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INVITEd REVIEW: Mineral and vitamin nutrition in ruminants 1

Article  in  The Professional Animal Scientist · April 2014

DOI: 10.15232/S1080-7446(15)30103-0

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 The Professional Animal Scientist 30 (2014):180–191
©2014 American Registry of Professional Animal Scientists

IvitaminRnutrition: Mineral
NVITEd and EVIEW
in ruminants 1
J. W. Spears,*2 PAS, and W. P. Weiss,† PAS
*Department of Animal Science, North Carolina State University, Raleigh 27695-7621; and
†Department of Animal Sciences, The Ohio State University, Wooster 44691

ABSTRACT (in feedstuffs and water) on performance UPDATE ON MINERALS

Requirements and factors that affect
dietary requirements for several trace
minerals and vitamins in ruminants are
poorly defined. Most B vitamins and
vitamin K are believed to be synthesized
by bacteria in the rumen in adequate
amounts to meet the requirement of the
animal. Nonetheless, several studies in-
dicate that supplementing high-producing
dairy cows with approximately 20 mg/d
of biotin can decrease hoof lesions and
lameness and in many instances increase
milk yield. In vivo synthesis of choline
is also likely limiting milk production in
early lactation. The vitamin E require-
ment for optimal immunity and health
in receiving cattle and transition dairy
cows continues to be an area of inter-
est, with responses to supplementation
varying. Requirements for vitamin D are
being reevaluated in light of its poten-
tial effects on immunity. Studies clearly
indicate that P requirements of cattle
are less than those recommended 20 yr
ago. Because of increased use of etha-
nol by-product feeds that are high in S,
considerable research has been conducted
to determine the effects of high dietary S
1
Presented at the American Registry of
Professional Animal Scientists (ARPAS)
Symposium: Applied Nutrition of
Ruminants—Current Status and Future
Directions, July 10, 3013, ADSA/ASAS Joint
Annual Meeting, Indianapolis, Indiana.
2
Corresponding author: jerry_spears@ncsu.
edu
and incidence of polioencephalomalacia.
Requirements for certain microminerals
are affected by antagonists. Sulfur and
Mo are important Cu antagonists that
can greatly affect dietary Cu bioavail-
ability. High dietary Fe, when present in
a bioavailable form, is a potent Cu and
Mn antagonist. Recent research suggests
that NRC recommendations for Co and
Mn might underestimate requirements.
An estimated requirement for Cr should
be considered in the future based on re-
sponses to Cr supplementation in cattle.
Key words: beef cattle, dairy cattle,
mineral, trace mineral, vitamin
INTRODUCTION
Minerals and vitamins are required
for the normal functioning of essen-
tially all metabolic processes in rumi-
nants. Dietary deficiencies or excesses
of certain minerals and vitamins can
result in substantial economic losses
in animal productivity. Vitamin and
mineral requirements for beef and
dairy cattle were last published by
the NRC (2000, 2001). Since that
time, considerable research has been
published dealing with minerals and
vitamins in cattle nutrition. This
paper will not attempt to cover all
minerals and vitamins but will focus
on minerals and vitamins for which
new information is available that
could affect future published require-
ments or recommendations.
FOR RUMINANTS
Phosphorus has received the most
attention of the macrominerals in
regard to defining requirements.
Limited research evaluating the
requirements of other macrominerals
has been published in the past 15 yr.
The concentration of S in many cattle
diets has increased in recent years as
a result of increased use of by-product
feeds that are high in S. High dietary
S can decrease DMI and ADG and
lead to polioencephalomalacia (PEM;
NRC, 2005). Considerable research
has been conducted to define upper
levels of dietary S that can be toler-
ated by cattle without adversely af-
fecting performance or health (Loner-
agan et al., 2001; Spears et al., 2011;
Sarturi et al., 2013).
It is important to note that trace
mineral requirements listed in the
beef and dairy NRC publications are
for total dietary concentrations (diet
plus supplemental), not supplemental
concentrations; however, requirements
of some trace minerals can be affected
by dietary antagonists, which could
affect requirements. Recent literature
with cattle related to responses to
supplementation of control diets with
chromium, cobalt, copper, manganese,
and zinc will be covered in this paper.
Phosphorus
Based on new research findings, P
requirements for lactating cows were

decreased considerably, whereas P re-
quirements for dry cows and growing
heifers were decreased slightly in the
last dairy NRC (2001) compared with
previous NRC editions. Phosphorus
requirements (percentage of dietary
DM) were approximately 0.36% dur-
ing early lactation, 0.22% for dry
cows, and 0.28% (6 mo old) to 0.18%
(18 mo old) for growing heifers (NRC,
2001). Research findings published
since 2001 suggest that P require-
ments listed in the dairy NRC are
adequate to support normal growth,
milk production, reproduction, and
health.
Milk yield, measures of ovarian
activity, and reproductive perfor-
mance did not differ among Holstein
cows fed 0.35 or 0.47% P from calving
through 44 wk of lactation (Tallam
et al., 2005). In a study conducted in
the United Kingdom over 4 succes-
sive lactations, lactating dairy cows
fed 0.36% total dietary P had similar
feed intake, milk yield, milk composi-
tion, fertility, SCC, bone ash, and
incidence of lameness and mastitis as
cows receiving 0.42 (summer period)
to 0.49% P (winter period; Ferris et
al., 2010a,b). Plasma P concentrations
were less and bone P concentrations
were slightly less in cows fed 0.36%
P. In a relatively short-term study,
varying P from 0.34 to 0.43 or 0.52%
P did not affect immune response of
lactating dairy cows (Mullarky et al.,
2009). Bone development in grow-
ing Holstein and Holstein × Jersey
crossbred heifers did not differ among
heifers fed 0.29 versus 0.39% P from 4
to 21 mo of age (Esser et al., 2009).
Studies with finishing cattle sug-
gest that P requirements are less
than those recommended in the beef
NRC (2000). Erickson et al. (1999,
2002) reported no improvements in
performance, carcass characteristics,
or bone ash from P supplementation
of finishing cattle fed diets contain-
ing 0.14 to 0.16% P. Experimental
diets used in these studies were higher
in fiber than diets typically used for
finishing cattle in the United States.
Phosphorus is known to function
in acid–base balance, and studies
are needed to determine the role of
Mineral and vitamin nutrition
dietary P in controlling acidosis in
cattle fed high-concentrate diets. The
major route of P excretion changes
from feces in ruminants fed high-fiber
diets to the urine when high-concen-
trate diets are fed (Scott and Buchan,
1985). Reed et al. (1965) found a
positive relationship between the P
content and titratable acidity of urine
in steers fed high-concentrate diets.
Phosphorus requirements of grow-
ing beef cattle and beef cows graz-
ing pasture are not well defined, and
responses to P supplementation have
been variable (Karn, 2001). Ternouth
et al. (1996) estimated P requirements
of growing cattle in Australia grazing
tropical pasture or being fed barley
straw–based diets. In cattle gaining
1.0 kg/d, estimated P requirements
decreased from 0.34% at 100 kg of
BW to 0.18% at 400 kg BW. A series
of long-term P supplementation stud-
ies under range conditions have been
conducted at the USDA Northern
Great Plains Research Laboratory in
Mandan, North Dakota (Karn, 1995,
1997). In these studies, the P content
of harvested roughages fed during the
winter and pastures grazed during
the growing season ranged from 0.10
to 0.24% P on a DM basis. In a 5-yr
study, providing beef cows supplemen-
tal P (4 to 6 g/d) twice a week in a
grain carrier increased calf weaning
weights in 3 of the 5 yr (Karn, 1997).
Phosphorus supplementation did not
affect percentage of cows calving or
average calving date. Weight gain
of growing Hereford and Hereford ×
Angus replacement heifers was not
affected by supplementing 4 to 6 g
of P/d during a 462-d study (Karn,
1995). In a subsequent study with
Herford × Simmental replacement
heifers, P supplementation increased
gain by approximately 0.04 kg/d
(Karn, 1995).
Chromium
Chromium functions by potentiating
the action of insulin in insulin-sensi-
tive tissues. Studies showing improve-
ments in immunity, glucose clearance,
and milk production as a result of Cr
supplementation of cattle diets were
181
discussed in the last beef NRC (2000)
and dairy NRC (2001); however, both
publications indicated that informa-
tion was not sufficient to estimate Cr
requirements of cattle. Considerable
research has been published with Cr
in the past 12 yr that may allow NRC
committees to estimate Cr require-
ments of beef and dairy cattle. The
FDA issued a regulatory discretion
letter in 2006, which permitted the
use of Cr propionate as a source of
supplemental Cr in cattle diets, at
levels up to 0.5 mg of Cr/kg of DM.
Studies have indicated that Cr
supplementation to diets of growing
cattle (Sumner et al., 2007; Spears et
al., 2012), dairy cows (Hayirli et al.,
2001), and beef cows (Stahlhut et al.,
2006a) can increase insulin sensitivity
following i.v. glucose administration.
A recent dose-titration study exam-
ined glucose and insulin metabolism
in growing heifers supplemented with
0, 3, 6, or 9 mg of Cr/animal daily
(Spears et al., 2012). These daily
doses corresponded to 0 (control diet
analyzed 0.20 mg of Cr/kg of DM),
0.47, 0.94, and 1.42 mg of supplemen-
tal Cr/kg of DM. All levels of supple-
mental Cr increased insulin sensitivity
based on lower insulin concentrations
and lower molar ratios of insulin to
glucose following i.v. glucose adminis-
tration. Results of this study indi-
cated that Cr requirements of growing
heifers, based on insulin sensitivity,
can be met by supplementing 0.47 mg
of Cr/kg of DM (Spears et al., 2012).
Responses to supplemental Cr might
be greatest under conditions that
decrease insulin sensitivity. It is well
documented that insulin resistance
occurs in late gestation and continues
during early lactation in both dairy
and beef cows (Sano et al., 1993).
Several studies (Hayirli et al., 2001;
McNamara and Valdez, 2005; Smith
et al., 2005) have reported that Cr
supplementation of dairy cow diets
during late gestation and early lacta-
tion increased DMI and milk produc-
tion during early lactation. Chromium
supplementation has improved repro-
ductive performance and decreased
postpartum BW loss in young beef
cows (Aragon et al., 2001; Stahlhut et
182
al., 2006b). Pregnancy rate in dairy
cows has tended to be improved by
Cr supplementation in some studies
(Bryan et al., 2004; Soltan, 2010).
Hormones produced during stress
can decrease insulin sensitivity and
also increase urinary excretion of
Cr in humans and rats (Spears and
Trivedi, 2013). In dairy cows exposed
to heat stress, Cr supplementation
increased DMI and milk production
(Al-Saiady et al., 2004; An Qiang et
al., 2009; Soltan, 2010). Increased re-
lease of cortisol during stress is known
to suppress a variety of immune
responses, and Cr supplementation to
diets of stressed cattle has decreased
blood cortisol concentrations (Spears
and Trivedi, 2013). Calves supple-
mented with 0, 0.1, 0.2, or 0.3 mg of
Cr/kg of DM exhibited a linear im-
provement in ADG and G:F during a
56-d receiving period (Bernhard et al.,
2012). Morbidity also tended to be
decreased by Cr supplementation in
this study. In addition, supplementing
0.2 mg of Cr/kg of DM decreased BW
loss in steers following an i.v. lipo-
polysaccharide challenge (Bernhard et
al., 2012).
Cobalt
Cobalt functions as a component of
vitamin B12. Ruminal microorganisms
are capable of synthesizing vitamin
B12 from dietary Co. When dietary
Co is adequate in the diet, ruminal
synthesis of vitamin B12 is generally
sufficient to meet the requirement of
the host animal. In the beef (NRC,
2000) and dairy NRC (2001) Co
requirements were estimated at 0.10
and 0.11 mg/kg of DM, respectively.
Considerable research has now been
published indicating that previous
recommendations for Co are prob-
ably too low. In Germany, long-term
studies (Schwarz et al., 2000; Stangl
et al., 2000) were conducted with
Simmental males to define dietary Co
requirements based on animal perfor-
mance and vitamin B12 concentrations
in plasma and liver. In these studies,
cattle were fed corn silage ad libitum
and 2.5 kg/d of an energy-protein
supplement for 280 d. Graded levels
Spears and Weiss
of Co were supplemented to the diet
to provide total dietary Co concentra-
tions ranging from 0.07 to 0.69 mg/
kg of DM. Cattle fed the control diet
(0.07 mg of Co/kg) had lower ADG,
ADFI, and carcass weights at slaugh-
ter than animals supplemented with
Co (Schwarz et al., 2000). Cobalt re-
quirements, based on maximal plasma
and liver vitamin B12 concentrations,
were estimated to be 0.26 and 0.24
mg/kg of DM, respectively (Stangl et
al., 2000). Based on regression analy-
sis, Co requirements were determined
to be 0.12 mg/kg of DM for maximal
BW gain and 0.16 to 0.18 mg/kg for
maximal feed intake (Schwarz et al.,
2000).
Two studies (Tiffany, 2003; Tif-
fany et al., 2003) have examined Co
requirements of growing and finishing
Angus and Angus-crossbred steers.
Control diets in these studies ana-
lyzed 0.04 to 0.05 mg of Co/kg of
DM, and Co was supplemented at
concentrations of 0, 0.05, 0.10, and
1.0 mg/kg of DM. Performance was
not affected by dietary Co during the
growing phase in either study; how-
ever, plasma B12 concentrations were
greater in Co-supplemented cattle
by d 56 of the growing phase. Cobalt
supplementation to the control diet
during the finishing phase increased
ADG and ADFI in both studies. Feed
intake was further increased when
supplemental Co was increased from
0.10 to 1.0 mg/kg of DM in one study
(Tiffany et al., 2003), but increasing
supplemental Co above 0.05 (total
diet Co of 0.10 mg/kg) did not signifi-
cantly increase ADG in either study.
Vitamin B12 concentrations in
plasma, liver, and ruminal fluid were
greatly increased by supplemental
Co in finishing cattle (Tiffany, 2003;
Tiffany et al., 2003). Increasing
supplemental Co from 0.10 to 1.0
mg/kg of DM also greatly increased
plasma and ruminal fluid B12 concen-
trations and moderately increased
liver vitamin B12. These results clearly
indicate that increasing supplemen-
tal Co above 0.10 mg/kg (0.15 mg/
kg of total Co) increases vitamin B12
status in finishing cattle; however,
the concentration of vitamin B12 in
plasma and liver required for normal
metabolic processes has not been
determined. Methylmalonyl CoA
mutase is an important vitamin B12
enzyme in ruminants that is involved
in the metabolism of propionate to
succinate, as it catalyzes the conver-
sion of l-methylmalonyl CoA to suc-
cinyl CoA. If vitamin B12 is deficient,
methylmalonic acid (MMA) increases
in plasma because of methylmalonyl
CoA not being efficiently converted to
succinyl CoA. Plasma MMA concen-
trations were elevated in steers fed
the control diet containing 0.05 mg of
Co/kg of DM (Tiffany, 2003). Cobalt
supplementation decreased plasma
MMA concentrations, and plasma
MMA concentrations were decreased
when supplemental Co was increased
from 0.05 to 0.10 mg/kg of DM; how-
ever, increasing supplemental Co from
0.10 to 1.0 mg/kg did not further
decrease plasma MMA concentrations.
Collectively, these results suggest that
the dietary Co (diet plus supplemen-
tal) requirement of finishing cattle fed
corn-based diets is approximately 0.15
mg/kg of DM. Limited research indi-
cates that finishing cattle fed barley-
based diets might have a greater Co
requirement than those fed corn-based
diets (Tiffany and Spears, 2005).
Increasing dietary Co in lactating
dairy cows fed control diets contain-
ing 0.19 (Kincaid and Socha, 2007),
0.37 (Kincaid et al., 2003), or 1.0 mg
of Co/kg of DM (Akins et al., 2013)
has not affected DMI, milk yield,
milk composition, or plasma or serum
vitamin B12 concentrations. Cobalt
supplementation has increased (Akins
et al., 2013) or tended to increase
colostrum and milk vitamin B12 con-
centrations.
In addition to mammalian metabo-
lism, Co also affects metabolism of
certain ruminal microorganisms. In
some bacteria, the pathway involved
in the conversion of succinate to
l-methylmalonyl CoA and then to
propionate is the reverse of that found
in the liver of ruminants and involves
vitamin B12-dependent methylmalo-
nyl CoA mutase (Tiffany and Spears,
2005). Kennedy et al. (1991) showed
that ruminal and plasma succinate

concentrations were greatly increased
in lambs fed Co-deficient barley diets.
Cobalt supplementation to corn- or
barley-based diets that were low in
Co increased molar proportion of
propionate in ruminal fluid and de-
creased the acetate:propionate ratio in
finishing steers (Tiffany et al., 2003;
Tiffany and Spears, 2005).
Copper
It is well documented that Cu
requirements of ruminants are greatly
affected by dietary concentrations of
Mo, S, and Fe. The beef NRC (2000)
indicated that 10 mg of Cu/kg of DM
should be adequate for beef cattle if
their diets do not exceed 0.25% S and
2 mg of Mo/kg. In the dairy NRC
(2001) Cu requirements were calcu-
lated assuming diets contained 0.25%
S and 1 mg of Mo/kg. Estimated Cu
requirements for dairy cattle were
approximately 11, 12 to 18, and 10
mg/kg of DM for lactating cows, late
gestation cows, and growing heifers,
respectively.
Breed might also affect Cu require-
ments, as well as susceptibility to Cu
toxicosis in cattle. Simmental and
Charolais cattle seem to have a higher
minimal Cu requirement than Angus
cattle (Ward et al., 1995; Mullis et al.,
2003). Recently, mRNA expression of
transporters involved in Cu absorp-
tion in the duodenum were found to
be lower in pregnant Simmental cows
than in Angus cows (Fry et al., 2013).
Jersey cows tended to have greater
liver Cu concentrations than Holstein
cows when Cu was supplemented to
diets at high (80 mg/kg; Du et al.,
1996) or moderately high concentra-
tions (20 mg/kg; Sol Morales et al.,
2000).
Milk yield, DMI, SCC, and plasma
Cu concentrations were similar in
lactating Holstein cows supplemented
with 0 (control diet analyzed 8 mg of
Cu/kg), 15, or 30 mg of Cu/kg during
an 83-d study (Chase et al., 2000).
Within each Cu treatment, cows in
this study received 0 or 500 mg of
supplemental Fe/kg of DM. High di-
etary Fe decreased liver Cu in control
Mineral and vitamin nutrition
cows but had little effect on liver Cu
concentrations in Cu-supplemented
cows. Supplementing 10 or 40 mg of
Cu/kg of DM to a control diet, con-
taining 0.24% S, 8.9 mg of Cu, 1.1 mg
of Mo, and 239 mg of Fe/kg of DM,
also did not affect milk yield, DMI,
SCC, or plasma Cu concentrations in
lactating Holstein cows (Engle et al.,
2001). Liver Cu concentrations did
not change during the 61-d study in
cows fed the control diet (374 on d 0
vs. 372 mg of Cu/kg of DM on d 61),
suggesting that 8.9 mg of Cu/kg of
DM was adequate to meet require-
ments.
Studies examining the effect of
dietary Cu on immune responses in
cattle have been reviewed (Weiss and
Spears, 2006). Scaletti et al. (2003)
evaluated the effect of dietary Cu on
responses of Holstein heifers to an
intramammary Escherichia coli chal-
lenge at 34 d of lactation. Heifers were
fed a control diet (6.5 mg of Cu/kg)
or the control diet supplemented with
20 mg of Cu/kg from 60 d prepartum
through 42 d of lactation. Heifers
supplemented with Cu had lower E.
coli numbers and SCC in milk, lower
clinical scores, and lower peak rectal
temperatures than controls. Although
the severity of E. coli infection was
decreased by supplemental Cu, the
duration of infection was not affected
by Cu (Scaletti et al., 2003).
Mullis et al. (2003) estimated the
Cu requirements of Angus and Sim-
mental females fed corn silage–based
diets. Based on a decline in liver Cu
over time, a diet containing 0.26% S,
6.4 mg of Cu, and 1.2 mg of Mo/kg
of DM did not meet the Cu require-
ments of growing Angus or Simmental
heifers during a 160-d study. A diet
containing 0.26% S, 4.4 mg of Cu,
and 1.2 mg of Mo/kg of DM was also
inadequate for first-calf heifers during
late gestation and early lactation. Ad-
dition of 7 mg of Cu/kg of DM to the
control diets met the requirements of
growing and first-calf heifers of both
breeds and resulted in increases in
liver Cu. Based on plasma Cu concen-
trations (Mullis et al., 2003), Sim-
mental females had a greater minimal
183
Cu requirement than Angus females.
Plasma Cu concentrations decreased
in Simmental heifers fed the control
diets over time to levels indicative of
at least marginal Cu deficiency. In
Angus heifers fed the control diets,
plasma Cu concentrations remained
within the normal range throughout
the study. Copper supplementation to
corn silage–based diets containing 5.2
(Ward and Spears, 1997) to 10.2 mg
of Cu/kg of DM (Engle and Spears,
2000a, 2001; Engle et al., 2000b) did
not affect performance of growing
steers. Furthermore, liver Cu concen-
trations did not decrease from initial
levels in growing steers fed control
diets containing 9.9 (Engle et al.,
2000b) or 10.2 mg of Cu/kg of DM
(Engle and Spears, 2000a). Supple-
mentation of Cu to diets containing 5
to 8 mg of Cu/kg and 1.7 to 1.8 mg
of Mo/kg also did not affect perfor-
mance of growing and finishing male
cattle intensively reared in Spain
(Garcia-Vaquero et al., 2011).
Performance responses to Cu sup-
plementation of corn-based finishing
diets have been inconsistent. Copper
supplementation (5 mg/kg of DM)
increased ADG and G:F in steers fed
a control diet containing 0.25% S, 2.9
mg of Cu, and 0.9 mg of Mo/kg of
DM (Ward and Spears, 1997). Gain
and DMI were also increased by Cu
supplementation (20 mg/kg of DM)
in finishing steers fed a control diet
containing 0.28% S, 4.9 mg of Cu,
and 0.6 mg of Mo/kg of DM (Engle
et al., 2000b). In other studies, Cu
supplementation to finishing diets
containing 4.9 to 5.2 mg of Cu/kg
has not affected performance (Engle
and Spears, 2000b, 2001; Engle et
al., 2000a) or decreased cattle per-
formance (Engle and Spears, 2000a).
Copper addition to finishing diets
containing 2.9 to 5.2 mg of Cu/kg has
decreased backfat without affecting
marbling and increased PUFA concen-
trations in muscle in several studies
with Angus and Angus-crossbred
cattle (Engle, 2011). Dietary Cu did
not affect backfat or PUFA in muscle
of finishing Simmental steers (Engle
and Spears, 2001).
184
Manganese
Results of studies published in the
past 10 yr suggest that the current
beef NRC (2000) Mn recommenda-
tion of 20 mg/kg of DM for grow-
ing and finishing cattle is adequate.
Performance of growing and finishing
male calves fed a corn silage–based
diet, containing 20.9 mg of Mn/kg of
DM, was not affected by Mn supple-
mentation (Kirchgessner et al., 1997).
Manganese supplementation to a
control diet at 10, 20, 30, 120, or 240
mg/kg of DM resulted in a linear in-
crease in liver and LM Mn concentra-
tions but did not affect performance
or carcass characteristics in growing
and finishing cattle (Legleiter et al.,
2005). The control diet used in this
study contained 29 mg of Mn/kg of
DM during the 84-d growing phase
and 8 mg of Mn/kg of DM during the
finishing phase, which averaged 112 d.
Manganese addition to a control diet
containing 16 mg of Mn/kg of DM
did not affect gain or feed efficiency
in growing beef heifers during a 196-d
study (Hansen et al., 2006a)
Manganese requirements for repro-
duction seem to be higher than for
growth. The beef NRC (2000) recom-
mends 40 mg of Mn/kg for breeding
cattle, whereas the dairy NRC (2001)
recommends 16 to 18 mg of Mn/kg
during gestation and 12 to 14 mg of
Mn/kg of DM for lactating cows. Re-
cent research indicates that the cur-
rent dairy NRC (2001) underestimates
Mn requirements for cows. Based on
regression of digestible Mn on Mn
intake, Weiss and Socha (2005) esti-
mated maintenance requirements for
Mn to be 49 and 28 mg/kg of DM, re-
spectively, for dry and lactating dairy
cows. The addition of 10 to 50 mg
of Mn/kg to a control diet contain-
ing 16 mg of Mn/kg of DM did not
affect age at conception or services to
conception in beef heifers (Hansen et
al., 2006a). Although pregnancy rate
tended to be higher in heifers supple-
mented with 50 mg of Mn/kg of DM,
differences among treatments were
not significant. In this study, treat-
ment diets were fed from shortly after
Spears and Weiss
weaning through the first trimester of
pregnancy. At the termination of this
study (Hansen et al., 2006a), pregnant
heifers from the control and 50 mg/
kg of supplemental Mn treatments
were selected to remain on treatments
through gestation and early lacta-
tion (Hansen et al., 2006b). Results
of this study clearly indicated that 16
mg of Mn/kg of DM was inadequate
for proper fetal development. Calves
born to control heifers were lighter
at birth and had lower whole-blood
Mn concentrations than calves from
Mn-supplemented heifers. Approxi-
mately 50% of calves born to heifers
fed the control diet (16 mg of Mn/
kg of DM) exhibited clinical signs
of Mn deficiency, including superior
brachygnathism, unsteadiness, swollen
joints, and disproportionate dwarfism
(Hansen et al., 2006b).
High dietary Fe might increase Mn
requirements. Manganese seems to use
the same transporter (divalent metal
transporter1) from the small intestine
as Fe, and high dietary Fe (810 mg/
kg of DM) decreased Mn concentra-
tions in duodenal mucosal scrapings
in calves (Hansen et al., 2010). Lim-
ited evidence also suggests that high
dietary Ca and P might decrease Mn
bioavailability (Spears, 2003).
Zinc
Based on the variable responses to
Zn supplementation that have been
observed, Zn requirements of cattle
seem to be affected by dietary factors,
but factors that affect Zn bioavail-
ability are not well defined (Spears,
2003). Limited research suggests that
growing heifers have a greater Zn
requirement for growth than bulls
and steers (Price and Humphries,
1980). The requirement for Zn was
estimated at 30 mg/kg of DM in the
last beef NRC (2000). Estimated Zn
requirements for dairy cattle were ap-
proximately 52, 21, and 32 mg/kg of
DM for lactating cows, late gestation
cows, and growing heifers, respectively
(NRC, 2001).
Performance and morbidity of
receiving cattle were not affected by
Zn supplementation (360 mg of Zn/d)
to a control diet containing approxi-
mately 25 mg of Zn/kg (Kegley et
al., 2001). In a 35-d receiving study
with heifers, supplementing 75 mg of
Zn/kg to a control diet that analyzed
52.5 mg of Zn/kg of DM did not af-
fect morbidity but tended (P = 0.11)
to decrease ADG and decreased G:F
(Nunnery et al., 2007).
Consistent with earlier studies
(NRC, 2000), responses to Zn supple-
mentation of growing and finishing
cattle diets have also been variable
in recent studies. The addition of
25 mg of Zn/kg to a corn silage–
based diet containing 33 mg of Zn/
kg of DM increased ADG of growing
steers by 0.10 kg/d during an 84-d
study (Spears and Kegley, 2002). In
other studies with growing diets, Zn
supplementation has not affected
performance of cattle fed control diets
containing 38 (Kegley et al., 2001), 35
(Kessler et al., 2003), or 28 mg of Zn/
kg of DM (Wright and Spears, 2004).
Performance did not differ among
finishing steers supplemented with 20
(total dietary Zn of 90 mg/kg), 100,
or 200 mg of Zn/kg of DM; however,
fat thickness and yield grade in-
creased quadratically with increased
dietary Zn (Malcolm-Callis et al.,
2000). Zinc supplementation (25 mg
of Zn/kg) to a control diet containing
26 mg of Zn/kg of DM did not affect
performance of finishing steers but in-
creased quality grades slightly (Spears
and Kegley, 2002). In contrast carcass
characteristics were not affected by
dietary Zn in finishing heifers, but
heifers fed the control diet (50.5 mg of
Zn/kg) tended to gain less (P = 0.11)
and have a lower (P = 0.06) G:F than
those supplemented with 75 mg of
Zn/kg of DM (Nunnery et al., 2007).
Increasing total dietary Zn from
41 to 63 mg/kg of DM did not affect
milk yield, milk composition, milk Zn
concentrations, or hoof hardness and
locomotion scores of lactating dairy
cows (Cope et al., 2009). Milk from
cows fed the higher level of Zn had
lower SCC and lower concentrations
of amyloid A, an acute-phase protein,
than cows fed 41 mg of Zn/kg of DM.

The lower SCC and milk amyloid A
concentrations suggest improved ud-
der health in cows receiving 63 mg of
Zn/kg of DM.
UPDATE ON VITAMINS
FOR RUMINANTS
Responses to supplemental vitamins
by ruminants include improved im-
mune function, fewer clinical health
problems, increased productivity,
and changes in meat characteristics.
Responses vary depending on the
vitamin, dose, and species or type of
animal supplemented. Not all the 14
recognized vitamins (for this review,
choline is considered a vitamin) have
been shown to elicit whole-animal
responses (e.g., health or production
measures) when supplemented to ru-
minants. Therefore, discussion will be
limited to vitamins A, D, E, and B12,
biotin, choline, niacin, and thiamine
because sufficient newer (1998 to
2013) whole-animal data are available
to reach conclusions regarding field
application.
Vitamin A
The beef NRC (2000) recommenda-
tions (IU/kg of DMI) for vitamin A
are 2,200, 2,800, and 3,900 IU/kg of
DMI for growing or finishing animals,
gestating animals, and lactating cows
or breeding bulls, respectively. Based
on a survey of nutritionists, finishing
beef cattle are commonly supplement-
ed with vitamin A at almost twice
NRC recommendations (Vasconcelos
and Galyean, 2007). Feeding supple-
mental vitamin A to finishing steers
at up to 4 times NRC recommenda-
tions did not enhance production or
carcass measurements (Bryant et al.,
2010). Indeed, some negative effects
were observed when diets contained 2
to 4 times the recommended amount
of vitamin A. In addition, the concen-
tration of α-tocopherol in liver was
inversely related with rate of vitamin
A supplementation, indicating the
possibility of a secondary vitamin E
deficiency when excess vitamin A was
fed. Some (Gorocica-Buenfil et al.,
2007; Pickworth et al., 2012; Ward
Mineral and vitamin nutrition
et al., 2012), but not all (Gorocica-
Buenfil et al., 2008; Bryant et al.,
2010), studies have reported improved
marbling and quality grade when
feedlot cattle were fed no supplemen-
tal vitamin A during the finishing
phase compared with cattle fed at ap-
proximately NRC recommended lev-
els. No negative effects on growth or
efficiency were associated with feeding
diets devoid of supplemental vitamin
A in any of the studies cited above.
Current data indicate no advantage
to feeding greater than NRC recom-
mendations for vitamin A to finishing
cattle. Indeed current data suggests
that vitamin A supplementation rates
for feedlot cattle could be decreased.
The current vitamin A requirement
for lactating cows is 110 IU/kg of BW
(NRC, 2001). That recommendation
is based mostly on animal health and
immune function data. Newer data
(LeBlanc et al., 2004; Bertoni et al.,
2008) supported the link between few-
er health problems and enhanced vita-
min A status but did not show that
higher rates of supplementation were
necessary to achieve good vitamin
A status. The commonly observed
decrease in plasma retinol concentra-
tions that occurs around calving was
eliminated by supplementing dry cows
with 880 IU/kg of BW of vitamin A
(Puvogel et al., 2005) or by feeding
300 mg/d of β-carotene (Chawla and
Kaur, 2004); however, the effect of
those treatments on cow health was
not determined because of insuffi-
cient animal numbers. Nonetheless,
the very high supplementation rate
of vitamin A (Puvogel et al., 2005)
decreased milk yield during the first
100 DIM. The current NRC (2001)
recommendation seems adequate for
health and milk production, although
because of potential losses of potency
during storage (Shurson et al., 2011),
including a modest safety factor is
recommended.
Vitamin D
The NRC beef requirement for
vitamin D is about 300 IU/kg of
DMI, but beef animals are usually
housed outside, with sun exposure
185
making the need for supplementa-
tion rare. The only recent research
on vitamin D for beef is the use of
very high supplementation rates (≥1
million IU/d) for approximately the
last week of the finishing period to
improve the tenderness of the beef.
This protocol usually decreases short-
term growth rates, feed intake, or feed
efficiency (Karges et al., 2001; Reiling
and Johnson, 2003; Montgomery et
al., 2004a). That vitamin D protocol
has improved tenderness of beef cuts
in some studies (Karges et al., 2001;
Montgomery et al., 2004b) but not in
others (Scanga et al., 2001; Reiling
and Johnson, 2003). The inconsisten-
cy of improved tenderness linked with
the consistent negative production
effects will likely limit application of
this protocol.
For dry and lactating dairy cows,
the vitamin D requirement is 30 IU/
kg of BW, which was based almost
entirely on the amount needed to
maintain calcium status. New data
mostly from nonruminant studies have
shown that vitamin D has a plethora
of effects other than calcium homeo-
stasis, including profound effects on
immune cell function. Studies that
evaluate the effects of supplementing
dairy cows with vitamin D on health
problems (other than hypocalcemia)
are lacking, but linkages between
vitamin D and bovine immunity have
been shown (Nelson et al., 2010). In
addition, infusing 25-OH vitamin D
into mammary gland quarters that
were infected with Streptococcus uber-
is significantly decreased the severity
of mastitis (Lippolis et al., 2011). In
humans, low concentrations of plasma
25-OH vitamin D is a risk factor for
several health problems (Christakos
and DeLuca, 2011), and similar data
are being generated with cattle. Dairy
cows that were seropositive for Myco-
bacteria antibodies had reduced con-
centrations of plasma 25-OH vitamin
D than cows that were seronegative
(Sorge et al., 2013). Plasma concen-
trations of 25-OH vitamin D in dairy
cows supplemented with NRC levels
of vitamin D and housed with limited
access to sunlight were significantly
lower than concentrations found in
186
cows with full exposure to sunlight
(Hymøller et al., 2009). This finding
might mean that the current recom-
mendation does not result in maximal
vitamin D status, but the ideal or
optimal concentration of 25-OH in bo-
vine plasma has not been determined.
A few older studies (conducted in the
1970s and early 1980s; see NRC, 2001,
for references) reported that feeding
vitamin D at approximately 2 times
the current recommendation increased
milk yields. Thus, the vitamin D
requirements need to be reevaluated
in light of its effects on milk yields,
immune function, and mastitis.
Vitamin E
The beef NRC (2000) suggested
that most beef cattle require little or
no supplemental vitamin E to main-
tain good health and productivity.
Production and reproductive studies
published since the last beef NRC
(2000) have generally found no benefit
of increased vitamin E supplementa-
tion (Cusack et al., 2005; Horn et al.,
2010a,b; Burken et al., 2012). Im-
mune function of feedlot cattle has
been enhanced with supplemental
vitamin E, but effects on morbidity
and clinical health measurements have
been small to nonexistent (Rivera et
al., 2002; Cusack et al., 2005, 2009).
In those studies, the supplementation
rate usually ranged from about 300 to
1,000 IU/animal daily. Supplement-
ing approximately 500 IU of vitamin
E during the last several weeks of
the finishing phase can increase color
stability in beef cuts, which might
become more important when animals
are fed diets with high concentra-
tions of PUFA, such as those with
high inclusion rates of distillers grains
(Burken et al., 2012).
The dairy NRC (2001) requirement
for supplemental vitamin E is ap-
proximately 500 IU/d for lactating
cows and 1,000 IU/d for dry cows.
These recommendations were based
on health and immune function.
Results of newer studies support the
concept that enhanced vitamin-E
status decreases retained fetal mem-
Spears and Weiss
branes and improves mammary gland
health in both dairy cows and dairy
ewes (Morgante et al., 1999; LeBlanc
et al., 2004; Rezamand et al., 2007;
Politis et al., 2012). In addition, low
concentrations of plasma tocopherol
have been found to be a risk factor
for displaced abomasum (Qu et al.,
2013); however, these studies do not
provide support for increasing the
vitamin E requirement for most types
of dairy cows. On the other hand, re-
sults of several studies have indicated
that supplementing 2,000 to 4,000
IU/d of vitamin E during the prepar-
tum period (2 to 3 wk before calv-
ing) improves health compared with
the current NRC recommendation of
1,000 IU/d (Weiss et al., 1997; Baldi
et al., 2000; Politis et al., 2004). Lon-
ger-term (60 d) supplementation of
vitamin E of 3,000 IU/d to dry cows
(3 times the current recommendation)
was found to increase clinical mastitis
on commercial farms, and the authors
suggested that high plasma concentra-
tions of tocopherol might be a risk
factor (Bouwstra et al., 2010a,b).
Newer research findings contradict the
link between high plasma tocopherol
and increased mastitis (Politis et al.,
2012), but these data show no benefit
and perhaps a negative effect of feed-
ing more than NRC requirement for
most of the dry period.
Biotin
Neither the beef (NRC, 2000) nor
dairy NRC (2001) established a
requirement for biotin; however, the
vast majority of data support supple-
menting cattle with biotin. Clinical
studies with beef (cows and growing)
cattle, sheep, and dairy cattle report
improved hoof health when animals
were fed 10 to 20 mg of biotin/d (3 to
5 mg/d for sheep) for several weeks or
months (Midla et al., 1998; Campbell
et al., 2000; Fitzgerald et al., 2000;
Hedges et al., 2001; Bergsten et al.,
2003; Pötzsch et al., 2003; Higuchi et
al., 2004; Bampidis et al., 2007; da
Silva et al., 2010). Milk yield also usu-
ally increases with biotin supplemen-
tation (Lean and Rabiee, 2011), which
is likely a metabolic response and not
caused by improved hoof health.
Choline
The NRC requirements have not
been established for choline, but more
recent studies have reported increased
ADG and improved feed efficiency
with beef cattle and sheep (Bryant
et al., 1999; Bindel et al., 2000) and
increased milk yield in early lactation
dairy cows (Sales et al., 2010) when
rumen-protected choline was fed.
Because of almost complete ruminal
degradation, choline must be rumen
protected to elicit responses. Supple-
mentation rates were approximately
5 and 15 g/d of actual choline for
beef and dairy cattle, respectively.
In addition to production responses,
supplemental rumen-protected choline
(approximately 15 g/d of actual
choline) during the peripartum period
might help prevent or decrease the
severity of fatty liver and fatty liver–
associated ketosis (Cooke et al., 2007;
Zom et al., 2011). The use of choline
to elicit production responses should
be based on potential rate of return,
and often, at least for dairy cows, the
response is profitable. The cost of
ketosis is difficult to quantify because
it is related to so many other health
problems (e.g., mastitis and displaced
abomasum), but decreasing its preva-
lence could be quite profitable.
Other B Vitamins
Although some positive milk pro-
duction responses have been reported
when vitamin B12 is supplemented or
injected (Girard and Matte, 2005),
most studies report no or very limited
increases (Preynat et al., 2009, 2010;
Akins et al., 2013). No new data
on vitamin B12 for beef cattle were
found. Based on the inconsistency of
response, routine supplementation
of vitamin B12 is not warranted, but
adequate cobalt must be fed. Niacin
supplementation is not normally prac-
ticed in the beef industry but is com-
mon for dairy cows. Milk and milk-
component yields have been increased
markedly by niacin supplementation

in some individual studies (e.g., +2.1
kg/d of FCM; Drackley et al., 1998),
but just as commonly, niacin has no
effect on production (Minor et al.,
1998). A meta-analysis determined
that on average, supplementing 12 g
of niacin daily increases milk protein
yield (Schwab et al., 2005). Early
lactation cows are more likely to
respond (Girard, 1998), but specific
dietary conditions that increase the
likelihood of a positive response have
not been identified. Ruminal degra-
dation of supplemental niacin seems
to be >90% (Santschi et al., 2005),
which has led to the development and
commercialization of rumen-protected
niacin. Milk production responses
to rumen-protected niacin have
been small (Yuan et al., 2011, 2012;
Zimbelman et al., 2013), but it might
help decrease heat stress (Zimbelman
et al., 2013).
Thiamine supplementation is not
common for dairy cows and has not
been studied extensively; however,
with the increased use of distillers
grains in beef feedlot diets, supple-
mentation is becoming more common
for those animals. High concentrations
of dietary S are a risk factor for PEM,
and some distillers grains have very
high concentrations of S. Injecting a
large amount of thiamine is standard
therapy for animals suffering from
PEM. Feeding supplemental thiamine
at approximately 120 mg/kg of DMI
to cattle consuming water with 1,000
mg/L of sulfate-S (Ward and Pat-
terson, 2004) or feeding 240 mg of
thiamine/kg of diet DM to sheep fed
a diet with about 0.6% S (Olkowski et
al., 1992) significantly decreased the
incidence of PEM. In the cattle study
(Ward and Patterson, 2004), con-
sumption of the high-sulfate water was
approximately equivalent to feeding
a diet with 0.5% added S. In a recent
study (Neville et al., 2010) with sheep
fed diets with 60% distillers grains
and 0.7% S in the total diet DM,
supplemental thiamine (up to 150
mg/d) had minimal effects on growth
and feed efficiency. The prophylactic
effects of thiamine (if any) could not
be evaluated because no cases of PEM
were observed with any of the diets.
Mineral and vitamin nutrition
IMPLICATIONS
Meeting mineral and vitamin re-
quirements is critical for optimizing
production and health in beef and
dairy cattle. Recommended require-
ments for minerals and vitamins were
last set by the NRC for beef cattle
and dairy cattle in 2000 and 2001,
respectively. Mineral and vitamin
recommendations may change as new
research enhances our understand-
ing of mineral and vitamin needs.
Supplemental biotin has consistently
improved hoof health in cattle, and
providing rumen-protected choline has
increased milk production in dairy
cows. Chromium supplementation has
enhanced insulin sensitivity in cattle
and resulted in improved production
and health in some studies. Estimat-
ing requirements for biotin, choline,
and Cr should be considered by future
NRC committees. Vitamin D require-
ments should be reevaluated based
on its potential effects on immunity.
Current recommendations appear
to overestimate P requirements of
beef finishing cattle. Recent research
suggests that NRC recommendations
for Co and Mn may underestimate
requirements.
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