RESEARCH PAPERS
Trace Element Deficiencies and Fertility in Ruminants:
A Review I
ABSTRACT
Various minerals (copper, cobalt,
selenium, manganese, iodine, zinc, and
iron) can influence reproductive perform-
ance of ruminants. Reproductive failure
may be induced by deficiencies of single
or combined trace elements and by
imbalances. This review is focused on
maladjustments of trace elements leading
to impaired breeding performance.
Opinion is diverse as to the existence of
various reproductive disturbances from
either a severe copper depletion or a
marginal dietary copper deficiency. Field
experience suggests that administration of
cobalt to ruminants on cobalt-deficient
diets improves their impaired breeding
performance. Selenium infertility in ewes
is more prevalent in some areas and in
some seasons, but the actual cause of this
malady and the continuing role of addi-
tional factors are unknown. Manganese is
necessary for normal fertility in rumi-
nants, and feeding low-manganese rations
depresses conception rates. Lack of
iodine impairs thyroid activity and also
ovarian function. Reproductive failure in
the female and in spermatogenesis are
manifestations of zinc deficiency. Despite
forages rich in iron, low availability in
certain instances could affect adversely
ruminant reproduction.
Knowledge of biochemical dysfunc-
tions from trace element deficiencies is
essential to determine the role which
trace elements play in fertility of ruminant
animals.
Received February 26, 1979.
t Contribution No. 827.
1979 J Dairy Sci 62:1195-1206
M. HIDIROGLOU
Animal Research Institute,
Research Branch
Agriculture Canada
Ottawa, Ontario K 1A 0C6
INTRODUCTION
Trace elements may function as cofactors, as
activators of enzymes, or stabilizers of secondary
molecular structure (103). Their study has
evolved from a recognition of their essential
function in cell metabolism. There has been
special interest in effects of dietary trace
element deficiencies on physiological functions
in general and in reproduction in particular
(56). Ruminants frequently are subject to
severe dietary deficiencies of trace elements
such as copper, cobalt, selenium, iodine,
manganese, and zinc. Concomitant infertility in
cattle is believed to be associated with enzymatic
dysfunctions resulting from these deficiencies.
Despite the accumulation of data suggesting
that reproductive performance in ruminants
depends upon adequate trace element nutrition,
there has been no comprehensive review of this
information.
Mineral deficiency usually involves several
minerals as well as other conditioning factors;
however, the deficiency symptoms of one
mineral may predominate and affect the
performance of the ruminant. The effects of
the lack of a single element may include repro-
ductive failure. In this respect effects of
copper, cobalt, selenium, manganese, iodine,
zinc, and iron malnutrition on reproduction
will be reviewed.
Copper
A variety of clinical symptoms have been
associated with copper deficiency. Hypocuprosis
in cattle is associated with female reproductive
disorders. Where such disorders have been
attributed to copper deficiency, the most
common symptom has been prenatal mortality,
particularly early embryonic loss. Most of the
relevant studies have been with sheep or goats.
A low copper content in the diet of the ewe
either prevented implantation or induced
1195
1196 HIDIROGLOU
embryonic loss and fetal death (61, 62). Of
nine ewes fed a severely copper-deficient diet
and then bred, five did not become pregnant
and died between the 23rd and 34th wk of the
experiment; of the remaining four ewes, two
aborted and two produced stillborn lambs.
Feeding female goats a semipurified diet low in
copper content lowered conception rates (4);
of the pregnant goats affected by copper
deficiency, 50% aborted. Fetal death occurred
between the 2nd and 5th mo of pregnancy. The
fetuses were mummified, and at abortion
placenta had degenerated with hemorrhagic or
necrotic lesions.
Contrary to expectations, concentration of
copper in blood does not appear to be related
directly to reproductive performance. Repro-
ductive performance of rams and their plasma
copper were unassociated (106). In the ewe,
low copper in blood did not reduce the size of
the lamb crop (45).
Copper deficiency may affect reproductive
behavior as well as performance; for example,
older female goats kept on a low copper diet
for an extended period showed nymphomania
(5). In these clinical field studies possible asso-
ciated changes in gonadal steroid metabolism
were not investigated.
Reliable information is limited for the
influence of absolute or conditional copper
deficiencies (such as results from a low copper
diet with excess m o l y b d e n u m and/or sulphur)
on the fertility of cattle. One of the earliest
signs of copper deficiency in cows is a decline
in fertility. Marked improvement results from
copper administration, provided there are no
other deficiencies (42, 43). Opinions from
casual observations in the field usually have
been contradictory with few well planned
studies yielding unequivocal data.
Generally, there is no clear relation between
copper in the blood and liver of cattle and their
fertility (54, 85). In some herds of cattle with
low blood copper, female fertility has been
impaired; yet in others conception rate has
been high (78). Administration of copper orally
or b y injection has led to a rapid and dramatic
improvement in breeding performance (58). In
New Zealand in herds in which blood copper
was marginally subnormal, a single administra-
tion of 400 mg of copper glycinate improved
fertility. The conception rate for the copper
treated cows was 72% versus 53% for the
Journal of Dairy Science Vol. 62, No. 8, 1979
untreated cows; treated cows required 1.00 and
untreated 1.15 services per conception (47).
Dietary supplement of copper has improved
the reproductive performance of cattle (96). A
high incidence of infertility in cattle in Australia
(11) was associated with cases of "falling
disease", anemia, and retarded growth in
areas where pasture copper was less than 3
ppm. This was overcome by parenteral adminis-
tration of copper. In Canada an increase in the
copper:molybdenum ratio was associated with
an improvement of fertility in cows fed si-
lage (72), and in other studies infertile cows
showed marked improvement in their concep-
tion rate when treated parenterally with copper
glycinate (13, 80).
In bulls, the administration of copper
improved semen quality (48). The improvement
was associated with increased sperm mobility
and fewer dead spermatozoa. However, situa-
tions have been described in which copper
deficiency in young livestock was severe, but
effects on breeding performance were not
adverse (96).
The pituitary gland and ovaries of cows
contained 9.5 and 4.8 /ag copper/g dry weight
(93). The average zinc and copper concentra-
tions in the fluid of normal and cystic follicles
of cows and heifers in (73) were 3 mg/liter Zn
and 2.1 rag/liter Cu. No differences between
cows and heifers in the follicular fluid content
of either element were detected, nor were there
significant correlations between liver and
follicular fluid copper and zinc. Zinc and
copper concentrations in fluid of follicular
cysts from anestrus cows decrease as the
volume of the fluid increases, indicating that
content per follicle is constant and suggesting
that these elements are intracellular in the
follicle. The uterine fluid of cows with cystic
endometrial hyperplasia contained 13 mg/liter
zinc and 12 mg/liter copper. Again there was
an inverse relation between volume of fluid and
copper and zinc concentrations (73).
Metabolism of copper appears to be altered
during pregnancy. Copper in blood plasma of
ewes fails during pregnancy and rises after
parturition (20). In pregnant cows, however,
ceruloplasmin and plasma copper have increased,
the highest attained in the 5th mo of pregnancy
(22). Parturition also may be influenced by cop-
per deficiency. Cows with retained placentas had
lower serum copper than normal cows' (105),
 REVIEW TRACE ELEMENTS AND FERTILITY
There seems to be an interaction between
estrogen and copper. Diethylstilbestrol adminis-
tration increased copper in the rat serum (98).
In humans receiving diethylstilbestrol therapy,
ceruloplasmin or copper, increased in plasma
apparently in proportion to the dose of the
estrogen administered (15). Changes in plasma
copper in rats parallel changes in plasma estrogen
during the estrous cycle, after ovariectomy, and
during administration of exogenous estrogen
(81). The relevance of these findings to similar
interactions in ruminants, however, is not clear.
The administration of estrogen to wethers and
ewes for 3 mo did not affect copper in their
blood or liver (69). Although the dosage of
estrogen was physiological, the lack of any
effect does not eliminate the possibility of
nonphysiological interactions having pharmaco-
logic interest.
Conclusions
Findings regarding the effect of copper
deficiency on the fertility of ruminants conflict.
It is difficult to ascertain whether reproductive
function is affected directly by the lack of
dietary copper or by some general dysfunction
produced by copper deficiency. That otherwise
normal cattle with a history of reproductive
failure can be cured by copper administration
suggests that the effect is specific. However,
when cattle are affected by severe hypocuprosis,
characterized by gross deficiency symptoms,
copper therapy alleviates the physical conditions
including infertility, thus suggesting that the
effect is general. Nevertheless, there are reports
of hypocupremic cows which appear normal
and are also fertile. One might compare these
differences in fertility to the variance in the
clinical manifestations of copper deficiency.
For example, there is a form of copper defici-
ency which is clinically silent and results in
only marginally suboptimal fertility in cattle.
This differs from the "persistent scouring"
symptom common in Holland and from the
"fibrosis of myocardium" which occurs in
Australia.
Further study is needed to clarify xhe
mechanism by which copper is involved in
reproductive processes. Response to copper
deficiency must take into consideration inter-
actions with other dietary factors such as the
availability of copper which may be influenced
by proteins which, in turn, affect copper
1197
absorption (101). In addition, complex forma-
tion between copper, molybdenum, and sulfur
may occur in the rumen which also will affect
the availability of copper (100, 1 0 2 ) a n d
produce a deficiency despite apparently normal
copper intake.
Cobalt
Field experience suggests that cobalt defici-
ency impairs breeding performance in both
cattle and sheep.
The most common manifestation of cobalt
deficiency is a marked reduction in conception
rate. In tropical areas such as Central Africa
(Zaire) or the northern part of South America
(Northern Brazil, French Guyana) in which
cobalt deficiency is endemic, the calving
interval encountered is often greater than 24
mo instead of the usual 14 mo (Hidiroglou,
unpublished). The reason for this is not yet
known. However, several studies appear relevant.
Involution of the uterus is completed within 3
wk after calving in cows treated with cobalt but
requires as much as 6 to 9 wk in cobalt-deficient
cows. Moreover, the cobalt treated cows have
stronger manifestations of estrus as well as a
lower incidence of irregular estrous cycles and a
higher conception rate (56). A Shorthorn herd
grazing a cobalt deficient pasture had a concep-
tion rate at first service of 53% compared with
67% in a similar herd grazing in the same
area but given copper therapy and 93% in a
herd given cobalt as well as copper therapy
(79).
In studies with sheep, cobalt deficiency was
associated with a reduction in estrus during the
normal breeding season. Dietary cobalt supple-
mentation increased the incidence of estrous
activity (110) and increased sperm counts
of males (7).
Conclusions
Cobalt is required to ensure fertility in
ruminants. In cobalt deficiency conception rate
of cattle declines while in the ovine lack of
estrus is one of the features of low dietary
cobalt intake.
Selenium
There are conflicting opinions of the effect
of selenium deficiency on reproductive perform-
ance in the ruminant. In those studies of
Journal of Dairy Science Vol. 62, No. 8, 1979
119 8 HIDI ROGLOU
impairment of fertility associated with selenium
deficiency, the locus of the impairment generally
has been embryonic or fetal loss. In several
cases of selenium deficiency, administration of
selenium prevented certain reproductive prob-
lems. Lambing percentages have increased in
controlled trials with ewes when selenium was
administered orally in monthly doses beginning
1 mo before mating and continuing through
pregnancy (33, 34, 35). Reduced fertility in the
untreated animals was attributable to embryonic
mortality at 20 to 30 days of pregnancy (3).
Ewes grazing on pasture of low selenium and
high estrogen content were given selenium 4 to
g wk before breeding. Their conception rate
increased from 49% to 76% (30). Significant
improvement in lambing rate and incidence of
twinning in ewes has been reported when the
diet was supplemented with selenium and
copper (46); the twinning rate was depressed by
a selenium deficiency with fewer lambs born to
the control ewes.
In contrast, studies on the effect of selenium
given to ewes in another selenium-deficient area
in Australia revealed no significant influence on
conception rate (23). Similarly, in areas where
white muscle disease in sheep had been diag-
nosed, selenium supplementation showed little
or no effect on sheep fertility (29). Nine trials
with ewes on low selenium pastures in Western
Australia failed to show an improvement in
reproductive performance when up to 50
mg of supplementary selenium was administered
prior to breeding (60).
In other studies, there were no significant
differences in rates of ovulation or embryonic
loss between ewes fed selenium-deficient hay
and those fed hay plus a selenium supplement.
Overall fertility in selenium-deficient ewes was
not lower than that of selenium treated ewes
(67).
In cattle also, there are contradictory reports
of a selenium requirement for the maintenance
of fertility. Low selenium (less than .05 ppm)
appears to account, at least in part, for infertility
of dairy herds in some areas in Uganda (55). In
other studies, however, prebreeding selenium
treatment of beef cows in areas deficient in
selenium did not improve subnormal pregnancy
rates (83, 91).
Interactions between selenium intake and
the availability of vitamin E have become
apparent through the research of a number of
Journal of Dairy Science Vol. 62, No. 8, 1979
workers who have indicated the difficulties of
interpreting observations based on situations in
which one of these factors is controlled. It is
possible that the onset o f infertility and its
severity under field conditions are associated
with seasonal changes in selenium and/or
vitamin E in forages. Such fluctuations may
account for some of the conflicting observations
regarding the influence of selenium on fertility.
Studies with purified diets have shown no
differences in the fertility of selenium-depleted
and of selenium-sufficient ewes, provided that
adequate amounts of vitamin E are supplied
(18, 19). In cattle, administration of vitamin E
and/or potassium selenate 1 mo before calv-
ing greatly reduced the incidence of retained
placenta in a herd of dairy cows fed little
selenium during the precalving period (97).
Effects of selenium and vitamin E, alone or
in combination, on fertilization of ova have
been studied in beef cows on either a low or an
adequate plane of nutrition (86); the fertilization
rate was 100% for females on an adequate
plane of nutrition given a supplement of
selenium and vitamin E. However, rates of
fertilization were lower when the diet was
deficient in either selenium or vitamin E or was
below maintenance requirement for protein and
energy (86).
Little information is available concerning the
influence of selenium on fertility in the male.
Only one study deals with ruminants, and this
indicates that selenium does not appear to
influence testosterone secretion in rams (18).
In rate, sperm mobility was poor in selenium-
deficient males. The spermatozoa showed
breakage near the mid-piece in most cases.
Vitamin E supplementation, even high, did not
alleviate these symptoms (108). The only
studies o f interactions of selenium with en-
docrine secretion, other than that just cited,
concerns the rat in which selenium of blood fell
between days 10 and 15 of gestation (9).
Conclusions
Reduced fertility sometimes is associated
with selenium-deficiency in sheep and cattle.
However, there are indications that other
conditioning factors are involved. Low
tocopherol in the roughage or the presence in
the feed of certain antagonists of vitamin E and
selenium may reduce fertility in animals raised
 REVIEW TRACE ELEMENTS AND FERTILITY
or maintained in selenium-deficient areas. The
problem o f selenium-related infertility appears
to be more prominent in sheep than in other
domestic ruminants.
Manganese
Although manganese occurs in most b o d y
tissues in low concentrations, only in extremely
low dietary and tissue concentrations does
deficiency result in manifestations of disturbed
or depressed reproductive function (24, 101).
The actual expression appears to depend on a
variety of factors such as degree and duration
of the deficiency.
While the most detailed and extensive
investigation of the consequences of manganese
deficiency have been with laboratory animals, a
number of studies in cattle have yielded essen-
tially similar results (5, 50). Principal among
these are suppression of estrus, reduction in
conception rates, increased incidence of abor-
tions, and small birth weights. The main clinical
sign of dairy cattle infertility caused by a low
manganese intake is anestrus or irregular return
to estrus, sometimes with extended periods of
anestrus (14, 107). Follicular development is
often poor, delayed ovulation is not infrequent,
and signs of estrus when it does occur are less
obvious. Conception rates of only 35 to 40%
are not unusual (14, 107). Dairy cattle with a
manganese intake of less than 40 ppm in the
feed maintained high fertility when the calcium
and phosphorus content of the diet were
adequate and well balanced (43, 44), but
when the calcium t o phosphorus ratio was high
and dietary manganese low (tess than 40 ppm),
fertility was depressed (45). However, when
manganese content of the diet exceeded 100
ppm, the best breeding performances were with
high calcium to phosphorus ratios. In Scandi-
navia, prophylactic administration of .25 g
manganese per day to dairy cattle increased
fertility (Sanstedt, cited by Kolb (50)).
Related reproductive observations have been
in sheep and goats as well. More services per
conception (2.5 vs. 1.5) were required for ewes
fed a low manganese diet (less than 8 ppm)
than w e r e required for ewes fed a manganese
supplemented diet (39). In female goats manga-
nese deficiency resulted in a poor conception
rate and increased embryonic mortality (5).
In South Australia manganese supplements
1199
increased the number of lambs born to a flock
o f Dorset-Horn ewes in which reproductive
performance previously has declined (27); this
further indicates the necessity for continued
awareness of the possible need for providing the
small grazing ruminant with manganese supple-
ments.
A study on the effect o f a dietary manganese
deficiency (less than 2 ppm) on the male
reproductive system of goats showed no effect
on the libido of adult males. However, there
were decreases in the motility of the sperma-
tozoa as well as reduced sperm numbers in the
ejaculate from both old and young male goats.
Although there was also a high proportion of
abnormal spermatozoa, fertility was not im-
paired. The ejaculate of the manganese-deficient
goats contained less ash, zinc, and copper than
that of controls, but manganese was normal
(31).
Abortions and cystic ovaries are prevalent on
the farms using feeds relatively rich in manga-
nese while incidence of anestrus is relatively
high in dairy herds with a manganese deficiency
(28). Manganese content o f corpora lutea
from cows without ovarian cysts was generally
higher than in luteal tissue from cows with
cystic ovaries although the difference was not
statistically significant (40). In the cortical
stroma, which contains ovarian follicles in
various stages of development and regression,
the manganese concentration was significantly
higher in cows without ovarian cysts than in
those with cysts. The reason for the lower
manganese content in cystic ovaries may be in-
creased degeneration of granulosa cells since
these are rich in mitochondria in which manga-
nese is concentrated (40).
While the precise loci of specific manganese
involvement in reproductive processes remain
unknown, some evidence suggests that manga-
nese plays a role in the activity of certain
endocrine organs. The pituitary and ovary are
relatively rich in manganese (5.9 and 5.2/ag/g).
Ovarian content in particular is sensitive to
dietary deficiencies of manganese. Manganese
dropped by one-third of normal during a 6-mo
imposition of a deficient diet (93). Large
ovarian follicles and corpora lutea of ewes have
taken up radioactively labeled manganese to a
greater extent than other ovarian or extraovarian
reproductive tissues (36). Uptakes also have
differed in relation to the estrus cycle. Maximal
Journal of Dairy Science Vol. 62, No. 8, 1979
1200 HIDIROGLOU
uptake occurred in the corpora lutea and this
was greatest between days 4 and 11 of the
estrus cycle when luteal progesterone production
was greatest. This suggests that manganese is
involved specifically in luteal metabolism
and/or activity (36, 37).
Although manganese is well known as an
activator of enzyme systems in many metabolic
pathways and as a constituent of certain
metalloenzymes (103), it also has been impli-
cated explicitly in the synthesis of sterols, and,
hence, of gonadal hormones (10). Gonado-
tropins such as HCG may influence manganese
transport and modify its availablility to differ-
ent tissues of organs (38).
Conclusions
Manganese is necessary for normal fertility
in ruminants, and a manganese deficiency
depresses conception. Because of the generally
low manganese content of corn silage there is a
need for constant alertness to the possibility of
reproductive failure caused by manganese
deficiency in farm animals whose diets contain
high proportions of corn silage, as well as in
animals kept on land manganese-deficient.
Iodine
The need for iodine for the prevention of
goiter in lambs long has been recognized. Its
involvement in thyroid function and homeo-
stosis through its incorporation with the
thyroid hormones, thyroxine and tri-
iodothronine, is understood reasonably well. It
generally has not been recognized, however,
that iodine-deficiency in ruminants has reper-
cussions in reproduction.
Iodine deficiency during pregnancy impairs
fetal thyroid functions, and abortions are
common in the iodine-deficient pregnant cow.
Normal development of the reproductive organs
and their proper functioning is dependent
on thyroid status. Thyroidectomized dairy
heifers cease to exhibit estrus at regular intervals
(66, 92). An iodine-deficient diet fed to dairy
cows resulted in anovulatory estrus caused by a
disturbance of the thyroid and pituitary func-
tions. This was corrected by the addition of
iodine to the diet (88). For "repeat breeder"
cows on iodine-deficient pasture, conception
rate was improved by adding iodine to their
Journal of Dairy Science Vol. 62, No. 8, 1979
feed (63). In Finland the administration of
iodine to cattle by sublimation resulted in
higher conception rate and lower incidence of
retained placentas than in controls (68).
The beneficial effect of iodine is believed to
involve stimulation of the anterior pituitary
gonodotrophic secretion mediated through the
thyroid gland. Intrauterine injections of iodine
to cows caused distinct lengthening of the
estrous cycle, but it was not ascertained whether
this was physiologic, pharmacologic, or toxic
(32). Rat uterus bound inorganic iodine specifi-
cally (101). Another phenomenon which may
be involved in the lengthening of the estrous
cycle is the effect of iodine on secretion of
thyrotropin-releasing factor which in turn
stimulates prolactin secretion (16). Infertility
associated with iodine deficiency has been in
sheep as well as in cattle. In areas in which a
high incidence of goiter occurs among lambs,
infertility problems in ewes have been alleviated
by provided salt supplements containing .07%
iodine (6).
Subnormal protein-bound iodine (PBI) in
cows has been associated with infertility (2). In
chronic repeat breeders, the average PBI was 37
/lg/liter, significantly below the average of 46
/ag/liter in normal cows. There was a signifi-
cant inverse relation between serum PBI and
interval between first breeding and conception
as well as number of services per conception
(53).
Improved reproductive performance was
associated with higher PBI; for every 10/~g[liter
increase in PBI, there was a decrease in the
service interval of 7.4 days and a decrease of .2
services per conception. Low serum PBI also
have been associated with abortion in cows
(52).
During the estrous cycle in cattle, iodine and
PBI in blood do not change (87). Blood iodine
is depressed significantly by pregnant mare
serum gonadotropin therapy but is not affected
by administration of estrogens (82). In preg-
nant cows PBI was as high as 128/~g/liter (1).
Iodine and PBI concentration of the thyroid
are reduced in cows with cystic ovaries and also
in cows in estrus as compared with pre-estrus
(1). Other observations have suggested influence
of estrogen on thyroid function; it is probable
that the same pituitary factors (gonadotropin)
which stimulate estrogen secretion have some
direct action on the thyroid gland.
 REVIEW TRACE ELEMENTS AND FERTILITY
Conclusions
Iodine influences reproductive functions
because of its vital role in thyroid function.
Thus, reproductive failure in cases of iodine
deficiency are likely to be secondary manifesta-
tions of thyroid dysfunctions resulting in
anestrus of irregular estrus, retained placenta,
abortion, and stillbirth.
Zinc
Zinc has been recognized for several decades
as indispensable for normal growth and health
in animals. Its lack causes various malformations
and has deleterious effects on sexual functions,
particularly in the male animal.
Although naturally occurring zinc deficiency
is rare in livestock, there are numerous reports
of improvements in the reproductive perform-
ance of animals given dietary zinc supplements.
In general, information as to the roles zinc may
play in reproductive processes has been obtained
primarily from studies on experimentally
induced zinc deficiencies.
In South Australia ewes given 140 mg of
zinc weekly produced significantly more Iambs
than did untreated ewes. It is not clear whether
this effect represents a remedy of a zinc defici-
ency. The zinc content of the pasture on
which the flock had grazed varied between a
subnormal 12 ppm dry grass and a normal or
optimal 45 ppm over 2 yr (27). Some workers
(84) have questioned an effect of zinc deficiency
on fertility in ruminants although they have
reported an apparent influence on the sex ratios
of offspring in zinc-deficient animals.
Fertility of sheep has been improved when
both rams and ewes received zinc supplementa-
tion (12, 51). Among the various factors
responsible for the poor fertility of ewes on a
low zinc ration was early embryonic mortality.
The fertilized ovum of ewes fed low zinc diets
does not implant in the uterine muscosa (8,
89). Feeding goats semipurified diets low in
zinc caused low conception rate and reduced
the number of kids per goat (5).
Cows given a zinc supplement had a 23%
higher conception rate than controls. The
ovarian zinc content of the treated cows was
higher than that of controls. Discontinuation of
the supplementation lowered fertility (71).
Zinc concentrations in plasma of nonpreg-
nant heifers at various stages of the estrous
1201
cycle were little changed (25). Zinc in plasma
decreased during parturition and was of greater
magnitude in cows suffering from dystocia (25,
76). Although large transitory depressions of
zinc have been recorded in some ewes with
dystocia, fall in the zinc concentration has not
been comparable to that in cattle (64). Possibly
these changes in zinc are secondary to the
dramatic increases in prostaglandin associated
with parturition, inasmuch as prostaglandins
bind zinc and facilitate its transport (90).
The highest zinc concentration in animals is
in human semen and spermatozoa. The concen-
tration of zinc in the semen of domestic animals
is considerably less and variable among animals
within species (26, 59). Considerable attention
has been given to zinc in the accessory glands
and in the semen of livestock in relation to
fertility. In a study of seminal plasma of bulls
of high and low fertility, the total zinc content
was related inversely to fertility; zinc was
lowest in the semen of highly fertile bulls and
rose as fertility decreased (94). It is difficult to
determine from available data whether high
concentrations of zinc have an adverse effect on
the fertilizing capacity of the semen or are
themselves the consequence of other deleterious
influences.
Zinc deficiency influences spermatogenesis
in the ram. In rams fed a zinc deficient diet (2.4
ppm) atrophy of the seminiferous tubules
accompanied complete cessation of spermato-
genesis (99).
In lambs, a diet with 17 ppm of zinz sup-
ported satisfactory somatic growth but was
insufficient for normal testicular growth and
spermatogenesis compared with lambs fed a
similar diet containing 32 ppm zinc.
Zinc-deficient calves had considerably smaller
testes than normal calves (74), and zinc-deficient
goats had seminiferous tubules which were
smaller than those of controls and contained an
immature germinal epithelium (65). Other
workers have found zinc-deficient goats had
testicular atrophy and lacked libido (70).
It appears that male accessory sex gland is
also dependent on zinc for normal function.
The prostate, in particular, contains a relatively
high concentration of zinc, and its uptake by
this gland is stimulated by testosterone (41)
which regulates prostate activity.
While zinc has been recognized as a cofactor
or constituent of certain enzymes, for example
Journal of Dairy Science Vol. 62, No. 8, 1979
1202 HIDIROGLOU
carbonic amhydrase, activity of zinc-dependent
enzymes in reproductive processes is unknown
(17, 75). It has been postulated (77) that a
possible role of zinc in ram seminal fluid might
be to prevent destruction of spermatozoal DNA
by inhibiting the DNase activity. Zinc is required
for spermatogenesis during the final stage of
maturation and also for maintenance of the
germinal epithelium (99). Although some of
these involvements may be mediated by influ-
ences on pituitary gonadotropin production
(109), follicle stimulating and luteinizing
hormones in serum of zinc-deficient cows are
not altered by zinc supplementation (49).
Conclusions
Zinc deficiency in ruminants, as in other
species, causes a more pronounced impairment
of fertility in the male than in the female. The
effect is severe and appears to be specific to the
final stages of spermatozoa maturation. Testicu-
lar size and concentration of zinc in semen may
be useful indicators of zinc deficiency as one of
the possible causes of reproductive dysfunction
in ruminants.
Iron
In view of the relatively large requirements
for iron and its occurrence in major quantities
in blood and muscle as well as in numerous
enzymes, it is arguable whether iron properly is
considered a trace element. Nevertheless,
iron traditionally has been included among the
essential trace minerals.
While there is no evidence of iron deficiency
ever in grazing sheep or cattle except as a result
of disease or infestation, a number of investiga-
tors have drawn attention to correlations
between the iron content of body fluids and
fertility.
Iron, manganese, copper, and zinc were
significantly higher in the sera of regular
breeders than in repeat breeders (57). Correla-
tions were high between iron and zinc content
of blood and number of inseminations required
per conception (51).
In a study of iron and copper in the accessory
gland fluids and semen of bulls, concentration
of iron was much greater in the spermatozoa
than in the seminal plasma (21). The average
concentration of iron in ampullary fluid and
in semen was 1.08 and .61 mg/ml, whereas in
Journal of Dairy Science Vot. 62, No. 8, 1979
seminal vesicular fluid and seminal plasma it
was .38 and .35 mg/ml. However, no compari-
sons have been made to allow deductions as to
differences related to altered reproductive
function.
Conclusions
Dietary iron is an essential element in
ruminants although its deficiency rarely has
been observed in grazing cattle or ovine. Iron is
abundant in all feeds, and a deficiency in adult
ruminant seems improbable. However, in
certain instances because of low availability of
iron in some roughages, ruminant reproduction
could be affected adversely by iron deficiency.
G E N E R A L REMARKS
Deficiencies of single trace elements rarely
occur in the field; combinations of mineral
deficiencies are much more common. The
problem is complicated further by close inter-
relations between functions o f many trace
elements and various metabolic processes so
that an imbalance in the intake o f one element
may change the requirement for others. For
example, excess calcium added to nonlegume
rations without added trace elements has
caused interference with the estrous cycle and
with ovulation in cattle; when alfalfa hay or
trace elements were added to the diet, fertility
was improved (95). A general elevation of the
mineral content of the diet appears to have
beneficial effects on fertility. By analysis of
feedstuffs fed to cows and blood samples
periodically over several months, trace elements
of the feed were reflected in the blood content,
and this in turn was paralleled with fertility
(104).
This review emphasizes the need for ade-
quately controlled experiments to determine
whether trace-element malnutrition, in itself,
impairs fertility. Little is known about the way
in which malnutrition of trace elements affects
the complex function of the reproductive
system. All metals in this review are involved in
enzymatic functions, and some hormonal
activities appear to be correlated with trace
elements. In future studies, therefore, it may
prove useful to examine the hormonal status of
cattle and sheep in the light of possible defici-
encies in reproductive dysfunction. Improved
assay techniques for hormones and minerals
 REVIEW TRACE ELEMENTS AND FERTI LITY
c o u l d lead to a b e t t e r u n d e r s t a n d i n g o f t h e
interrelationships between reproductive abnor-
12
malities and deficiencies o f specific trace
elements. 13
The r e l a t i o n s h i p b e t w e e n iodine, h y p o -
t h y r o i d i s m , and cystic ovaries in cows and
b e t w e e n zinc a n d s p e r m a t o g e n e s i s are p a r t i c u l a r 14
i n t e r a c t i o n s w h i c h n e e d to be s t u d i e d in detail 15
as d o e s t h e role o f s e l e n i u m in r u m i n a n t fertility.
O t h e r p r o b l e m s inviting f u r t h e r s t u d y include
investigation o f t h e possible i n v o l v e m e n t o f 16
zinc i n t a k e in t h e seasonal quality o f ram s e m e n
and t h e physiological role o f m a n g a n e s e in
s p e r m a t o g e n e s i s o f rams. 17
ACKNOWLEDGMENTS
The a u t h o r is grateful t o D. W. F r i e n d , K. J. 18
J e n k i n s , a n d G. J. Marcus for t h e i r helpful
suggestions in p r e p a r i n g this m a n u s c r i p t .
19
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