Milk Fat Depression in Dairy Cows Influence of Supplemental Fats

Charles R. Staples
Professor, Department of Animal Sciences, University of Florida
P.O. Box 110910, Gainesville, FL 32611, (352) 392-1958, FAX (352) 392-1931
Email: [email protected]

Introduction
Supplementing cows with fat can have several beneficial effects. It usually
increases the energy density of the diet when starch or fiber is replaced with fatty acids.
If milk production is increased, then feed efficiency may be improved which usually
translates into more profit. Less heat may be produced in the rumen during digestion of
fat-supplemented diets as fatty acids are not digested in the rumen. Less heat produced
during digestion would help cows during heat stress conditions. Also palatability of the
diet might be improved and feed particle separation may be reduced if a liquid fat is
added to the TMR. As a result, fat inclusion can be a good choice for diet formulators.

Dietary supplementation of fat also can change the concentration of fat in milk, its
daily yield, and its fatty acid composition. What happens in the rumen during microbial
fermentation of feed and metabolism of fats as well as the synthesis and incorporation of
fats by the mammary gland make predicting of these relationships difficult. Some fat
sources will be reviewed as to their effects on milk fat, how these effects might occur,
and what can be done through diet formulation to help control these effects.

Sources and Descriptions of Fats

Choices of fats for feeding to dairy cows are extensive and include oilseeds (fed
whole, rolled, ground, roasted, extruded), rendered fats such as tallow and yellow grease,
vegetable oils, blends of animal and vegetable oils, marine oils, and fats that have been
modified to reduce their metabolism by ruminal microbes such as calcium salts of fatty
acids and prilled fats. Due to time and space limitations, those fat sources that are most
commonly fed will be discussed in more detail. Each fat source is composed of different
fatty acids and those of selected fats are shown in Table 1. Each fatty acid can be
described easily by listing how many carbons make up the length of the molecule and
then how many double bonds occur in each carbon chain. Palmitic acid is a 16-carbon
long molecule with zero double bonds, thus C16:0. The carbon chains that contain zero
double bonds are classified as saturated fatty acids whereas those containing one or more
double bonds are classified as unsaturated fatty acids. Thus fat sources such as the
oilseeds are considered highly unsaturated whereas tallow is considered moderately
unsaturated. This is important because the degree of unsaturation influences microbial
metabolism in the rumen and thus milk fat.
Table 1. Major fatty acid composition of some fat sources (partially adapted from
Kennelly, 1996).
C16:0 C18:0 C18:1 C18:2 C18:3
Fatty acid Palmitic Stearic Oleic Linoleic Linolenic
Tallow 26 19 40 5 1
Yellow grease 21 11 44 14 3
Choice white grease1 24 11 48 12 1
2
Booster Fat 25 22 45 2 -
Megalac; EnerG-II 2 51 4 35 8 -
Canola oil 4 2 52 25 13
Cottonseed oil 25 3 17 54 -
Linseed oil 5 3 20 16 55
Safflower oil 7 2 9 80 <1
Soybean oil 8 3 24 58 8
Sunflower oil 6 4 20 66 <1
Menhaden fish oil 3 17 3 7 1 1
1
Onetti et al., 2001.
2
Commercially prepared, partially ruminally inert fat sources (Church and Dwight Co.;
Virtus Nutrition).
3
Also contains ~10% C16:1, ~11% C20:5, and 12% C22:6.

Fate of Fatty Acids in the Rumen

The ruminal microbes will convert unsaturated fats to saturated fats in a sequence
of events called biohydrogenation. Some scientists have speculated that this act of
biohydrogenation by bacteria is an attempt to protect themselves, as unsaturated fats can
be toxic to bacteria, primarily the bacteria that digest fiber. If the feeding of unsaturated
fats reduces the numbers or activity of fiber-digesting bacteria in the rumen, then feed
intake can decrease, milk production can decrease, and milk fat concentration can
decrease. During the process of biohydrogenation of unsaturated fats in the rumen, the
conversion to the saturated state may be incomplete. This will result in the synthesis of
several forms (isomers) of fatty acids including some trans fatty acids (Figure 1). Some
of the trans fatty acids such as the trans-10, cis-12 conjugated linoleic acid (CLA) and the
trans-10 C18:1 can have a tremendous impact on milk fat when they leave the rumen, are
absorbed into the blood stream, and are taken up by the mammary gland. These trans
fatty acids can be formed by a diverse range of bacteria in the rumen during
biohydrogenation of unsaturated fatty acids (Bauman et al., 2000). This will be
discussed in more detail later.

Figure 1. Proposed schematic for isomerization and biohydrogenation of unsaturated

fatty acids by anaerobic gut microbes. Adopted from Harfoot and Hazelwood, 1997;
Bauman and Griinari, 2003; Mosley et al, 2002.) Solid and dotted lines represent the
major and minor pathways respectively. All compounds may leave the rumen within the
digesta. In addition, the trans monoenes may be isomerized to other trans monene
isomers by microbial isomerases (Proell et al., 2002). In addition, Destaillats et al.
(2205) detected minute amounts of cis-9,trans-13,cis-15 C18:3 and of cis-9,trans-13
C18:2 in milk fat, possibly produced in the rumen.

Oleic Acid Linoleic Acid Linolenic Acid

Cis-9 C18:1 Cis-9, cis-12 C18:2 Cis-9, cis-12, cis-15 C18:3

Conjugated Linoleic Conjugated Linoleic Cis-9, trans-11, trans-10, cis-12,

Acid (CLA) Acid (CLA) cis-15 C18:3 cis-15 C18:3
Cis-9, trans-11 C18:2 trans-10, cis-12 C18:2

trans-11,
cis-15 C18:3 trans-10,
cis-15 C18:3
Trans-11 C18:1 Trans-10 C18:1

Trans-6 to 16
isomers Stearic Acid
C18:1 C18:0

Dietary Fats are Used for Milk Fat

Milk is composed of fatty acids of varying chain length. The 4- to 16-carbon long
fatty acids found in milk are made by the mammary gland from acetate and butyrate, 2-
and 4-carbon long fatty acids respectively, produced by microbes in the rumen. These
fatty acids make up about 50% of the milk fat. The other 50% of the fat in milk comes
directly from fat absorbed from the blood. These fatty acids are primarily 16- to 18-
carbon long fatty acids. All of the 18-carbon long fatty acids and about 30% of the 16-
carbon long fatty acids come primarily from the diet (Akers, 2002). This fat also can
come from the ruminal microbes digested in the small intestine and from body fat
reserves that are mobilized mainly during times of negative energy balance. As fat
appears in the blood stream during digestive metabolism and body fat mobilization, the
mammary gland transfers it into milk fat. It is well established that the more dietary fat
consumed, the greater the proportion of the milk fat is made up of the 18-carbon fatty
acids and thus a smaller the proportion of the shorter chain fatty acids. For example,
when soybeans, cottonseeds, and canola seeds were fed, the proportion of milk fatty acids
of less than 14 carbons, of 14 carbons, and of 16 carbons decreased an average of 25, 29,
and 20%, respectively, whereas the fatty acids of 18 carbons increased 45% for C18:0,
40% for C18:1, 59% for C18:2, and 11% for C18:3 (Ashes et al., 1997). The reduction in
synthesis of the shorter chain fatty acids may be due to a reduction in activity of the key
enzymes responsible for milk fat synthesis. Thus more of the dietary fats were
incorporated directly into milk fat and fewer fatty acids were synthesized by the
mammary gland when supplemental oilseeds were fed.

Milk Fat Depression and Fat Supplementation The Traditional Theory

For some time, the negative effect of supplemental fats on milk fat concentration
was attributed to the fats reduction on ruminal digestion of dietary fiber. It was
proposed that the physical coating of the fiber or of the microbes by fatty acids reduced
fermentation by cellulolytic microbes resulting in less acetic acid being produced in the
rumen. With less acetic acid being delivered to the mammary gland, less milk fat was
synthesized and milk fat concentration was reduced. In addition, forage-digesting
bacteria are pH sensitive and their number decrease with increasing acid conditions in the
rumen. So this negative effect of supplemental fat on fiber digestion was aggravated
when higher grain diets were fed. Interestingly, trans isomers of C18:1 were reported to
increase in the milk following the feeding of soybeans, cottonseeds, or cod liver oil in the
1970s (Storry, 1988). It is only recently that scientists have been able to link some of
these isomers to milk fat depression and thus caused us to rethink the mechanism causing
milk fat depression via fat supplementation. Information on this new theory involving
supplemental fat sources will be discussed below.

Effects of Individual Supplemental Fat Sources on Milk Fat

Tallow. Seven studies fed diets of 0 or 2 to 2.5% tallow in corn silage-based diets
(38 to 50% of dietary DM) (Table 2). Feeding tallow significantly decreased DMI in 4 of
the 6 studies in a range from 1.7 to 3.7 lb/day. This decreased feed intake was
accompanied by reduced milk production of 3.6 lb in one study (Onetti et al., 2001).
However in three other studies, milk production was increased by 4.2 lb/d (Smith et al.,
1993), 5.1 lb/d (Onetti et al., 2002), and 5.3 lb/d (Weiss and Wyatt, 2003). In all seven
studies, milk fat % was significantly depressed from as little as 0.20% to as much as
0.68% units, with the average depression being 0.38% units. Based upon this limited
number of studies, what might be expected when tallow is added to corn silage-based
diets at 2 to 2.5% of the dietary DM? Assuming a similar response to that shown in
Table 2, the efficiency of milk production will be increased (from 1.42 to 1.49 lb of milk
per lb of DM intake in these studies) and milk fat % will be decreased by 0.4 percentage
units (e.g. from 3.35 to 2.97%).

Table 2. Effect of feeding tallow on performance of lactating dairy cows fed diets containing 38
to 50% corn silage (DM basis).
% tallow in DM intake, Milk, lb/day Milk fat, %
Reference diet lb/day
Smith et al., 1993 0 56.0 50.9 3.33
2.5 57.8 55.1* 3.13*
Adams et al., 1995 0 48.1 58.9 3.65
2.5 47.2 58.4 3.35*
Onetti et al., 2001 0 58.0 93.3 3.30
2 54.7* 89.7* 2.83*
Onetti et al., 2002 0 50.9 77.6 3.11
2 49.2* 82.7* 2.82*
Ruppert et al., 2003 0 49.8 71.2 3.18
2 47.2* 73.2 2.89*
Weiss and Wyatt, 2003 0 49.2 77.4 3.76
2.35 48.3 82.7* 3.08*
Onetti et al., 2004 0 60.8 99.0 3.12
2 57.1* 97.7 2.68*
* P 0.05 (estimated if reference did not report exact comparison).
 This negative influence of tallow on milk fat concentration may be eliminated if
some or all of the corn silage is replaced with a different forage such as alfalfa hay.
When Smith et al. (1993) replace one-fourth of the corn silage with alfalfa hay, tallow no
longer depressed milk fat%. Milk fat% was 3.42 vs. 3.47% from cows fed diets of 0 or
2.5% tallow, respectively when diets were 37.5% corn silage and 12.5% alfalfa hay.
When alfalfa hay replaced even more of the corn silage (25% corn silage and 25% alfalfa
hay), milk fat% tended to be increased by feeding tallow (3.35 vs. 3.70% for cows fed 0
and 2.5% tallow diets, respectively). In a Wisconsin study, replacing 25% or 50% of the
corn silage with alfalfa silage did not improve milk fat% when tallow was added to the
diets (Onetti et al., 2002). However a later Wisconsin study documented that the milk
fat-depressing effect caused by tallow supplementation to corn silage-based diets was
alleviated when half of the corn silage was replaced with shortly-chopped alfalfa hay or
with alfalfa silage (Onetti et al., 2004). However feeding long-stemmed alfalfa hay was
not as effective likely due to cows selecting against the long-stemmed forage. Other
forages such as bermudagrass hay and cottonseed hulls also have reversed the milk fat-
depressing effect of tallow when partially replacing corn silage (Adams et al., 1995).
Feeding tallow at 2.5% of the diet reduced milk fat% from 3.65 to 3.35% when corn
silage was the sole dietary forage. Replacing 25% of the corn silage with bermudagrass
hay maintained or improved milk fat % from 3.37 to 3.47% when tallow was fed.
Likewise, supplementing tallow into corn silage-based diets that contained some
cottonseed hulls did not reduce milk fat % (3.53 vs. 3.60% for cows fed 0 or 2.5% tallow
diets, respectively).

A study from the University of Illinois (Ruppert et al., 2003) also indicates that
changing the forage in the diet from predominantly corn silage to alfalfa silage can
alleviate the depressing effect that tallow can have on milk fat % (Table 3). As tallow
increased in the corn silage-based diet from 0 to 2 to 4% (% of dietary DM), milk fat %
tended to decrease from 3.18 to 2.89 to 2.70% whereas milk fat% was unchanged when
tallow was added to an alfalfa silage-based diet (tallow by forage interaction, P = 0.12).
The reason for this response was likely due to what was going on in the rumen. Cows fed
the corn silage-based diets had a more acidic ruminal fluid (average pH of 5.92 vs. 6.04)
likely due to the greater intake of starch. As shown in Table 3, the concentration of trans
C18:1 fatty acids in milk fat tended to increase to a greater extent when tallow was fed in
the corn silage-based diets than in the alfalfa silage-based diets (P = 0.11). Therefore the
introduction of unsaturated fatty acids (predominantly cis-9 C18:1 in the case of tallow)
into a more acidic environment (caused by feeding more corn silage) produced more
trans C18:1 fatty acids in milk (Figure 1). Kalscheur et al. (1997) documented that the
trans C18:1 fatty acids appear in greater amounts in the small intestine when cows are fed
higher grain diets that are not adequately buffered. Therefore cows fed diets that cause a
more acidic pH in the ruminal fluid may be more susceptible to the incomplete
biohydrogenation of dietary unsaturated fatty acids, thus more trans fatty acids are
formed (Figure 2). Feeding diets in which corn silage is the sole or main forage source is
likely to result in a more acidic ruminal fluid than feeding diets containing alfalfa because
alfalfa has 1) greater natural buffering capacity than corn silage and 2) often stimulates
greater buffer production via saliva due to greater chewing of longer particle length
forage. Can these trans fatty acids be affecting synthesis of milk fat?
Table 3. Effect of feeding tallow on selected measurements from lactating dairy cows
fed diets based upon corn silage (CS) or alfalfa silage (AS) (Ruppert et al., 2003).
40% CS : 10% AS 10% CS : 40% AS
Measure 0% 2% 4% 0% 2% 4%
tallow tallow tallow tallow tallow tallow
Milk fat, % a 3.18 2.89 2.70 3.39 3.44 3.41
Rumen pH b 5.94 5.88 5.93 6.09 6.10 6.03
Milk trans-C18:1,
% of milk fat c 1.45 2.95 4.86 1.19 1.89 3.05
a
Dietary forage by tallow interaction, P = 0.12.
b
Dietary forage, P < 0.01.
c
Dietary forage by tallow interaction, P = 0.11.

Figure 2. Cows fed high corn silage diets are more suscep-
suscep-
tible to milk fat depression during fat supplementation

Unsaturated Lower
fat ruminal pH
Trans fatty acids may
decrease synthesis
Increased production
of milk fat
of trans fatty acids by
Rumen bacteria

RUMEN

Trans Fatty Acids. The trans C18:1 fatty acids are a mixture of several isomers,
that is, the single double bond can be located at different carbons along the carbon chain,
anywhere from the 6th to the 16th carbon position (Mosley et al., 2002). Of the trans
C18:1 fatty acids, scientists in New York identified the trans-10 C18:1 fatty acid in milk
fat as being most closely associated with milk fat depression caused by feeding an
unsaturated fat (corn oil) in a higher grain diet compared to a lower grain diet (Griinari et
at., 1998). Onetti et al. (2004) documented that the trans-10 C18:1 fatty acid in milk fat
increased most dramatically when milk fat % was depressed by adding tallow to corn
silage-based diets and returned to normal levels when alfalfa silage replaced the corn
silage. Weiss and Wyatt (2003) reported that cows fed tallow experienced a milk fat
depression and they also had more than a three fold increase in the concentration of trans-
10 C18:1 isomer in their milk fat.
 The major fatty acid in most oil seeds and grains including corn is linoleic acid
(C18:2) (Table 1). Linoleic acid can be converted to trans fatty acids in the rumen. Just
as there are many isomers of trans C18:1, several isomers of linoleic acid can be formed
in the rumen called conjugated linoleic acid (CLA). The most common one is cis-9,
trans-11 CLA but includes the trans-10, cis-12 form of C18:2. This latter fatty acid can
be further converted to trans-10 C18:1 by ruminal microbes (Bauman and Griinari, 2003;
Figure 1). As the C18:2 in corn oil fed by Griinari et al. (1998) underwent
biohydrogenation in the rumen, more of it was converted to trans-10 C18:1 under the
more acidic rumen conditions created by a high grain diet (pH of ~6.4 vs. ~6.1). Ruminal
pH is a determinant of microbial populations in the rumen (Russell, 1979). Those
ruminal microbes that increase in numbers under more acidic ruminal conditions may
possess isomerase enzymes to convert linoleic acid to the trans-10, cis-12 CLA. Trans-
C18:1 acids were formed as the major end product of biohydrogenation rather than C18:0
when increased levels of C18:2 were present (Polan et al., 1964).

However tallow is generally quite low in C18:2, having cis-C18:1 as the dominant
unsaturated fatty acid (Table 1). AbuGhazaleh et al. (2005) demonstrated using
continuous culture techniques that cis-C18:1 was converted to 10 different isomers of
trans-C18:1 (trans-6 to trans-16 except trans-8) with the trans-10 isomer in greatest
concentration. At a ph of 5.5, the dominant trans isomer was trans-10. In addition, more
of the trans-10 isomer was found at a pH of 5.5 than at a pH of 6.5. Interestingly, they
reported that more of the trans-10 isomer originated from sources other than cisC18:1;
that is, a significant part of the trans-10 isomer was made from other trans C18:1 isomers.
This work provides valuable physiological evidence for the increased production of trans-
10 C18:1 isomers from cisC18:1. Therefore a significant portion of the oleic acid in the
form of tallow that was supplemented to the corn silage-based diets fed by Onetti et al.
(2001, 2004) and Ruppert et al. (2003) likely was directly or indirectly converted to trans-
10 C18:1 in the rumen. The trans-10 C18:1 and the trans-10,cis-12 CLA leave the rumen
with the digesta, are absorbed into the blood from the small intestine, and are taken to the
mammary gland where they are incorporated into the milk fat. There is no evidence that
the mammary gland can synthesize trans-10 C18:1 and trans-10, cis-12 CLA fatty acids
(Piperova et al., 2002). However these trans fatty acids may inhibit the synthesis of the
short and medium chain fatty acids by partially inhibiting the enzymes responsible for
milk fat synthesis by the mammary gland (Bauman and Griinari, 2003), thus potentially
accounting for the depressed milk fat % due to the feeding of tallow. It is these trans
fatty acids that may be the cause of the lowered milk fat by tallow supplementation to
corn silage-based diets. Other yet unidentified intermediates in addition to trans-10
C18:1 and trans-10, cis-12 CLA may be responsible for milk fat depression (Bauman and
Griinari, 2003). By partially replacing corn silage with alfalfa (Onetti et al., 2004 and
Smith et al., 1993), the ruminal fluid is less acidic and less of the trans-10 isomers are
produced so milk fat synthesis is not reduced at the mammary gland.

In addition to trans fatty acids being synthesized by the ruminal microbes in the
cow, some dietary fat sources may contain trans fatty acids. Some processed vegetable
oils can contain trans fatty acids such as recycled restaurant grease (yellow grease) due to
the high temperatures reached during the frying process or hydrogenated soybean oil.
Wonsil et al. (1994) supplemented a control diet for lactating dairy cows with either
hydrogenated soybean oil, menhaden oil, or hydrogenated tallow at 3.3% of dietary DM.
Intake of trans C18:1 was 0, 69, 0, and 12 g/d for the 4 diets respectively and the amount
of trans C18:1 appearing in the small intestine was 37, 152, 163, and 38 g/d. This
demonstrates that the trans C18:1 fatty acid was synthesized by microbes in the rumen in
all feeding situations but was found in greater amounts when some was fed (soybean oil)
and when marine oil was fed (more will be said about marine oil in the next section).
The milk fat % was depressed in cows fed diets that delivered the most trans C18:1 fatty
acids for absorption; that is, 3.26, 2.95, 2.78, and 3.18%, respectively. A commercial
calcium salt of primarily (57%) trans C18:1 (EnerG TR; Virtus Nutrition) is being
marketed as a fat supplement to decrease milk fat concentration in markets based on milk
fat quotas. Feeding a calcium salt form of CLA (Virtus Nutrition) at a rate of ~0.33 lb/d
prepartum and ~0.5 lb/d for 7 weeks postpartum resulted in a drop in milk fat
concentration from 3.49 to 2.99% but only a tendency for a decrease in milk fat yield
from 3.04 to 2.62 lb/d (Selberg et al., 2004). This product contained ~8% of the CLA as
trans-10, cis-12 C18:2 so intake of trans-10, cis-12 CLA was ~20 g/d, indicating its
potency to reduce milk fat.

Marine oils. The feeding of oils manufactured from salt-water fish in the oil form
has depressed milk fat of lactating dairy cows. As intake of fish oil increased from 0 to
0.16 to 0.33 to 0.66 lb/day (0%, 0.38%, 0.8%, and 1.8% of dietary DM), milk fat %
decreased linearly (3.95, 4.05, 3.31, and 2.88% respectively) and the trans-10 C18:1 fatty
acid increased linearly (0.29, 0.46, 1.11, and 4.15% of milk fatty acids) (Arola et al.,
2002). Fish oil contains less than 9% C18:1 and C18:2 combined, those fatty acids that
are the usual substrates used by ruminal microbes to produce trans-10 C18:1 and trans-
10, cis-12 CLA. However, AbuGhazaleh and Jenkins (2004) showed that the C20:5 and
C22:6 fatty acids in fish oil increased the trans C18:1 fatty acids and reduced the
biohydrogenation of C18:1 and C18:2 in vitro. Therefore fish oil may act as a modifier
of ruminal bacteria. On a commercial basis, dairy cows are sometimes fed fish meal. As
dietary concentration of fish meal increased in corn silage-based diets (0, 2.6, 5.2, and
7.8% of dietary DM), milk fat concentration decreased linearly (3.5, 3.2, 3.1, and 3.0%,
respectively) (Spain et al., 1995). However in their second study when the diets
contained a nearly equal mix of corn silage and alfalfa silage, fat concentration and yield
were similar (3.8 vs. 3.7% and 2.4 vs. 2.4 lb/d) in milk from cows fed diets containing 0
or 3.8% fish meal (DM basis). Although ruminal fluid pH was not measured by Spain et
al. (1995), it is likely that pH was more acidic when alfalfa silage was omitted from the
diet and therefore may have created an environment leading to a less complete
hydrogenation of unsaturated fatty acids in the rumen; that is, greater trans fatty acid
production.

Whole cottonseeds. Whole cottonseeds are a commonly fed feedstuff supplying

fat, protein, and fiber to dairy cows. About 50% of seeds are processed by oil mills and
50% are fed directly to livestock (National Cottonseed Products Assoc. 1995). About
70% of the fatty acids in cottonseeds are unsaturated. This unsaturated fat can reduce
milk fat % just as tallow has done. The milk fat % was numerically depressed by feeding
whole cottonseeds (10 to 15% of dietary DM) in all nine studies in which corn silage was
the main forage fed but was significantly lower in only two studies (Figure 3). The
average depression was 0.29% units. In a longer term study from calving through 17
weeks postpartum, Jersey cows were fed diets of 0 or 12.9% whole cottonseeds in which
all of the forage came from corn silage. Over the course of the study, cows fed whole
cottonseeds produced milk of significantly lower milk fat % (4.60 vs. 4.88%) (Bertrand et
al., 1998). As was the case with tallow, milk fat % responded differently to the feeding
of whole cottonseeds if alfalfa hay partially or completely replaces corn silage. Milk fat
% was increased by addition of whole cottonseeds to diets when alfalfa hay replaced 25%
(3.30 vs. 3.55%) or 50% (3.25 vs. 3.46%) of the corn silage whereas whole cottonseeds
had little effect on milk fat % when corn silage was the only forage (3.33 vs. 3.27%)
(Smith et al., 1993). In another study, milk fat % was increased by addition of whole
cottonseeds to diets when bermudagrass hay replaced 25% of corn silage (3.37 vs.
3.60%) or when cottonseed hulls replaced 25% of corn silage (3.53 vs. 3.73%) whereas
whole cottonseeds had little effect on milk fat % when corn silage was the only forage
(3.65 vs. 3.54%) (Adams et al., 1995). In this last experiment, alfalfa hay did not have
the positive benefit that it did in the Smith et al. (1993) paper. Therefore the inclusion of
a second forage such as alfalfa hay, bermudagrass hay, or cottonseed hulls with corn
silage should be practiced when diets containing whole cottonseeds in order to generate a
positive milk fat response. Applying the forage lessons learned from feeding tallow, the
second forage should help buffer the rumen in order to be effective.

Figure 3. Feeding Whole Cottonseed (10 15%) in

Corn Silage-
Silage-Based Diets Reduces Milk Fat %

4.6
4.4 * * P < 0.05
4.2
Milk fat, %

4
3.8 * 0 WCS
3.6 + WCS
3.4
3.2
3
2.8
FL, FL, FL, MS, AL, TX, FL, FL, FL,
1984 1984 1987 1989 2001 1985 1995 1990 1993

Location of Study

Other oil seeds. In studies in which cows were fed the same diet throughout the
experiment (continuous study), milk fat concentration was unchanged when cows were
fed whole or rolled sunflower seeds (7 to 10% of dietary DM) or rolled safflower seeds
(10% of dietary DM) in diets containing at least 50% of the forage as alfalfa (Markus et
al., 1996; Stegeman et al., 1992). Again in continuous studies, the feeding of roasted
whole or ground soybeans (12 to 18% of dietary DM) had no effect on milk fat
concentration of cows fed diets in which at least 25% of the forage was alfalfa (Faldet
and Satter, 1991; Pires et al, 1996; Weiss and Wyatt, 2003). Feeding rolled sunflower
seeds, typically high in C18:2 (9% of dietary DM), resulted in a drop in milk fat
concentration from 3.14 to 2.43% whereas feeding sunflower seeds high in C18:1 did not
affect milk fat concentration (2.92%) (Casper et al., 1988). The trans fatty acid content in
milk fat was greatest in cows fed the regular sunflower seeds and the dietary forage was
75% corn silage. Extrusion of the oilseeds appears to consistently depress milk fat
concentration across a number of oilseed sources. Feeding a 50:50 mix of extruded
linseeds and rapeseeds (7.6% of dietary DM) in diets in which the main forage was corn
silage depressed milk fat concentration from 3.65 to 2.98% but this was reversed when
supplemental vitamin E was fed at ~20 times above the NRC recommendation (9616
IU/d) (Focant et al., 1998). However this reversal of milk fat depression by vitamin E
was not accompanied by a reversal in the trans C18:1 increase in milk fat caused by
feeding the oil seeds, suggesting that the milk fat depression was caused by something
other than increased trans C18:1. However the trans-10 C18:1 was not reported. In
studies in which cows were fed the same diets throughout, feeding extruded soybeans
(17% of dietary DM) reduced milk fat content by an average of 0.41% units (3.20 vs.
2.79%) even when dietary forage was at least 25% alfalfa. However the daily production
of fat was not changed due to an increase in milk production (68.6 vs. 75.2 lb/d) caused
by feeding extruded soybeans (Kim et al., 1991; Kim et al., 1993). Again extrusion of
canola seeds (8.5% of dietary DM) resulted in a lowered milk fat concentration (3.14 vs.
3.86%) whereas feeding whole or ground canola seeds did not affect milk fat (Bayourthe
et al., 2000) in corn silage diets. Feeding extruded flaxseeds (13% of dietary DM)
resulted in a numerical reduction in milk fat % (3.82 vs. 3.56%) in diets of 35% grass
silage and 21% corn silage (Gonthier et al., 2005). The trans C18:1 fatty acids were in
greater concentration in the plasma and milk fat. The extrusion process of oil seeds likely
results in a faster and a greater availability of oil in the rumen than when whole or roasted
oil seeds are fed.

Yellow Grease. This fat is derived from the collection and processing of
primarily restaurant grease but it can contain dead stock fat. Because restaurants have
been using primarily vegetable oils for frying, yellow grease (YG) is primarily vegetable
oil and thus has ~2 times more unsaturated fat than does tallow (Table 1). Because these
cooking oils are exposed to higher temperatures and moisture, they have a higher
maximum free fatty acid level (10 to 25%) and are therefore more likely to turn rancid.
When purchasing YG, select ones that have lower free fatty acid levels and contain
antioxidants. Including YG at 2% of dietary DM in diets containing 45% alfalfa hay
increased milk yield 5.5 lb/day and fat yield 60 g/day and did not affect milk fat % (Avila
et al., 2000). In this same study, scientists substituted YG for tallow in order to examine
how changing the degree of saturation affected cow performance. In these alfalfa hay-
based diets, yield of milk (5.5 lb) and fat were increased without changing milk fat %. In
corn silage-based diets, lactating dairy cows fed YG at 5% of dietary DM had lower
digestibility of dietary ADF (21.6 vs. 31.6%) than cows no fed YG, thus contributing to a
5.3 lb drop in DM intake (Jenkins and Jenny, 1989). Although milk production remained
unchanged (69.4 and 70.5 lb/d for cows fed YG or control diet respectively) milk fat %
was depressed from 3.50 to 2.83% by feeding YG. This negative effect on feed intake
and milk fat % may have been minimized if the feeding rate would have been reduced
from 5 to 2%.

Choice White Grease (CWG). This is an animal fat typically composed of a

similar fatty acid profile as yellow grease (Table 1). Milk fat % was depressed from 3.5
to 3.0% when cows were fed diets (36% corn silage and 12% alfalfa hay) of 6% CWG
(Tackett et al., 1996). Similarly milk fat % was depressed from 3.30 to 2.83% when
cows were fed diets (50% corn silage) of 2% CWG (Onetti et al., 2001). In this study,
the trans-10 C18:1 isomer but not the trans-10, cis-12 isomer was in greater concentration
in milk fat of cows fed the CWG.
 Calcium salts of palm oil (CSPO). This product is prepared by reacting palm oil
distillate with sodium hydroxide and water by heating and then precipitating calcium salts
using calcium chloride (CaCl2). Unsaturated fatty acids prepared in this way are less
reactive in the rumen and so negatively affect microbial fermentation to a lesser degree
than untreated unsaturated fats (Chalupa et al., 1986). The acid conditions of the
abomasum split the calcium from the fatty acids so that the fats are absorbed for
metabolism in the small intestine. A review by Chilliard et al. (2001) concluded from
studying 29 different groups that cows fed an average of 0.6 kg/day of CSPO produced
significantly more milk, an additional 2.0 lb/day, with no change in milk fat % compared
to cows not fed CSPO. This was across several experiments and types of diets. Diets
based on corn silage were not examined apart from those based upon a non grain-
containing forage.

How Much Supplemental Fat Can Be Safely Fed?

What management steps can be taken to prevent milk fat depression when
supplemental fats containing unsaturated fatty acids are fed? If a low ruminal fluid pH
is an important factor contributing to milk fat depression, then those factors that help
buffer pH need to be managed. If an unsaturated fat is brought into a ration and the cows
respond with lowered milk fat, it may suggest that that group of cows were borderline
acidotic to begin with. Certainly the effective fiber supply, a forages buffering capacity,
and the presence of supplemental buffering and alkalizing agents should all be
considered. Also the amount of fat consumed has an influence. As concentration of fat
increases in the diet, negative effects of fat can be increased. As one example, when
choice white grease was increased from 0 to 2 to 4% of dietary DM in corn silage-based
diets, milk fat % (3.30, 2.93, and 2.85%) and yield (3.06, 2.67, and 2.38 lb/day)
decreased (Onetti et al, 2001).

Dr. Tom Jenkins at Clemson University has developed some fat-feeding

guidelines based upon the fiber concentration of the diet and the proportion of the
unsaturated fatty acids in the fat supplement (Jenkins, 1993). His equations consider
dietary fiber concentration and the degree of unsaturated fatty acids in the supplement.
The higher the fiber concentration of the diet, the more fat can be included in the diet.
Cows fed diets containing more fiber usually have a less acidic rumen and therefore
fewer trans fatty acids are produced and milk fat depression is minimized. The greater
the proportion of unsaturated fatty acids in the fat supplement, the less fat can be included
in the diet because of the effect these unsaturated fats can have on trans fatty acid
production in the rumen. Two equations exist, one using the NDF and a second using the
ADF concentration of the diet. The current equations do not distinguish between diets
that contain corn silage or alfalfa as the sole or major forage species.

Maximum dietary concentration of supplemental fat =

(6 x % dietary ADF) (% of unsaturated fatty acids as a % of total fatty acids in the fat
supplement % of total fatty acids in fat supplement) x 100% and

(4 x % dietary NDF) (% of unsaturated fatty acids as a % of total fatty acids in the fat
supplement % of total fatty acids in fat supplement) x 100%.
 The unsaturated fatty acids considered are generally C18:1, C18:2, and C18:3.
For tallow shown in Table 1, these add up to 47.1%. Tallow is considered to be 100%
fatty acids. Therefore, a diet that contains the minimum ADF concentration of 19% may
have a maximum dietary concentration of tallow of 2.4% [(6 x 19) (47.1%) (100%) x
(100%)] without resulting in a milk fat depression. Using the NDF values from the corn
silage-based diets from references cited in Table 1, no milk fat depression would be
expected if tallow was fed at less than 2.6% (Onetti et al., 2001), 2.9% (Onetti et al.,
2002), 2.8% (Ruppert et al., 2003), 2.9% (Adams et al., 1995), and 3.1% (Smith et al.,
1993). However, milk fat depression was observed when tallow was fed at 2 to 2.5% of
diet DM indicating that the equations may need to be adjusted if corn silage is the sole
forage source in the diet. Equations that would have maximized tallow to <2 to 2.5% of
dietary DM in the previous references have the initial coefficients reduced as follows:

(4.5 x % dietary ADF) (% of unsaturated fatty acids as a % of total fatty acids in the fat
supplement % of total fatty acids in fat supplement) x 100% and (2.5 x % dietary
NDF) (% of unsaturated fatty acids as a % of total fatty acids in the fat supplement %
of total fatty acids in fat supplement) x 100%.

As more studies are conducted with fat supplements in corn silage-based diets, proper
equations can be developed.

Using the equation guidelines of Jenkins (1993), maximum feeding of whole

cottonseeds is 8.9% when diets are of minimum fiber concentration. Whole cottonseeds
are 18% fat with 71% of the fatty acids as unsaturated fatty acids. Therefore the
maximum feeding of whole cottonseeds = (6 x 19) (71) 18 x 100 = 8.9%. The studies
in Figure 3 fed diets of 10 to 15% whole cottonseeds. The pattern of milk fat depression
across these corn silage-based studies may have been eliminated if level of dietary
cottonseed had been reduced. Palmquist (1984) has recommended that dietary fat intake
should not exceed milk fat output in order to prevent milk fat depression.

Summary
A variety of fat sources have depressed milk fat concentration including tallow
(2%), yellow grease (5%), choice white grease (2%), hydrogenated soybean oil
(3.3%), extruded oil seeds (17%), and fish oil (0.75%) (dietary DM basis).

Milk fat depression is thought to be caused by a reduced synthesis of short-chain

fatty acids by the mammary gland due to the formation of trans fatty acids
(especially trans-10 C18:1 and trans-10, cis-12 CLA) synthesized by ruminal
microbes in a more acidic ruminal environment in the presence of mono- and/or
polyunsaturated fatty acids.

Cows consuming diets that contain corn silage as the only or major forage source
appear to be more susceptible to milk fat depression when unsaturated fats are
supplemented. Partial substitution of corn silage with another forage such as
alfalfa may alleviate this negative effect by making the rumen less acidic.
Keeping ruminal pH above 6.0 for most of the day will likely minimize the
formation of trans isomers by ruminal microbes.
 References

AbuGhazaleh, A.A., and T.C. Jenkins. 2004. Disappearance of docosahexaenoic and

eicosahexaenoic acids from cultures of mixed ruminal microorganisms. J. Dairy Sci.
87:645-651.

AbuGhazaleh, A.A., M.B. Riley, E.E. Thies, and T.C. Jenkins. 2005. Dilution rate and
pH effects on the conversion of oleic acid to Trans C18:1 positional isomers in
continuous culture. J. Dairy Sci. 88:4334-4341.

Ackers, M.R. 2002. Page 95 in Lactation and the Mammary Gland. Iowa State Press.
Ames, IA.

Adams, A.L., B. Harris, Jr., H.H. Van Horn, and C.J. Wilcox. 1995. Effects of varying
forage types on milk production responses to whole cottonseed, tallow, and yeast. J.
Dairy Sci. 78:573-581.

Arola, A., K.J. Shingfield, A. Vanhatalo, V. Toivonen, P. Hahtanen, and J.M. Griinari.
Biohydrogenation shift and milk fat depression in lactating dairy cows fed increasing
levels of fish oil. J. Dairy Sci. 85(Suppl.1):143.

Ashes, J.R., S.K. Gulati, and T.W. Scott. 1997. Potential to alter the content and
composition of milk fat through nutrition. J. Dairy Sci. 80:2204-2212

Avila, C.D., E.J. DePeters, H.Perez-Monti, S.J. Taylor, and R.A. Zinn. 2000. Influence
of saturation ratio of supplemental dietary fat on digestion and milk yield in dairy cows.
J. Dairy Sci. 83:1505-1519.

Baker, J.G., J.E. Tomlinson, D.D. Johnson, and M.E. Boyd. 1989. Influence of two
whole oilseed sources supplemented with Megalac on the performance and milk
composition of early lactating cows. J. Dairy Sci. 72:483 (Abstr.).

Bayourthe, C., F. Enjalbert, and R. Moncoulon. 2000. Effects of different forms of

canola oil fatty acids plus canola meal on milk composition and physical properties of
butter. J. Dairy Sci. 83:690-696.

Bertrand, J.A., F.E. Pardue, and T.C. Jenkins. 1998. Effect of ruminally protected amino
acids on milk yield and composition of Jersey cows fed whole cottonseed. J. Dairy Sci.
81:2215-2220.

Bauman, D.E., L.H. Baumgard, B.A. Corl, and J.M. Griinari. 2000. Biosynthesis of
conjugated linoleic acid in ruminants. Proc. Amer. Soc. Anim. Sci. 1999.
htpp//www.asas.org/jas.symposium/proceedings/0937
Bauman, D.E., and J.M. Griinari. 2003. Nutritional regulation of milk fat synthesis.
Annu. Rev. Nutr. 23:203-227.

Chalupa, W., B. Vecchiarelli, A.E. Elser, D.S. Kronfeld, D. Sklan, and D.L. Palmquist.
1986. Ruminal fermentation in vivo as influenced by long- chain fatty acids. J. Dairy
Sci. 69:1293-1301.

Casper, D.P., D.J. Schingoethe, R.P. Middaugh,, and R.J. Baer. 1988. Lactational
responses of dairy cows to diets containing regular and high oleic acid sunflower seeds.
J. Dairy Sci. 71:1267-1274.
Chik, A.B. 1987. Effects of dietary whole cottonseed on lactational performance and
digestibility as related to roughage source, protein source and calcium treatment. Univ.
of Florida. Ph.D. Dissertation.

Chilliard, Yves, Anne Ferlay, and Michel Doreau. 2001. Effect of different types of
forages, animal fat or marine oils in cows diet on milk fat secretion and composition,
especially conjugated linoleic acid (CLA) and polyunsaturated fatty acids. Livestock
Prod. Sci. 70:31-48.

Coppock, C.E., J.W. West, J.R. Moya, D.H. Nave, J.M. LaBore, K.G.Thompson, L.D.
Rowe, Jr., and C.E. Gates. 1985. Effects of amount of whole cottonseed on intake,
digestibility, and physiological responses of dairy cows. J. Dairy Sci. 68:2248-2258.

Destaillats, F., J.P. Trottier, J.m.. Galvez, and P. Angers. 2005. Analysis of -linolenic
acid biohydrogenation intermediates in milk fat with emphasis on conjugated linolenic
acids. J. Dairy Sci. 88:3231-3239.

Faldet, M.A. and L.D. Satter. 1991. Feeding heat-treated full fat soybeans to cows in
early lactation. J. Dairy Sci. 74:3047-3054.

Focant, M., E, N. Mignolet, M. Marique, F. Clabots, T. Breyne, D. Dalemans, and Y.

Larondelle. 1998. The effect of vitamin E supplementation of cow diets containing
rapeseed and linseed on the prevention of milk fat oxidation. J. Dairy Sci. 81:1095-
1101.

Gonthier, C., A.F. Mustafa, D.R. Ouellet, P.Y. Chouinard, R. Berthiaume, and H.V. Petit.
2005. Feeding micronized and extruded flaxseed to dairy cows: Effects on blood
parameters and milk fatty acid composition. J. Dairy Sci. 88:748-756.

Griinari, J.M., D.A. Dwyer, M.A. McGuire, D.E. Bauman, D.L. Palmquist, and K.V.V.
Nurmela. 1998. Trans-octadecenoic acids and milk fat depression in lactating dairy
cows. J. Dairy Sci. 81:1251-1261.

Harfoot, C.G. and Hazlewood, G.P. (1997) Lipid metabolism in the rumen. In TheRumen
Microbial Ecosystem. P.N. Hobson. Ed. Elsevier Applied Science, London, p. 382-426.
Jenkins, T.C. 1993. Strategies for including fat to dairy rations. Pages 14-25 in Proc.
Clemson Dairy Conf., Clemson, SC.
Jenkins, T.C. and B.F. Jenny. 1989. Effect of hydrogenated fat on feed intake, nutrient
digestion, and lactation performance of dairy cows. J. Dairy Sci. 72:2316-2324.

Kalscheur, K.F., B.B. Teeter, L.S. Piperova, and R.A. Erdman. 1997. Effect of dietary
forage concentration and buffer addition on duodenal flow of trans-C18:1 fatty acids and
milk fat production in dairy cows. J. Dairy Sci. 80:2104-2114.

Kennelly, J.J. 1996. The fatty acid composition of milk fat as influenced by feeding
oilseeds. Anim. Feed Sci. Tech. 60:137-152.

Kim. Y.K., D.J. Schingoethe, D.P. Casper, and F.C. Ludens. 1991. Lactation response
of dairy cows to increased dietary crude protein with added fat. J. Dairy Sci. 74:3891-
3899.

Kim, Y.K., D.J. Schingoethe, D.P. Casper, and F.C. Ludens. 1993. Supplemental dietary
fat from extruded soybeans and calcium soaps of fatty acids for lactating dairy cows. J.
Dairy Sci. 76:197-204.

Lubis, D., H.H. Van Horn, B. Harris, Jr., K.C. Bachman, and S.M. Emanuele. 1990. J.
Dairy Sci. 73:3512 3925.

Markus, S.B., K.M. Wittenberg, J.R. Ingalls, and M. Undi. 1996. Production responses
by early lactation cows to whole sunflower seed or tallow supplementation of a diet based
on barley. J. Dairy Sci. 79:1817-1825.

Mosley, E.E., G.L. Powell, M.B. Riley, and T.C. Jenkins. 2002. Microbial
biohydrogenation of oleic acid to trans isomers in vitro. J. Lipid Res. 43:290-296.
National Cottonseed Products Assoc. 1995. The Cottonseed Processing Industry
Statistical Handbook. Natl. Cottonseed Prod. Assoc., Inc. Memphis, TN.

Onetti, S.G., R.D. Shaver, M.A. McGuire, and R.R. Grummer. 2001. Effect of type and
level of dietary fat on rumen fermentation and performance of dairy cows fed corn silage-
based diets. J. Dairy Sci. 84:2751-2759.

Onetti, S.G., R.D. Shaver, M.A. McGuire, D.L. Palmquist, and R.R. Grummer. 2002.
Effect of supplemental tallow on performance of dairy cows fed diets with different corn
silage:alfalfa silage ratios. J. Dairy Sci. 85:632-641.

Onetti, S.G.,S.M. Reynal, and R.R. Grummer. 2004. Effect of alfalfa forage
preservation method and particle length on performance of dairy cows fed corn silage-
based diets and tallow. J. Dairy Sci. 87:652-664.

Palmquist, D.L. 1984. Use of fats in diets for lactating dairy cows. Chap. 18 in Fats in
Animal Nutrition. J. Wiseman, ed. Butterworths, London, England.

Pires, A.V., M.L. Eastridge, and J.L. Firkins. 1996. Roasted soybeans, blood meal, and
tallow as sources of fat and ruminally undegradable protein in the diets of lactating dairy
cows. J. Dairy Sci. 79:1603-1610.
Piperova, L.S., J. Sampugna, B.B. Teter, K.F. Katscheur, M.P. Yurawecz, Y. Ku, K.M.
Morehouse, and R.A. Erdman. 2002. Duodenal and milk Trans Octadecenoic acid and
conjugated linoleic acid (CLA) isomers indicate that postabsorbtive synthesis is the
predominant source of cis-9-containing CLA in lactating dairy cows. J. Nutr. 132:1236-
1241.

Polan, C.E., J.J. McNeill, and S.B. Towe. 1964. Biohydrogenation of unsaturated fatty
acids by rumen bacteria. J. Bacteriol. 88:1056-1064.

Proell, J.M., E.E. Mosley, G.L. Powell, and T.C. Jenkins. 2002. Isomerization of stable
isotopically labeled elaidic acid to cis and trans monoenes by ruminal microbes. J. Lipid
Res. 43:2072-2076.

Ruppert, L.D., J.K. Drackley, D.R. Bremmer, and J.H. Clark. 2003. Effects of tallow in
diets based on corn silage or alfalfa silage on digestion and nutrient use by lactating dairy
cows. J. Dairy Sci. 86:593-609.

Selberg, K.T., A.C. Lewis, C.R. Staples, N.D. Luchini, and L. Badinga. 2004.
Production and metabolic responses of periparturient Holstein cows to dietary conjugated
linoleic acid and trans-octadecenoic acids. J. Dairy Sci. 87:158-168.

Smith, W.A., B. Harris, Jr., H.H. Van Horn, and C.J. Wilcox. 1993. Effects of forage
type on production of dairy cows supplemented with whole cottonseed, tallow, and yeast.
J. Dairy Sci. 76:205-215.

Spain, J.N., C.E. Polan, and B.A. Watkins. 1995. Evaluating effects of fish meal on milk
fat yield of dairy cows. J. Dairy Sci. 78:1142-1153.

Stegeman, G.A., D.P. Casper, D.J. Schingoethe, and R.J. Baer. 1992. Lactational
responses of dairy cows fed unsaturated dietary fat and receiving bovine somatotropin. J.
Dairy Sci. 75:1936-1945.

Storry, J.E. 1988. The Effect of Dietary Fat on Milk Composition. Pages 111 1441 in
Recent Developments in Ruminant Nutrition 2. Butterworths, London.
Tackett, V.L., J.A. Bertrand, T.C. Jenkins, F.E. Pardue, and L.W. Grimes. 1996.
Interaction of dietary fat and acid detergent fiber diets of lactating dairy cows. J. Dairy
Sci. 79:270-275.

Umphrey, J.E., B.R. Moss, C.J. Wilcox, and H.H. Van Horn. 2001. Interrelationships in
lactating Holsteins of rectal and skin temperatures, milk yield and composition, dry
matter intake, body weight, and feed efficiency in summer in Alabama. J. Dairy Sci.
2680-2685.

Van Horn, H.H., B. Harris, Jr., M.J. Taylor, K.C. Bachman, and C.J. Wilcox. 1984. By-
product feeds for lactating dairy cows: effects of cottonseed hulls, sunflower hulls,
corrugated paper, peanut hulls, sugarcane bagasse, and whole cottonseed with additives
of fat, sodium bicarbonate, and Aspergillus oryzae product on milk production. J. Dairy
Sci. 67:2922 - 2938.

Weiss, W.P., and D.J. Wyatt. 2003. Effect of dietary fat and vitamin E on alpha-
tocopherol in milk from dairy cows. J. Dairy Sci. 86:3582-3591.

Wonsil, G.J., J.H. Herbein, and B.A. Watkins. 1994. Dietary and ruminally derived trans-
18:1 fatty acids alter bovine milk lipids. J. Nutr. 124:556-565.