter, Munich, fore, only the distribution free rank coeffi-
West Germany) showing the very high reli- cient of Spearman was used to calculate the
ability of the E2 determination, especially a t correlations between hormone levels and the
the low E2 levels of our subjects. other variables. Analyses of variance (unbal-
anced ANOVA, General Linear Model) and
Consumption of food and Tukey tests were carried out in order to test
alcoholic beverages the differences in mean sex hormone levels of
To assess the dietary habits of the sub- groups with different dietary patterns. Stu-
jects, the men were required to give infor- dent’s t-test was used to analyze group dif-
mation about their usual consumption pat- ferences in sex hormone concentrations in
tern over the last two months according to relation to alcohol consumption.
their own memory. Although the diet recall
method is not a highly valid technique, this RESULTS
method is inexpensive and not time consum- Sex hormone levels
ing, and therefore is widely used in field The mean hormone levels, standard devi-
research settings (Quandt, 1987). The de- ations, skewness, and kurtosis are given in
tailed information from our subjects was Table 1.Numbers of subjects are inconstant
classified into four categories: category I for various reasons. We had great difficulties
(cat I) comprises intake of food obtained only with some probands a s in obtaining enough
from traditional hunting and gathering; cat- serum through venipuncture due to the very
egory I1 (cat 11) includes food from tradi- cold desert mornings soon after sunrise. In
tional resources and additionally very rarely addition, some saliva and serum samples
consumed domestic food products like beef, were inadvertantly lost in transport from the
40 K.H. CHRISTIANSEN
TABLE 1. Means, standard deuiation. skewness (k3), and kurtosis (k4) o f sex hormone levels in San men
Hormone N Med
Tser (ng/ml) 107 4.58
DHT (ng/ml) 107 1.15
Tsal (pg/ml) 110 76.69
E2 (pg/mU 102 23.57
DHT/Tser ,100 107 26.34
Tsal/Tser ,100 103 1.67
Tser/E2 99 205.71
DHT/E2 99 5n fifi
*One-tailed test of skewness (k3) and kurtosis (k4): P < .01.
field or during processing in the laboratory.
There were no losses of saliva samples be-
cause of contamination with blood after oral
bleeding. Although 15 men had minor, non-
visible free haemoglobin in their saliva
(corresponding to 10 to 50 erythrocytes/pl:
11 men; corresponding to 50 erythrocytes/
p1: 2 men; corresponding to 250 erythro-
cyteslpl: 2 men), we could not detect a signif-
icant systematic rise in their salivary test-
osterone concentrations a s compared with
uncontaminated saliva samples.
Sex hormone levels and age
Rank correlation of sex hormone levels
and age of subjects yielded no significant
coefficients. In our study group, with a n age
range of 18 to 38 years, only small (rho < 5
0.16) positive or negative correlation coeffi-
cients were found.
Food and alcohol consumption
Only one out of 114 !Kung reported having
sustained himself exclusively from tradi-
tionally gathered or hunted food during the
preceding two months (cat I). All other men
supplemented their main diet of traditional
food either rarely (cat 11: n = 8) or more
regularly (cat 111: n = 40)with domestic food
or, in addition to traditional and domestic
products, ate some Western food (cat IV:
n = 65). Only men from categories I1 to IV
were included in the further analysis.
Moderate alcohol consumption (< 1 1 of
beer) during 24 hours preceding the veni-
puncture occurred in 72 cases. One man had
consumed 3 to 4 1 of beer and 41 !Kung had
remained abstinent. The one subject with a
higher alcohol level was excluded from sta-
tistical group comparisons.
Diet, consumption of alcohol, and sex
hormone concentrations
The results of unifactorial variance analy-
ses carried out for absolute hormone values
-
X S1) k.? k4
4.83 2.04 0.98* 1.08*
1.30 0.60 1.14* 1.38*
80.10 36.02 0.94' 2.22
24.89 10.63 0.97* 0.87*
32.35 20.92 1.17' 0.59'
1.94 1.20 2.16' 6.90'
219.81 107.76 1.41* 2.54
59 62 32.06 1.27* 2.50
~
and hormone ratios of groups with different
dietary habits (cat 11,111,and IV) are given in
Table 2. Mean levels ofTsal (F-test, P < .Ol),
DHT (F-test, P < . l o ) and TsaYTser (F-test,
P < .001) significantly decrease with in-
creasing intake of domestic and Western
food. A significant F-value ( P < .05) for the
ratio DHTPTser, however, is not connected
with such a systematic decrease among the
three diet categories. Additional Tukey tests
(Cochran and Snedecor, 1974) revealed that
the subjects eating mainly traditional food
(cat 11)had only significantly elevated mean
concentrations of Tsal and TsaLTser as com-
pared with subjects eating a mixed diet of
traditional, domestic, and Western food (cat
IV). However, neither of these group means
is significantly different from that of the
middle category (cat 1111, representing men
with a mixed diet of traditional and quite
regularly consumed domestic food.
Men who drank alcohol (up to 1 1 of self-
brewed beer) during the 24-hour period be-
fore the venipuncture had significantly
lower DHT concentrations in comparison
with subjects who had remained abstinent
(Table 2). Such a tendency (P < . l o ) could
also be observed for Tsal and the ratio DHT/
E2. When the amount of time after the last
intake of alcohol was considered, a positive
correlation between time elapsed and DHT
(r = 0.21, rho = 0.17) and DHTA32 (r = 0.25,
rho = 0.17) could be found ( P < .05 for r and
P < .10 for rho).
DISCUSSION
Sex hormone Eeuels
Previous researches on the endocrine sta-
tus of !Kung men provided information for
only two (Tser and E2) of the sex hormones
which were investigated in the present
study. Up to now, no data have been avail-
able for DHT and the free fraction of tes-
tosterone, Tsal, a s well as for the interrela-
tion of the androgens and the ratio of Tser
 SEX HORMONE LEVELS IN !KUNG MEN 41

TABLE 2. Analysis o f variance f o r mean sex hormone levels and usual diet (cat II: mainly traditional food; cat III:
traditional and domestic food products; cat IV: traditional, domestic, and Western food); t-tests between
the group o f alcohol consumers and abstinent men during the preceding 24 hours
Tser DHT Tsal E2 DHT/Tser Tsal/Tser Tser/E2 DHT/E2
Type of diet n=106 n=106 n=109 n=101 n=106 n=102 n=98 n=98
Diet cat I1 X 4.881 1.641 101.3g1 30.61l 34.39' 2.99l 167.04' 57.18l
Diet cat Ill f 4.341 1.401 89.89',2 24 25' 38.58' 2.301,2 206.69' 64.61'
Diet cat IV X 5.19l 1.211 71.822 24.71' 28.12' 1.572 235.07I 57.13)
F-value 2.10 2.28* 4.60*** 1.23 3.08** 7.66**** 1.67 0.60
Alcohol abstinent X 4.77 1.48 88.31 25.81 36.20 2.17 206.82 68.11
1 1 beer X 4.82 1.21 76.18 24.39 30.56 1.84 224.88 55.85
t.V~ll1P -0.12 2.04** i.m* 0.63 1.33 1.31 -0.77 1.80'
"Group means with same number do not differ significantly
* P < .in.
** P < .n5.
***P< .01.
****P< .001

and DHT to E2. Therefore, our results have
to be discussed mainly in comparison with
sex hormone values from Caucasoid samples
and with data obtained from Kavango men,
a neighboring Bantu-speaking population
from northeastern Namibia, who were inves-
tigated one month later during the same
field research period (Christiansen, 1991).
The mean level of total testosterone, Tser,
in the !Kung subjects corresponds well with
the findings of van der Walt et al. (1977,
1978) and also with data from Worthman
and Konner (19871, who monitored the Tser
concentrations in !Kung men during a six-
day hunt. All values reported for San men lie
within the normal range of healthy young
men but definitely on the lower end of the
distribution of the Tser mean values from
European or American males. In the major-
ity of studies, the morning peak values of
Tser are around 6 ng/ml, and several au-
thors published mean Tser concentrations in
males clearly above 7 ng/ml (Moll et al., 1981:
8.0 ng/ml; Bremner et al., 1983: 7.4 ng/ml;
Christiansen et al., 1984: 8.1 ng/ml; Knuss-
mann et al., 1985a: 7.67 ng/ml; Spratt et al.,
1988: 7.54 ng/ml). Moreover, the comparison
of the !Kung Tser concentrations with the
Tser levels of Kavango men from Namibia
shows that the latter have a significantly
higher mean Tser value (x = 6.26 ng/ml; t-
test: P < .0001). Thus, our results together
with previous findings (van der Walt et al.,
1977, 1978; Worthman and Konner, 1987)
suggest that San men have, on the average, a
comparatively low level of total testosterone.
A very similar result as for the level of Tser
can be observed for Tsal, the free fraction of
testosterone in the saliva, which is not bound
to carrier proteins such a s the sex hormone
binding globulin (SHBG). Compared to Cau-
casoid samples of matching age whose peak
values usually range between 100 and 200
pg/ml with a majority around 150 pg/ml
(Khan-Dawood et al., 1984; Christiansen
et al., 1984; Meikle et al., 1987; Knussmann
et al., 1985a, for a review of the pertinent
literature before 19841, the San men defi-
nitely exhibit relatively low Tsal concentra-
tions. They also have significantly lower
Tsal levels than the Kavango men from
northern Namibia (x = 97.18; t-test:
P < .0002; Christiansen, 19911, and a sam-
ple of male Efe, pygmy hunter-gatherers in
the Ituri Forest of Zaire (Ellison et al., 19891,
whose morning levels of Tsal lie at 420 pmoM
(approx. 145.5 pg/ml). As both Tsal and Tser
levels have a likewise rather low level as
compared to other samples, the ratio TsaV
Tser (approximately 2%) is equivalent to
findings in Caucasoid samples (e.g., Ver-
meulen et al., 1971; Baxendale et al., 1980;
Moll et al., 1981; Wang e t al., 1981; Chris-
tiansen et al., 1984; Knussmann et al.,
1985a; Navarro et al., 1986) and in Kavango
males (Christiansen, 1991).
DHT, which is the main metabolite of
testosterone, reaches serum concentrations
from 0.51 to 1.34 ng/ml in European males
and the majority of the reported values lies
under 1 ng/ml (Pirke and Doerr, 1975; Ver-
meulen, 1976; Christiansen et al., 1984;
Knussmann et al., 1985a;Meikle et al., 1987;
Dericks-Tau and Taubert, 1987). The mean
DHT level of the San males lies within these
usual limits, but it points to a comparatively
high peripheral activity of 5a-reduction of
testosterone in our sample. However, in the
neighboring Bantu-speaking population of
Kavango males even higher DHT values
42 K.H. CHRISTIANSEN
were found (x = 1.67; t-test: P < .0002). As
the Namibian samples belong to the same
age group as the Caucasoid samples referred
to, these findings of rather high DHT levels,
especially in the Kavango males, cannot be
explained by age effects. It is more likely
that elevated DHT concentrations are con-
nected with nutritional factors as is indi-
cated by decreasing DHT levels when the
traditional hunter-gatherer diet is increas-
ingly replaced by domestic and Western food.
When DHT is related to Tser concentra-
tions in the San sample, the resulting hor-
mone ratio DHTITser is comparatively high
due to the more elevated DHT together with
relatively low Tser levels. Data from Euro-
pean men yielded ratios of DHT/Tser be-
tween x = 13.97 and x = 16.94 (Chris-
tiansen et al., 1984; Knussmann et al.,
1985a) which are only half as high as the
mean DHT/Tser level of the San.
Comparing the mean E 2 levels of our
study group with values from non-San sam-
ples shows that, on the average, the !Kung
men have estradiol concentrations which are
normal for Caucasoid males of the same age
(Kley, 1975; Kley et al., 1979; Kicovic et al.,
1980; Christiansen et al., 1984; Knussmann
et al., 1985a;Meikle et al., 1987; Spratt et al.,
1988) and for their Kavango neighbors
(Christiansen, 1991). The E2 levels in our
!Kung men correspond well with data of two
other San samples previously published (van
der Walt et al., 1977, 1978; Worthman and
Konner, 1987). In these studies E2 was also
determined with the radioimmunoassay
method. Neither of these studies on !Kung
men could confirm the earlier finding of
Tobias (1967) that San men show an “over-
oestrogenization,” as he concluded from ele-
vated urinary estrogen levels in his small
sample (and on the basis of the less reliable
laboratory methods of the 1960s). Thus,
there is no longer support for his hypothesis
that the paedomorphous bodybuild of the
San is due to their elevated estrogen levels
(Christiansen and Winkler, 1990).
The ratio TserlE2 in San men is lower than
in other samples of healthy men, a conse-
quence of relatively low Tser levels in !Kung
men of our study. Their mean value differs
considerably from TserE2 levels in Euro-
pean (x < 350.0: Christiansen et al., 1984;
Knussmann et al., 1985a) and Namibian
males (X = 265.23, Christiansen, 1991)
whose Tser and E2 concentrations were all
determined with the same assay technique
in the same laboratory. On the other hand,
no significant differences in the DHTlE2
level could be found between San men and
other published data for healthy young
males.
Diet, consumption of alcohol, and
hormone concentrations
Varying degrees of contact with other cul-
tures led to differences between our subjects
with regard to the supplement of their regu-
lar diet with Bantu and Western food. This
enabled us to investigate the hormonal sta-
tus of San men in relation to their food
intake. However, the significant results in
the present study cannot be interpreted as
showing a causal relationship between food
intake and sex hormone levels on the basis of
our research design alone. But more elabo-
rate investigations which focused on the ef-
fects of nutritional factors on the sex hor-
mone status in men support our findings and
thus indirectly support our hypothesis of a
causal relationship between the San men’s
diet and their androgen levels.
The traditional hunter-gatherer diet proves
to be high in protein (average daily consump-
tion of protein in a San sample was 93 grams
per person [Lee, 19681) and exceeds the US
government recommended daily allowance
for people of the size, stature, and vigorous
activity of the San by 33 grams of protein
(Campbell, 1983). Bantu and Western diets
provide higher levels of carbohydrate or fat
and less protein. By comparison with the
men who ate mainly traditional hunter-
gatherer food (cat II), those men who supple-
mented their diet regularly with consider-
able amounts of Bantu and Western food
(cat IV) exhibited a significant decrease in
mean Tsal levels as well as in the ratio
Tsal/Tser. Our findings of parallel changes
in the proteidcarbohydrate balance of the
San diet and in the Tsal and TsalPTser con-
centrations are supported by Knussmann et
al. (1985b). They found, in healthy young
men, a significantly positive correlation be-
tween Tsal and total protein levels in the
serum, and the percentage of free testoster-
one, expressed by the ratio TsalPTser, was
also positively correlated with total serum
protein. Similar results were published by
Smith et al. (1975),who reported a decrease
of TsalPTser in Indian men suffering from
protein malnutrition. The mechanisms by
which an alteration in the protein/
carbohydrate ratio of a diet can produce
 SEX HORMONE LEVELS IN !KUNG MEN 43
changes in free testosterone concentrations ested San men. Assistance in the field was
may be explained by the study ofAnderson et given by H Muth, L Penny, the !Kung inter-
al. (1987),who concluded that changes in the preters //Ci//que and !Kwi, and K. Gaerdes.
proteidcarbohydrate ratio of a diet influ-
ence plasma steroid concentrations and also
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SHBG concentrations were significantly Wissel PS, and Kappas A (1987) Diet-hormone in
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and to Dr. K.F.R. Budack, Windhoek, for his Kicovic PM, Luisi M, Cortes-Prieto J, and Franchi M
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not have been carried out without the partic- Kley HK (1975) Ostrogene im Plasma des Mannes.
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