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OCR for page 328
Molybdenum
Molybtlenum (Mo), discovered about 1782, now is a recognized ubiqui-
tous element in the earth's surface and living matter. Molybdenite, the
principal molybdenum ore, is found in close association with tin ore and
is used industrially in the manufacture of various alloys. Present knowI-
edge regarding the biological importance of molybdenum developed, in
large part, from studies of its metabolic Interrelationship with copper.
The true nature of this interrelationship is still being researched.
Several comprehensive reviews on molybdenum exist, including those
of Schroeder et al. (1970) and Underwood (1976~. Aspects of mob
denosis have been described by Ammerman and Miller (1975), Buck et
al. (1973), Clarke and Clarke (1975), Kubota (1976), Ward (1976), and
Poitevint and Nelson (19781.
ESSENTIALITY
A biological requirement for molybdenum was first demonstrated by
Bortels (1930), who found molybdenum to be an essential media nu-
trient for the growth of Azotobacter sp. Subsequently, nitrogen-fixing
bacteria, such as the symbiotic Rhizobia sp. in legume roots, were
shown to require molybdenum (Steinberg, 1936~. Molybdenum-
deficient soils were subsequently recognized, and significant improve-
ments in their yields of pasture legumes were made by appropriate
molybdenum applications.
328
OCR for page 329
Molybdenum
329
When a long recognized severe diarrhea of cattle (tears scours) in
England was shown to be caused by molybdenum toxicosis, attention
turned to the metabolic significance of molybdenum in animals. The
copper-molybdenum interrelationships were revealed when Ferguson
et al. (1938) found molybdenum toxicosis could be controlled by copper
supplementation and when Dick and Bull (1945) showed chronic copper
toxicosis of sheep in Australia could be alleviated by molybdenum
supplementation .
The major biochemical role of molybclenum in animals is currently
believed to be in the formation and activity of xanthine oxidase
(xanthine dehydrogenase), a molybdenum-contain~ng metalloprotein
essential for the metabolic degradation of punnes to uric acid (DeRenzo
et al., 1953; Richert and Westerf~eld, 1953~. Xanthine oxidase is present
in microorganisms, animal tissues, and milk, especially cow's milk.
Aldehyde and sulfide oxidases are also molybdenum-dependent en-
zymes present in animal tissues.
METABOLISM
Most dietary forms of molybdenum, except molybdenite (MoS2), are
absorbed from the gastrointestinal tract, but the rates of absorption and
routes of excretion may differ with species (Underwood, 1977~. In
swine, peak blood levels of molybdenum occur within 4 hours after an
oral dose of 99molybdenum, and the urinary tract is the main excretory
route for absorbed molybdenum. In cattle, peak blood levels from an
equivalent dose of 99molybtienum are not reached until 96 hours post-
administration, and the main route of molybdenum excretion is via the
feces (Bell et al., 19641. Peak plasma molybdenum levels in cattle fed
100 ppm supplemental molybdenum in cottonseed meal for 12 months
were not reached until 7 months after the start of the feeding trial
(Lesperance and Bohman, 1963~. Absorbed molybdenum is also ex-
creted via the milk from cattle and sheep in proportion to the levels of
orally or parenterally administered molybdenum.
The rates of absorption, retention, and excretion of molybdenum are
inversely related to the level of dietary inorganic sulfate. In sheep, for
instance, increasing the dietary sulfate from 0.1 to 0.3 percent in a diet
supplemented with 10 mg molybdenum per day decreased the molybde-
num retention from 37 to 4 percent. A working hypothesis for the effect
of sulfate on molybdenum retention is that sulfate inhibits membrane
transport of molybdenum, thus decreasing absorption of molybdenum
in the intestine and decreasing reabsorption of molybdenum by the
OCR for page 330
330 MINERAL TOLERANCE OF DOMESTIC ANIMALS
renal tubules (Dick, 1956b). Other factors influencing molybcIenum
metabolism, in addition to copper and sulfate, include dietary levels of
manganese, zinc, iron, lead, tungstate, ascorbic acid, methion~ne
cyste~ne, and protein.
SOURCES
Naturally growing herbage usually reflects the molybdenum content of
the soil. Concentrations of molybdenum in normal herbage often range
from 0.1 to 3 ppm (Underwood, 1977) on a dry weight basis. The
molybdenum in herbage is present as water-soluble sodium and ammo-
nium molyb~ate and as insoluble molybdenum oxide (MoO3), calcium
molybdate (CaMo04) and molybdenum sulfide (MoS21. Only MoS2 am
pears to be very poorly absorbed. Plants growing on soils ~ndustriaRy
contaminated with molybclenum or containing naturally high levels of
molybdenum have contained up to 231 ppm molybclenum (Gardner and
Hag-Patch, 1962~.
TOXICOSIS
LOW LEVELS
Manifestations of molybdenum toxicosis in cattle include diarrhea,
anorexia, achromotnchia, and posterior weakness. Natural foodstuffs
containing up to 6.2 ppm molybclenum were found by Smith et al. (1975)
to be associated with bone malformations In calves. Cunningham et al.
(1953) have reported that natural forages containing 25.6 ppm molyb-
denum were responsible for diarrhea, emaciation, anemia, achromotri-
chia, and even death in several age-groups of cattle. Huber et al. (1971)
and Vanderveen and Keener (1964) reported that molybdenum levels
up to 100 ppm had no effect in cattle, yet Gardner and Hall-Patch (1962)
found cattle grazing industr~aNy contaminated forages containing BS
ppm molybdenum developed diarrhea and locomotor disturbances. The
achromotrichia, diarrhea, and reduced weight gains have also been
demonstrated in cattle consuming molybdenum at 100 ppm
(Lesperance and Bohman, 1963), at 17~300 ppm(Hubere' al., 1971),
at 400 ppm (Cunningham et al., 1953), and at 2.34 g per day (Bntton and
Goss, 1946~. Molybdenum toxicosis has been observed in young lactat-
ing cattle consuming as little as 40 ppm molybdenum when the diets
contained 0.3 percent sulfate (Vanderveen and Keener, 19641. These
.
OCR for page 331
Molybdenum
331
authors also reported that 200 ppm molybdenum, In conjunction with
0.3 percent dietary sulfate, produced posterior paresis in the young
lactating cattle. It appears that 100 200 ppm dietary molybdenum are
required to significantly increase the molybdenum content of milk
(Cunningham et al., 1953~. Cattle with molybdenosis have also been
reported to have an increased incidence of partunent hemogiobinuria
(Goold and Smith, 1975~. Thomas and Moss (1951) have observed de-
creased libido and testicular degeneration In young bulks fed 1-2 g
sodium molybdate dibydrate dady for a period of 120 days.
The effects of molybdenum on hepatic copper levels depend upon the
levels of dietary molybdenum and copper (Lesperance and Bohman,
19631. Levels of molybdenum up to 40 mg per day tend to decrease
hepatic copper levels, while dietary molybdenum levels beyond 40 mg
per day may alter hepatic copper levels very little (Ammerman and
MiDer, 19751. Dick (1956a) has reported that dietary copper levels of
~10 ppm protect cattle against dietary molybdenum levels of approxi-
mately 5 - ppm.
Sheep appear more resistant to molybclenosis than cattle and tolerate
plasma molybdenum levels of 0.1~.2 mg/dI, or approximately 2~0
times the normal plasma molybdenum levels, without affecting cerulo-
plasmin levels. This is true providing the dietary sulfate intake is about
0.1 percent (Dick, 1953a; Suttle, 19751. The manifestations of
molybdenum-induced, secondary hypocuprosis include reduced crimp
and pigmentation of wool, anemia, alopecia, and reduced weight gains.
Neonates born to hypocupremic dams exhibit enzootic ataxia
(swayback), a demyelinating disease that may also be accompanied by
blindness. Dick and Bull (1945) found young ewes that consumed
1~100 mg molybdenum per day as ammonium molybdate for a 6-
month period had significantly lower than normal liver copper levels
regardless of the dietary copper level. Thus, the daily intake of mob
denum, which will alter liver copper levels in sheep, approximates 10
ma. In cases of elevated liver copper levels, Ross (1970) found that 100
mg molybdenum per day for 12 weeks caused a significant reduction in
hepatic copper levels in sheep. Sheep fed 120 mg molybdenum and 7.4
g sulfate daily for 20 months were found to have reduced hemoglobin
levels, reduced levels of copper in the wool, and increased levels of
albumin-bound copper in blood (Bingley, 19741. Normal plasma copper
levels were noted at molybdenum intakes of 12 mg/day.
Goodrich and Tillman (1966) investigated the effect of 2 and 8 ppm
molybdenum on lambs receiving either 10 or 40 ppm copper and either
0.1 or 0.4 percent sulfate. Eight parts per million molybdenum elimi-
nated the detrimental effects of the high sulfate level upon rate of gain
OCR for page 332
332 MINERAL TOLERANCE OF DOMESTIC ANIMALS
and feed efficiency and also reduced liver copper levels. The latter
effect was reversed by the addition of 40 ppm copper. A direct effect
of molybdenum upon rumen sulfate levels has been demonstrated by
Gawthorne and Natler (1976) in rumen fistulated sheep. Marcilese et al.
(1976) suggested a higher turnover rate of ceruloplasmin in cattle than
in sheep accounted for the difference in molybdenum tolerance be-
tween these two species.
Other ruminants are even more tolerant of molybdenum than sheep
or cattle. For instance Nagy et al. (1975) found that mule deer tolerated
up to 1,000 mg molybdenum per day without clinical signs. Daily
molybdenum intakes by the mute deer of 2,500 to 5,000 mg caused
diarrhea, and 5,000 to 7,500 mg caused anorexia. Appetite returned as
soon as dietary molybdenum supplementation ceased.
Gipp et al. (1967) and Kline et al. (1973) have demonstrated little to
no effect of 26 to 50 ppm molybdenum upon swine growth in the
presence of supplemental copper and sulfate, while Davis (1950) re-
ports no apparent effect of 1,000 ppm molybdenum in growing swine.
Standish et al. (1975) have fed 1,500 ppm molybdenum to growing
swine for 69 days in the presence of 17.S ppm copper. These levels of
molybdenum and copper caused a marked reduction in the rate of gain
of these swine; however, the effects were reversible by 0.4 percent
sulfate. High molybdenum levels in swine diets tent! to promote copper
storage in liver and kidney in contrast to an opposite effect in rumi-
nants.
Avian species appear comparable to rodents in their susceptibility to
molybdenum. Kratzer (1952) demonstrated a slight growth inhibition in
young chickens fed 200 ppm molybdenum and a 25 percent growth
inhibition in poults fed 300 ppm molybdenum. Davies et al. (1960) fed
molybdenum to young chicks at levels ranging from 500 to 8,000 ppm.
The effects ranged from growth depression and anemia at the low levels
to 61 percent morality at the highest level. Arthur et al. (1958) also
induced anemia in young birds fed 4,000 ppm molybdenum for ~ weeks.
Lepore and Miller (1965) indicated 500 ppm molybdenum in laying hens
caused decreased hatchability, and 1,000 ppm molybdenum caused
decreased egg production. Davies et al. (1960) have also indicated that
ammonium molybciate is more toxic for birds than sodium molybdate.
Horses seem resistant to molyb~ienosis, for they can graze, without
apparent problems' the same pastures that are known to cause diarrhea
in cattle. However, clinical cases of rickets in foals and yearlings have
been thought to be due to molyWenosis from pasture or dam's mink
(Walsh and O'Moore, 1953~.
Arrington and Davis (1953) have fed up to 4,000 ppm molybdenum to
OCR for page 333
Molybdenum 333
rabbits consuming a basal diet containing approximately 16.4 ppm
copper. Rabbits fed 1,000 or more ppm molybclenum experienced
anorexia, loss of weight, alopecia, a slight dermatosis, anemia, splayed
front legs, and premature deaths. Manifestations of molybdenum toxi-
cosis became apparent within 4 weeks in young rabbits and after a
longer period in older rabbits. The toxic manifestations of molyWenu~n
were alleviated by 200 ppm copper for at least a 4-month period. Molyb-
denosis in rabbits may decrease phosphorus absorption, increase phos-
phorus excretion, and result in a r~cketic syndrome.
The tolerance of rats for dietary molybclenum is, as with other spe-
cies, dependent upon the dietary levels of copper and sulfate. Gray and
Daniel (1964) found young copper-deficient rats experienced reductions
In growth rate, liver copper, and blood hemoglobin levels when fed
1~1,000 ppm molybdenum. Dietary copper supplementation for these
rats at 3 ppm caused their growth rate, liver copper, and blood hemo-
gIob~n levels to return to normal. Miller et al. (1972) reported reduced
growth rate of young rats fed 100 ppm molybdenum could be prevented
by SO4 supplemention. The reports of Whanger and Weswig (1970),
Gray and ElBis (1950), Gray and Daniel (1954), Halverson et al. (1960),
Compere et al. (1965), and Nielands et al. (1948) indicate that dietary
molybdenum levels in excess of 500 ppm in rats impair growth, increase
blood and liver copper levels, decrease ceruloplasmin, and increase
tissue molybdenum levels. In general, all these effects are usually aBe-
viated by copper and/or sulfate supplementation.
In guinea pigs fed 100 ppm molybdenum in the presence of adequate
copper, Smith and Wright (1975) found a tncholoracetic acid-~nsoluble,
copper-molybdenum complex in plasma, which they felt accounted for
the absence of a significant elevation in liver molybdenum during
molybdenosis in this species.
HIGH LEVELS
Very few molybdenum toxicity studies In which sufficient molybdenum
was used to cause death of the animals have been reported. Davies et
al. (1960) found 6,000~,000 ppm molybdenum as sodium or ammonium
molyb~ate caused 30 60 percent morality in a Week period in grow-
ing chickens, and the ammonium molybdate appeared to be the more
toxic. Robitaille and Bilek (1976) found the Ado for molybdenum in tank
water for fish to be 7,340 ppm for trout in 96 hours. The ~D50 for oral
ammonium molybdate in guinea pigs is reported to be 2.2 g/kg of body
weight, and the MAD of sodium molybdate intraperitoneally in rats is 290
mg/kg (Stecher et al., 19681.
OCR for page 334
1
334 MINERAL TOLERANCE OF DOMESTIC ANIMALS
FACTORS INFLUENCING TOXICITY
As has been implied, the effects of excess molybdenum are essentially
those of copper deficiency. The integumental changes, including rough
hair coat, achromotrichia, and loss of crimp in the wool, are related to
a deficiency of the copper~ependent enzyme, tyrosinase. The anemia
of molybdenum toxicosis is related to a deficiency of a second copper-
dependent enzyme, ferroxidase. The skeletal and/or colIagenous mani-
festations of molybdenum toxicosis are also related to a deficiency of
a third copper~ependent enzyme, dopamine ,l3 hydroxylase. It is also
probable that the general growth retardation and anorexia associated
with molybJenosis may relate to deficiencies of a fourth copper-
dependent enzyme, cytochrome C oxidase.
Inorganic sulfate supplements appear to reverse all the manifesta-
tions of molybdenosis except the increased copper storage by the liver.
There is also some indication that the effects of molybdenum on copper
levels of milk, ceruloplasmin, albumin, and urine are not reversed by
sulfate. The apparent effects of molybdenum are also influenced by
manganese, zinc, iron, lead, tungstate, ascorbic acid, methionine,
cysteine, protein, and alkalinity of soils. The bases for many of these
interactions are yet unexplained.
Miller et al. (1972) present paradoxical data suggesting that ruminal
processes decrease the biological availability of molybdenum. Gaw-
thorne and Nader (1976) report that molybdenum and sulfate must be
supplied in the diets simultaneously before copper concentration in
ruminant liver is decreased. In rodents, endogenous sulfur from the
metabolism of sulfur-containing amino acids appears not to function as
dietary sulfate in altering the effects of molybdenum as occurs in rumi-
nants (Mills et al., 1958; Cook et al., 1966~.
TISSUE LEVELS
Solid tissue, blood, and milk levels of molybdenum are readily altered
by changes in dietary molybdenum levels, and the magnitude of the
tissue response to elevations in dietary molybdenum depends on con-
comitant inorganic sulfate levels (Dick, 1953a), tungstate (Davies et al.,
1960), and copper. The concentrations of molybdenum in liver of
animals on normal diets range from 2 to 4 ppm on a dry matter basis and
may be as high as 30 ppm if the animals were consuming high levels of
molybdenum (Gray and Daniel, 19641. Renal molybdenum concentra-
tions approximate 50 percent of the liver concentrations of molybde-
OCR for page 335
Molybdenum
335
num (Underwood, 1977), and other tissues in a declining order of usual
molybdenum concentration are: spleen, lung, brain, bone, and muscle.
The total quantity of molybdenum in a skeleton is greater than 50
percent of the total molybdenum in the body (Dick, 1969~. The molyb-
denum levels of whole blood of sheep and cattle on low molybdenum
diets can range from 1 to 6 ,ug/41 (Beck, cited by Underwood, 19771.
The concentration of molybdenum in milk of cattle fed standard diets
ranged from 18 to 120 ,u g/ 1 with a mean of 73 ,u g (Archibald, 1951) and
is primarily associated with the nonlipid fraction of milk, specifically,
the xanthine oxidase. Extreme levels of molybdenum in excess of 1
ppm in milk have been associated with high molybdenum pastures.
MAXIMUM TOLERABLE LEVELS
Estimates of the maximum tolerable levels for molybdenum for several
species are presented in Table 25. These range from levels of 5 to 10
ppm, which have been weakly associated with impaired bone develop-
ment in young horses and cattle, respectively, to very high tolerance
levels approximating 1,000 ppm for swine. It must be emphasized that
substantially higher levels of molybdenum would be tolerated in the
presence of adequate copper and inorganic sulfate.
SUMMARY
Molybdenum is an essential trace element and a component of xanthine
oxidase that is important in purine metabolism. The soils and resulting
herbage in some geographic areas have relatively high molybdenum
levels that account for a regional incidence of molybdenosis in live-
stock. This disease is essentially a secondary copper deficiency mani-
fested by diarrhea, anorexia, depigmentation of hair or wool, neuro-
logic disturbances, and premature death. A wide variation in the
apparent susceptibility of various livestock species to molybdenum
toxicity is due to variations in concurrent dietary levels of copper, zinc,
sulfur, silver, cadmium, and sulfur-containing amino acids. The wide
tolerance limits range from 6.2 ppm in growing cattle to approximately
1,000 ppm in adult mule deer.
OCR for page 336
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OCR for page 342
342 MINERAL TOLERANCE OF DOMESTIC ANIMALS
REFERENCES
Ammerman, C. B., and S. M. Miller. 1975. Molybdenum in ruminant nutrition: A review.
(Unpublished).
Archibald, J. G. 1951. Molybdenum in cows' milk. J. Dairy Sci. 34:1026.
Arrington, L. R., and G. K. Davis. 19S3. Molybdenum toxicity in the rabbit. J. Nutr.
51:295.
Arthur, D. I., I. Motzok, and H. D. Branion. 1958. Interaction of dietary copper and
molybdenum in rations fed to poultry. Poult. Sci. 37:1181.
Bell, M. D., G. B. Diggs, R. S. Lowrey, and P. L. Wright. 1964. Comparison of Mo99
metabolism in swine and cattle as affected by stable moly~date. J. Nutr. 84:367.
Bingley, J. R. 1974. Effects of high doses of molybdenum and sulphate on the distribution
of copper in plasma and in blood of sheep. Aust. J. Agric. Res. 25:467.
Bortels, H. 1930. Molydan als Katalysator bei der biologischen Stockstoffbundung.
Arch. Mikrobiol. 1:333.
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Representative terms from entire chapter:
molybdenum levels
