Seasonal
food intake and growth
From ASTF Final Report, April
2003 -- Feeds for qiviut production: trace mineral nutrition of muskoxen.
The amount and quality of forage is critical to management of
muskoxen because grass hay provides 97% of the dry matter ingested by
adults. Demands for hay increase in late summer when food intakes increase
by 74% in castrated males between April and August in Fairbanks. The
efficiency of digesting fiber is sustained at high intakes indicating a
remarkable plasticity of ruminal function. High intakes at high
efficiencies late in the summer support rapid fat gains. This seasonal
adjustment in nutrient uptake would allow reproductive females to rapidly
replenish tissues that were depleted during milk production and gain fat
reserves before winter. High food intakes also permit muskoxen to load
nutrients such as copper and zinc that are only present at low
concentrations in the diet. Grass hay is typically low in sodium but high
in potassium. Therefore, high forage intakes are associated with excess
intakes of potassium that exacerbate the low levels of dietary sodium.
Supplements provide 78% of dietary sodium for captive muskoxen and
probably facilitate high food intakes and body mass gains by compensating
for the large urinary loss of electrolytes during autumn.
Seasonal growth of lean tissues and hair (qiviut) alter demands
for nutrients in muskoxen. Young muskoxen mostly gain lean tissues such as
muscle and bone during the first summer and autumn. Consequently, young
muskoxen are only 5% body fat in the middle of winter but soon gain both
lean and fat tissues during the second summer to reach adult fat contents
of approximately 20% body mass before the second winter. Low stores of
subcutaneous fat could make young muskoxen more vulnerable to thermal
stress especially if wet conditions reduce the insulative capacity of
their coats during the first winter. Males grow more rapidly than females
during the second year probably in response to testosterone production.
Rapid growth of males will probably increase their food intakes when
compared with females in the second year. Captive females are often able
to complete gestation in the second winter because adequate reserves of
lean and fat tissues are deposited during the second summer. Breeding of
these young females may however, curtail further growth and impair
delivery of large calves.
Supplemental protein above the concentration of grass hay (10%
crude protein in dry matter) does not increase hair growth or lean gains
in young muskoxen. The excess protein is probably deposited as fat after
eliminating the excess nitrogen. Therefore sustaining high intakes of hay
with 10% protein is probably adequate for non-reproductive animals as long
as supplements are provided to maintain supplies of minerals such as
sulfur, sodium, copper and zinc. Protein supplements of 14% crude protein
(dry matter basis) may be useful for lactating females to maintain sulfur
and nitrogen supplies when deposition of hair, milk and tissues coincide
in late summer.
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Trace mineral
status
From ASTF Final Report,
April 2003 -- Feeds for qiviut production: trace mineral nutrition of
muskoxen.
Adults
Preliminary research indicated that the trace mineral copper was
a limiting nutrient for muskoxen. Sedges and grasses that form the bulk of
the diet of captive and wild muskoxen are typically low in copper (2 to
10ppm). Storage of copper in the liver varies widely in muskoxen from less
than 10 µg/g (wet mass) in wild animals on Banks Island to over 150 µg/g
(wet mass) in captive animals at Fairbanks. Low liver copper correlates
with a low levels of serum copper and the metaloenzyme ceruloplasmin. The
exhaustion of liver copper in adult females is predicted at serum
concentrations of 0.7µg copper/ ml and 7.5 IU ceruloplasmin /L. These
criteria may be used to monitor copper reserves in captive and wild herds
when sampling the liver is not feasible.
Accumulation of copper in adult females probably relies on high
intakes of forage during autumn. Castrated males consume a diet of less
than 5 ppm copper when grass hay (2ppm copper) and supplement (15 ppm
copper) are combined. Liver copper reserves are used to support growth of
the fetus and accumulation of copper in the fetal liver. Although demands
for energy and protein are both high during milk production, the
deposition of copper in milk is low at only 2 to 3 ppm in dry matter.
Supplementation of females with injectable copper does not affect the
concentration of copper in their calves or in their milk.
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Calves
Muskoxen are born with large reserves of copper in the liver
which is stored at concentrations greater (>200 µg/g wet mass) than
that of the maternal liver. Liver copper is apparently used through the
early development of the calf because concentrations of copper decline to
approximately 50µg/g and 25 µg/g at 30 days and 60 days of age
respectively. Calves mainly consume milk for the first 30 days but
progressively shift to other foods such as pelleted rations and forages as
milk production declines.
Weaning is possible at 100 days of age when mothers are placed in
breeding harems with bulls. Calves accompanying their mothers may be
stressed by the herd activity. Wet conditions from rain and poorly drained
pens may also contribute additional stress by increasing the rate of
infection from pathogens in the ground. An increased frequency of diarrhea
during August can occur with or without separating calves from their
mothers at 100d. Recurrent bouts of diarrhea are probably associated with
repeated intestinal infections that ultimately slow growth and
progressively debilitate the calf. Prolonged debilitation could weaken the
immune system resulting in atrophy of lymph glands in the intestine and
mesentery.
Copper is used for several reactions in the immune response.
Copper demands of muskox calves are probably increased by high rates of
exposure to pathogens. Captive calves in Fairbanks suffer slowed growth
and high rates of diarrhea when provided complete diets of 5 ppm copper
without injectable supplements. That is, dietary copper concentrations
that are similar to sedges and grass hays are not adequate for these
captive animals. Injections of copper-gluconate to supplement this diet
reduced frequency of diarrhea and increased the concentration of liver
copper when compared with untreated calves (153 vs. 20 µg/g wet mass).
The problem of inadequate copper supply to calves is confounded
by the transition of the digestive tract and the liver from a diet of milk
to forage. Furthermore, calves may be more susceptible than adults to
other factors such as fungal toxins in hay. Therefore signs of intestinal
infection, poor growth, muscle wasting and low liver copper can be the
result of more than one condition. Provision of approximately 9 ppm copper
to muskox calves did not prevent the onset of these signs even though this
dietary concentration is apparently adequate for development of reindeer
and caribou reared at high animal densities in the same facility. Our
comparison of muskoxen and reindeer born in captivity indicate that the
digestive tract and the liver of reindeer are more developed at birth and
that reindeer mature more quickly than muskoxen. Therefore slow
development may make muskoxen more susceptible to digestive disruption
than reindeer even though both species are born at the same time of year.
A complete diet of 30 ppm copper (dry basis) provided from birth
to 160 days of age was associated with serum concentrations of 1.7 µg
copper/mL and 59 IU ceruloplasmin/mL after 100 d of age. These levels are
considerably higher than the minima associated with exhaustion of liver
copper in adults. This suggest that the diet was apparently adequate to
sustain copper supply even though the animals ultimately succumbed to
repeated bouts of intestinal infection and wasting. High dietary
concentrations of copper may be essential to rearing captive muskoxen
especially when the rate of infection is high and commensurate with the
density of animals.
Long term management of a captive herd may therefore require
restriction of calving areas to pastures that are actively managed to
minimize infection rate by limiting animal numbers, rotation of livestock
and tilling the substrate.
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Feeding
standards
From ASTF Final Report,
April 2003 -- Feeds for qiviut production: trace mineral nutrition of
muskoxen.
Formulated feeds for muskoxen are available from Alaska Pet and
Garden, Anchorage under license agreement with the University of Alaska
Fairbanks. Young muskoxen are provided a complete ration with the typical
composition listed in Table 1. This product is designed to be fed free
choice from birth to approximately 180 days of age when animals should be
transitioned to a diet of grass hay with supplemental rations. The calf
ration and the supplements should not be fed free choice because the
concentrations of trace minerals are high and designed to augment the low
levels in hay. Consequently, free choice feeding of these products could
precipitate toxic accumulations of trace nutrients. The products are not
designed for multi-species feeding, for example, the high concentrations
of copper may be toxic to sheep.
Weekly rates of supplementation for muskoxen are 70 g/kg0.75
body
weight for yearlings and 35 g/kg0.75 body
weight for adults. These weekly rates are plotted against body mass of the
animal in Figure 1. The weekly
ration should be provided in 2 or 3 equal allotments during the week to
provide an even rate of nutrient supply and to minimize rejection, wastage
or over supplementation of some animals in the herd. Supplements can be
changed to low protein formulations in winter to reduce costs of feed.
The largest component of feed intake is grass hay. Predicted
intakes of hay are based on the difference between the rate of
supplementation and the daily dry matter intakes: 45 g/kg0.75 body
mass in winter and 78 g/kg0.75 body mass in summer. Hay intakes are plotted
in Figure 2. Those intakes may
be reduced considerably by consumption of fresh forages.
Table 1. Typical composition of dry matter in the calf ration,
supplements and grass hay provided to captive muskoxen at the Institute of
Arctic Biology, University of Alaska Fairbanks.
|
Component
|
Calf Ration
|
Summer Supplement
|
Winter Supplement |
Grass Hay
|
|
Organic Matter
(g.100g-1)
|
>80
|
>80
|
>80
|
> 90
|
|
Neutral Detergent Fiber
(g.100g-1)
|
< 25
|
< 25
|
< 25
|
60
|
|
Acid Detergent Fiber
(g.100g-1)
|
< 13
|
< 13
|
< 13
|
30 - 35
|
|
Acid Lignin (g.100g-1)
|
< 3
|
< 3
|
< 3
|
3 - 4
|
|
Crude Protein (g.100g-1)
|
16
|
13 - 17
|
> 10
|
6 - 10
|
|
Sulfur (g.kg-1)
|
3
|
3 – 4
|
3 – 4
|
0.6 - 1.2
|
|
Calcium (g.kg-1)
|
1 - 2
|
2 – 3
|
2 – 3
|
2 - 3
|
|
Magnesium (g.kg-1)
|
1
|
1.3
|
1.3
|
0.6 - 0.9
|
|
Potassium (g.kg-1)
|
6
|
< 8
|
< 8
|
10 -12
|
|
Sodium (g.kg-1)
|
1
|
7
|
7
|
0.1 - 0.2
|
|
Copper (mg.kg-1)
|
30
|
30
|
20-30
|
2 - 4
|
|
Zinc (mg.kg-1)
|
100
|
120
|
80-120
|
7 - 11
|
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|