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Vet Clin Food Anim 23 (2007) 247–268

Feed Value of Supplemental Fats


Used in Feedlot Cattle Diets
Richard Avery Zinn, PhDa,*,
Alejandro Plascencia Jorquera, PhDb
a
Department of Animal Science, University of California, Davis,
Davis, CA 95616-8521, USA
b
UABC, Mexicali, Me´xico

Supplemental fat is added to animal diets for many reasons. It can im-
prove palatability, feed efficiency, and reproductive efficiency, and can
help alleviate heat stress [1]. Certain fatty acids (FA) (ie, conjugated linoleic
acid) may influence nutrient partitioning [2]. Supplemental fat reduces the
evolution of dust generated during the processing and handling of feed.
Chiba and colleagues [3] demonstrated that supplementation with 2.5%
animal fat reduced airborne dust by 82% when feed delivery augers were
operating. Added fat also functions as a lubricant to reduce wear on feed-
mixing equipment and reduce particle separation to improve the uniformity
of feed mixes [4].
Nutritionally, supplemental fats are a concentrated source of energy
(nearly three times the net energy [NE] value of corn [5]), and essential
FA [6]. In the United States market, inedible fats and oils typically have
a least cost advantage (cost/Mcal NE) over cereal grains as an energy
source. The NE of supplemental fat for feedlot cattle are 6.00 and 4.85
Mcal/kg for maintenance and gain, respectively [5]. Supplemental fats are
composed largely (90%) of FA [7,8]; because FA fatty acids areis not di-
gested within the rumen, the energy value of supplemental fats for feedlot
cattle depends on FA digestibility within the small intestine [9].
The performance response to supplemental fat has been quite variable.
Consequently, great attention has been paid to gaining a better understand-
ing of the factors that affect its feeding value. These factors include the type
or source of fat [10–12], the free fatty acid (FFA) concentration [13,14], the

* Corresponding author.
E-mail address: razinn@ucdavis.edu (R.A. Zinn).

0749-0720/07/$ - see front matter Ó 2007 Elsevier Inc. All rights reserved.
doi:10.1016/j.cvfa.2007.03.003 vetfood.theclinics.com
248 ZINN & JORQUERA

degree of saturation or titer [15–17], and the method [18,19] and level of sup-
plementation [9,20–24].

Industry overview
Much of the perishable animal by-products produced in the United
States are handled by renderers and converted into beneficial products
used in the feed and oleochemical industries. In the United States, 25% to
55% of the live slaughter weight of livestock and poultry is considered waste
and unsuitable for retail sale. A total of 24.5 billion kg per year (67 million
kg/d) of animal by-product waste is generated here [25]. During 2005, the
United States rendering industry produced roughly 3 billion kg of inedible
fats and oils. Tallow and yellow grease are the major source of fat used in
animal feeds, representing approximately 80% of total inedible fats and
oils consumed by livestock. Historically, rendered animal fats were used
largely in soap production [26]. However, because of an increasing abun-
dance (and preference) for natural and synthetic alternatives [27], industrial
uses gradually have decreased in importance, increasing the competitiveness
of fats and oils as an energy-dense feed ingredient for livestock. In 2005, 1.8
billion kg of supplemental fat was fed to livestock, 4.8% and 2.9% more
than in 2003 and 2004, respectively.
Numerous sources of fats and oils are available for animal feed; however,
because of price considerations, the more common sources of supplemental
fats are lower grades of tallow, recycled restaurant grease (or yellow grease),
choice white grease, poultry fats, and acidulated animal/vegetable soap
stocks. Usage of tallow and yellow grease by the United States feed industry
increased from 0.9 million metric tons in 1990 to more than 1.3 million met-
ric tons in 2005 (Table 1).

Quality characteristics of feed fats


Although much attention has been directed toward understanding the
factors influencing the NE value of supplemental fat, alterations in NE do
not necessarily form the basis for constraints on supplementation. Indeed,

Table 1
Tallow and grease consumption in animal feed during the 2002–2005 period
Year
Fat source 2002 2003 2004 2005
Inedible tallow 401.2 444.0 428.2 541.9
Greasea 649.8 292.0 769.1 740.9
Total 1050.8 736.0 1197.3 1282.8
a
Yellow grease and other grease sources.
Data from US Census Bureau. Fats and oils. Production, consumption and stocks (millions of
kilograms). Available at: http://www.census/gov/mcd. Accessed February 26, 2006.
SUPPLEMENTAL FATS USED IN FEEDLOT CATTLE DIETS 249

greater concern often is directed at the potential detrimental effects of sup-


plemental fat on diet acceptability (palatability) and feed intake [16,28]. The
reasons for the occasional negative impact of supplemental fats on diet ac-
ceptability by cattle are far from clear, although each time the problem
arises, attention is drawn to the importance of fat ‘‘quality.’’ Feed mills
use measurements of moisture, insoluble impurities, unsaponifiables
(MIU), total fatty acids (TFA), FFA, titer, iodine value, and initial peroxide
value to assess fat quality.

Moisture, impurities, and unsaponifiables


Some condensation moisture is unavoidable with any feed fat. However,
the level should be less than 1.5%. Moisture permits the formation of rust,
and rust accelerates autocatalytic (nonenzymatic) oxidative rancidity. Mois-
ture in the presence of high levels of FFA and high temperature also promotes
autocatalytic hydrolysis of glycerides. The practice of clearing lines with steam
may increase the moisture content of fat in bulk tanks and should be avoided.
Impurities refer to filterable materials insoluble in kerosene, such as fine par-
ticles of hair, bone, hide, minerals, metals, and so forth. Thus, it is not a measure
of, nor does it in any way represent, potentially hazardous contaminants such
as pesticide residues. Feed fats should not contain more than 1% impurities.
Because impurities tend to settle out, they may accumulate as sludge at the bot-
tom of the bulk tank, ultimately clogging valves, lines, and nozzles. Conse-
quently, tanks should be examined and cleaned on a regular basis.
Unsaponifiables refer to that material which is soluble in petroleum ether
but does not react with sodium or potassium hydroxide to form soap. These
materials include various compounds such as sterols, pigments, fat-soluble
vitamins, fatty alcohols, fatty–fatty esters (condensation products), waxes,
mineral oils, pesticides, and so forth. Unsaponifiables usually represent
less than 1% of most feed fats, with the exception of soap stocks or feed
fats containing blends of soap stocks, which may contain more than 4%.
Unsaponifiables apparently contribute very little to the energy value of
feed fat. However, aside from that, a high unsaponifiable value is not any
more indicative of an animal health safety hazard than a low value is indic-
ative of wholesomeness. The potential for feed fats to become contaminated
with pesticides or other toxic chemicals is real. In 1957, large losses were
noted in the poultry industry, presumably because of the presence of di-
oxin-contaminated tallow [29]. It would be expensive to analyze every ship-
ment of feed fat for pesticide residue, but every shipment should be certified
by the supplier to be pesticide-free.

Total fatty acids


TFA is another measure of the purity of the feed-fat source. Triglycerides
contain approximately 90% FA and 10% glycerol. Thus, FA levels of less
250 ZINN & JORQUERA

than 90% reflect dilution with other ingredients. Because FA are the pri-
mary energy source in feed fats, the value of a feed fat should be discounted
based on TFA content [9].

Free fatty acids


FFA refers to FA not esterified to glycerol. In ‘‘whole’’ fats, the presence
of high levels of FFA may be an indication of improper storage or handling
of the fat. Hydrolysis may occur as either enzymatic hydrolysis during stor-
age or before rendering, or as autocatalytic hydrolysis. Often, the latter is
associated with oxidative rancidity. Antioxidants (ie, ethoxyquin) should
be added to all feed fats to prevent rancidity from occurring, particularly
in the presence of high levels of FFA. Limited evidence suggests that fats
with high proportions of FFA may inhibit ruminal biohydrogenation,
thereby increasing intestinal FA digestion [30].

Iodine value
Iodine value refers to the grams of iodine taken up by 100 g of fat. It is
a measure of the degree of saturation of FA (each double bond takes up two
atoms of iodine). The hardness or softness of fats (titer) is in direct relation-
ship to the FA profile of the lipid: the more saturated the FA, the harder the
fat; the more unsaturated, the softer. Feed fats with high iodine values
(O60) typically contain vegetable soap stocks. With the trend away from
the use of tallow in cooking, the iodine value of yellow grease has increased
markedly (titer has decreased).

Initial peroxide value


Peroxide value refers to the current state of oxidative rancidity, and is
measured in mEq/kg of fat. An initial peroxide value of less than 5 mEq/kg
indicates that the sample is not rancid. Properly handled fat should not
exceed an initial peroxide value of 10. However, the rancidity of fat can
change quickly, depending on conditions. Consequently, the best indicator
of the rancidity might be simply to smell the fat.
The presence of trace amounts of copper in complete mixed diets can accel-
erate rancidity greatly, particularly if the fat source has a high iodine value.
Oxidation, or rancidity, does not appear to have a detrimental effect on the
palatability or use of the fat, per se, in swine and poultry [26]. However, the
presence of oxidized fat may lead to the loss of fat-soluble vitamins in the diet.

Sources and types of fat


Forage lipids are found mainly in the form of polyunsaturated esterified
FA such as galactoside-glyceride; FA concentration of this form is rarely
SUPPLEMENTAL FATS USED IN FEEDLOT CATTLE DIETS 251

above 1.5% of diet dry matter. On the other hand, FA concentration in ce-
real grains, oil seeds, and commercial feed fats is variable, higher, and in the
form of triglycerides. Diverse sources of commercial feed fats may be used in
feedlot cattle diets; these differ mainly in impurities content, FFA, and de-
gree of saturation (Table 2) [31–33]. Terminology differences between
traders of fats and oils and nutritionists can lead to confusion. Fats and
oils are classified according to the American Fats and Oils Association [7]
standards, not their origin or composition. For example, tallow may not
be 100% beef fat. It can contain other sources of fats, as long as it meets
the appropriate specifications for titer, FFA, color, and MIU. Within the
rendering industry, tallows are anything with a titer (temperature at which
melted FA obtained from a fat source coagulate or solidify when cooled)
of 40 C or higher, whereas anything softer is considered to be grease. Oils
generally are considered to be polyunsaturated and of vegetable origin.
Commonly used fat sources are described in Table 2.

Yellow grease
The term ‘‘yellow grease’’ is descriptive of its yellowish appearance. It also
may be referred to as ‘‘restaurant grease’’ or ‘‘kitchen grease,’’ because it is
composed of any combination of waste greases collected from bakeries, restau-
rants, school cafeterias, and the like, and of rendered animal fat. During recent
times, cooking has shifted to being done more and more with vegetable oils, so
that most recovered yellow grease is from vegetable origin that has been par-
tially hydrogenated. The unsaturated/saturated FA ratio of yellow grease is
approximately 2.6:1 [34,35]. Because of the source diversity, yellow grease is

Table 2
Average chemical composition of commercial feed fats fed to feedlot cattle
Yellow Bovine Blended Calcium
grease tallow animal- Soap salts
[11,13, [11,15,31, vegetable stocks [32,33,39,
Concept 14,37,50] 40,50,67] fat [9,39,67] [7,13,66] 41,45,67]
Humidity 0.40 0.12 0.88 1.40 d
Impurities 0.22 0.08 0.56 4.90 d
Unsaponifiable 0.71 0.31 3.88 3.46 d
matter
Iodine value 82.06 54.04 67.16 102.60 d
Total FA 92.60 92.48 92.90 85.70 81.30
FFA 13.95 7.80 51.00 54.80 d
FA profile (%)
C16:0 18.03 25.23 22.30 21.50 49.80
C18:0 10.32 15.73 13.70 6.00 4.03
C18:1 46.88 42.18 35.50 26.50 36.30
C18:2 17.16 5.26 18.70 40.20 7.46
C18:3 1.42 0.47 1.55 3.10 0.30
252 ZINN & JORQUERA

not uniform in its composition and may vary from one area to another or one
manufacturing plant and another. According to the standards established by
the American Fat and Oils Association [7], titer must be below 40 C and it
must not contain more than 15% FFA or a maximum of 2% of impurities.

Tallow
Tallow is a by-product operation of the meat industry. Its raw materials in-
clude butcher’s shop scrap, bones, offal, fleshing grease from hides, dead stock,
and chicken grease. All these materials are handled in the same way, through
one of three process variants, dry rendering, wet rendering, and solvent extrac-
tion. Tallow is characterized by a greater uniformity, a higher titer (O40 C)
and lower humidity, impurities, and FFA content (!1.5%), compared with
other fat sources [10,36]. Because of its high titer, tallow can be a greater chal-
lenge to handle or convey into feed mixes when ambient temperature is low.

Blended fats
Blended animal-vegetable fats are a mixture, in any proportion, of ren-
dered animal fat or grease, hydrolysate animal fat or grease, yellow grease,
hydrolysate animal fat or vegetable oil, and acidulated vegetable or animal
soap stocks. Like yellow grease, blended animal-vegetable fat is not uniform
in composition, and thus, it may be misleading to generalize or typify its
characteristics. Compared with yellow grease, it is darker in appearance
and usually higher in FFA and unsaponifiable matter. Blended animal-veg-
etable fats also tend to be higher in iodine value. Typical quality specifica-
tions for blended animal-vegetable fats are 90% minimum TFA, 50%
maximum FFA, 1.5% maximum moisture, 1% maximum impurities, and
3.5% maximum unsaponifiables [9].

Soap extracts and other grease sources high in free fatty acids
Soap stocks are a by-product of animal-vegetable oil refining. The FA
composition is similar to the original oil, but is much higher in nonesterified
FA (O75% unsaturated, O50% FFA). Another high-FFA fat is recycled
‘‘griddle grease.’’ This material is obtained in traps in the rinse water lines
in cafeterias and restaurants. Compared with yellow grease, griddle grease
has a threefold greater concentration of FFA [37]. Griddle grease may be
blended with restaurant grease before processing, resulting in a yellow
grease that is higher in FFA and MIU.

Protected fats
The term ‘‘protected fat’’ may be a misnomer. It generally refers to feed
fats that have been sequestered in a solid form, greatly simplifying handling.
The actual degree of ‘‘protection’’ from ruminal biohydrogenation can be
SUPPLEMENTAL FATS USED IN FEEDLOT CATTLE DIETS 253

variable. Examples of ‘‘protected fats’’ include calcium /magnesium FA


soaps, high-titer fat blends composed predominately of palmitate or stea-
rate, FA alginates, and protein-complexed fat. The last is prepared by
spray-drying a homogenate of highly unsaturated vegetable oil and soluble
protein, resulting in the formation of microparticles (15-50 m in size), each
particle containing lipid surrounded by a layer of protein. The unsaturated
acids are protected from microbial lipases and hydrogenases by treating the
lipid-protein complexes with formalin (approximately 5% by weight).
Formaldehyde crosses links with amino acids to render the protein resistant
to proteolysis in the rumen.

Factors that influence fat digestion in feedlot cattle


Biohydrogenation
Dietary triglycerides are hydrolyzed by ruminal lipases to yield glycerol
and FFA. Glycerol is metabolized further to propionic acid and absorbed
from the rumen. Unsaturated, long-chain FFA undergo extensive biohydro-
genation (a process that reduces their toxicity toward ruminal microorgan-
isms). The degree of biohydrogenation averages roughly 70%, ranging from
60% to 90% [14,37–39]. Assuming a ruminal passage rate of 0.05/h, the rate
of ruminal biohydrogenation averages 0.093/h, with a range of 0.075/h to
0.450/h. Variation in degree of biohydrogenation may be due to physical
form, dietary FA profile, rate of passage (ruminal FA turnover), level of
supplementation, and ruminal pH. The rate of biohydrogenation of C18:1
does not appear to be affected by the level of FA intake. However, for every
percentage unit increase in dietary level of C18:2, the rate of biohydrogena-
tion of C18:2 decreases (0.012/h) [40]. Biohydrogenation of calcium salts of
unsaturated FA is roughly 20% less than that of conventional liquid feed
fats (47%–57%) [39,41]. Long-chain fatty acids (LCFA) resulting from ru-
minal hydrolysis of glycerol esters are not digested or absorbed from the ru-
men, passing through the omasum and abomasum intact. As a consequence
of extensive ruminal hydrolysis and biohydrogenation, lipid flow to the
small intestine is composed mainly of saturated, nonesterified LCFA
(Fig. 1). Because of lipid synthesis by ruminal microbes, FA flow to the
small intestine of cattle exceeds dietary intake.

Emulsification
Lipids are insoluble in water. Their absorption from the small intestine
depends on emulsification and the formation of micelles. The greater the
surface area of lipid micellar structures, the greater their absorption from
the small intestine. The surface area of the micelle is enhanced by the inter-
action of FA with bile salts, and by the proportion of unsaturated FA [14].
Intestinal digestion of unsaturated FA is not affected by the level of
254 ZINN & JORQUERA

Feed

Lipids RUMEN
(Average insaturated:saturated
ratio 70:30)

Lipolysis
LCFA

BH
Insaturated Saturated
FFA FFA

Absorption Propionate Microbial lipids

Small intestine

Fig. 1. Route and transformation of lipids in ruminant gastrointestinal tracts.

supplementation [42–44]. Intestinal digestion of saturated FA (ie, palmitic


and stearic acid) is roughly 80% that of the unsaturated FA (ie, oleic and
linoleic acids) [13,37,45]. But this percentage varies, depending on bile pro-
duction and the proportion of unsaturated FA entering the small intestine.
For example, Børsting and colleagues [46] observed that intestinal digestion
of C18:0 was 92% when cattle were fed diets supplemented with emulsified
vegetable fats protected against ruminal hydrolysis by a formaldehyde-
casein matrix, whereas, with conventional supplemental fats (tallow, yellow
grease), C18:0 digestion was generally less than 75%.
Ruminants have a greater capacity for digestion of saturated FA than do
nonruminants because of lower duodenal pH (2.0–2.5) [47] and greater bile
secretion of phosphatidylcholine and taurocholates [48]. However, because
of extensive ruminal biohydrogenation, most FA entering the small intestine
are saturated. Thus, change in intestinal FA digestion is a predictable func-
tion of the proportion and supply to the small intestine of saturated FA (pri-
marily palmitic and stearic acids) [42,43,49,50].
Micelle formation, the process of emulsification, is facilitated greatly by
the action of bile salts. As shown in Fig. 2, bile flow alone explains 69% of
the variation in FA digestion. Bile production in cattle is independent of
time of day or feeding frequency [51], oscillating between 0.5 and 1.45 mL/
h/kg body weight (BW) [52–55]. Variation in bile production has been attrib-
uted to the level of fat supplementation [55], reduced resorption of bile in dis-
tal parts of the intestine (enterocolic reflex) [56], irritation in the biliary duct
[57], and dietary characteristics. Plascencia and colleagues [55] observed that
SUPPLEMENTAL FATS USED IN FEEDLOT CATTLE DIETS 255

90

FA digestibility, % 85

80

75

70

65

60

55
16 21 26 31 36 41 46
Bile production, mL bile/g of lipids in duodenum

Fig. 2. Relationship of bile production (mL/g of fat) and FA postruminal digestibility, FA di-
gestibility (%) ¼ 65.58 þ 0.431BP, R2 ¼ 0.69, where PB ¼ mL of bile/g of lipids in duodenum.
(From Plascencia A, Mendoza GD, Vasquez C, et al. Influence of levels of fat supplementation
on bile flow and fatty acids digestion in cattle. Journal of Animal and Veterinary Advances
2004;3:763–8; with permission.)

bile production decreased linearly from a high of 40 mL bile/g fat entering


the small intestine at low levels of supplementation, to a low of 17 mL
bile/g of FA entering the small intestine at high levels of supplementation.
Most of the FA not digested within the small intestine are excreted in the
feces as metal (calcium and magnesium) soaps that are not soluble in petro-
leum ether. Consequently, using conventional ether extraction to measure
the lipid content of feeds greatly underestimates fecal lipid excretion.

Free fatty acids


The presence of high levels of FFA in supplemental fats may indicate in-
appropriate storage or handling [58]. Hydrolysis may occur as enzymatic li-
polysis during storage or before processing, or as a result of autocatalytic
hydrolysis, also known as oxidative rancidity [59]. A high proportion of
FFA, per se, in supplemental fats fed to swine and poultry does not affect
growth performance [60]. However, the effect in ruminants is less certain.
Czerkawski [61] observed that the digestibility of flaxseed oil FA was 24%
lower when fed as 100% FFA than when fed esterified as triglycerides. In
contrast, Zinn [9,12] did not detect appreciable differences in growth perfor-
mance, dietary NE, or intestinal digestibility of FA when comparing low-
FFA yellow grease (10% FFA) with a blend of animal-vegetable fat (50%
FFA) supplemented into cattle finishing diets at the rate of 4% and 8%
of diet dry matter.
Considering the chemical composition of triglycerides, glycerol is a dilu-
ent of the NE content. Therefore, increasing the proportion of FFA in a sup-
plemental fat source should increase its energy value slightly [25]. Plascencia
and colleagues [37] evaluated the influence of the proportion of FFA (15%,
256 ZINN & JORQUERA

28%, and 42% FFA) in yellow grease on growth performance and charac-
teristics of digestion in feedlot cattle fed a finishing diet containing 5% yel-
low grease. Although FFA proportion did not affect FA digestion,
increasing the proportion of FFA increased (linear component) daily gain,
feed intake, and gain efficiency, but not dietary NE, demonstrating that
the proportion of higher levels of FFA in supplemental fats may not have
detrimental effects on diet acceptability and feeding value.

Unsaturated/saturated ratio
In vitro studies [62,63] demonstrated that unsaturated FA may have
a more inhibitory effect on the growth and activity of ruminal microbes, es-
pecially cellulolytic bacteria, oleic acid (C18:1) having the greater inhibitory
effect. However, growth performance responses in feedlot cattle have not
been supportive of this finding. Brandt and Anderson [10], comparing tallow
(lower unsaturated/saturated ratio) and yellow grease (higher unsaturated/
saturated ratio), observed similar responses to both sources of fat in the first
of a two-trial study. In the second trial, dry matter intake and daily weight
gain were lower for cattle fed the yellow-grease–supplemented diet. In other
studies [64,65] comparing tallow versus yellow grease, weight gain and dry
matter intake was lower for the tallow-supplemented diets. For the most
part, it can be generalized that the unsaturated/saturated ratio, per se, of
supplemental fat, does not affect the feeding value of fat for feeding cattle,
even at supplementation levels of 6% or more [9,12,13,17,66–69]. When the
unsaturated/saturated ratio of a supplemental fat is very low, the titer, or
melting point, of the fat can be so high that intestinal emulsification and di-
gestion is reduced appreciably. For example, the intestinal FA digestibility
was reduced from 74% for tallow to 37% for high hydrogenated tallow
[70,71]. Likewise, the intestinal digestibility of yellow grease was reduced
by 23% when it was fed in a hydrogenated form [72].
Biohydrogenation of unsaturated FA is a process that reduces their tox-
icity toward ruminal microorganisms due to detergent action on the micro-
bial cell membrane [73]. Microbial biohydrogenation of unsaturated FA
requires a free carboxyl to proceed [74]. Thus, ruminal hydrolysis of lipids
by lipase, galactosidase, and phospholipase produced by ruminal bacteria
(mainly Anaerovibrio lipolytica and Butyrivibrio sp) [75] is rate limiting.
Some of the most relevant factors affecting biohydrogenation rate are rumi-
nal pH [76], microbial population [77], nature of lipids intake [78], and flow
rate (ruminal turnover of FA) [79]. The rate of ruminal biohydrogenation of
C18:1 is not affected appreciably by the level of fat supplementation [80]. In
contrast, for each percentage unit increase in C18:2 supplementation, the
rate of biohydrogenation of C18:2 decreased by 0.12%/h [40]. Considering
that the rate of ruminal triglyceride lipolysis is intense (O90%/h) [81], the
degree of biohydrogenation of unsaturated FA is also intense (70%, range
60%–93%) [14,37–39,82,83]. For example, Wu and colleagues [39], feeding
SUPPLEMENTAL FATS USED IN FEEDLOT CATTLE DIETS 257

cows with an blend of animal-vegetable fat (59% unsaturated FA), noted


that linoleic acid intake increased from 171 to 296 g/d, whereas the flow
of linoleic acid to the duodenum only increased from 45 to 54 g/d. Likewise,
in steer fed diets with supplemental soy oil soap stock (84% unsaturated
FA), intake of C18:1 increased from 38 to 99 g/d, whereas the flow of
C18:1 to the duodenum only increased from 19 to 27 g/d [66]. Even with
many of the ‘‘protected’’ fats, ruminal biohydrogenation remains high
(47% to 57%) [14,39,41,84]. Thus, the unsaturated/saturated ratio of sup-
plemental fats has less relevance for ruminants, compared with nonruminant
species [60,85].
Because the digestibility of the saturated FA C16:0 is greater than the di-
gestibility of saturated FA C18:0, increasing the proportion of C16 FA in
the supplemental fat may improve its feeding value [86,87]. Accordingly,
Corona and colleagues [88] observed that a 50:50 blend of a supplemental
fat consisting of 97% C16:0 with yellow grease (O65% unsaturated FA) in-
creased intestinal FA digestibility, compared with the original fat sources
fed separately.
Zinn and colleagues [14] reported that, when ruminal biohydrogenation
was reduced by protecting fat in a protein-formaldehyde matrix, intestinal
FA digestion increased (87.8% versus 80.3%). Additionally, they observed
that for each percentage unit increase in the proportion of unsaturated
FA in relation to TFA entering the small intestine, the digestibility of
C18:0 increased by 1%.

Method of supplementation
Devendra and Lewis [89] proposed that the physical coating of feed parti-
cles with supplemental fat may impede microbial enzymatic activity and
thereby reduce diet digestibility, particularly fiber digestion. Indeed, approx-
imately 80% of total lipids in ruminal chyme are associated with feed particles
[90]. However, at practical levels of supplementation (!6%), the method of
supplementation (mixing fat with grain or forage before adding other dietary
ingredients versus adding fat as a final step in batch mixing) does not affect
appreciably the characteristics of either ruminal or total tract organic matter
digestion [19]. Zinn and colleagues [18], evaluating the feeding value of yellow
grease supplemented at 5% of dietary dry matter in a steam-flaked, corn-
based finishing diet, did not observe differences in either cattle growth perfor-
mance or digestive function when 20% supplemental fat was added to the
grain initially versus adding the fat at the end of the batch-mixing process.
Likewise, Plascencia and Zinn [91] did not observe differential effects on the
feeding value of supplemental fat when the fat was concentrated in the forage
portion of the diet (alfalfa hay was saturated with 20% yellow grease). Be-
cause the titer (melting point) of supplemental fat can affect the degree of li-
polysis [40], it may be reasoned that supplemental fats with higher titers
may have differential feeding values, depending on method of
258 ZINN & JORQUERA

supplementation. However, following a protocol similar to that of Zinn and


colleagues [18], Plascencia and colleagues [17] did not observe interactions be-
tween method of supplementation and the feeding value of tallow (a higher-
titer fat source) versus yellow grease (a lower-titer fat source).

Level of supplementation
Cattle require a brief period of adaptation to supplemental fat. Although
adapted cattle can tolerate high levels of fat supplementation (6%) without
depressing growth performance [9,19,20,24,92], as little as 3% supplemental
fat can affect detrimentally the growth performance of nonadapted cattle
[11,28]. This effect is not related to the energy value of the fat, per se, but rather
to the effect of fat supplementation on diet acceptability (palatability). Conse-
quently, receiving diets should not contain more than 2% supplemental fat.
The feeding value of fat varies, depending on the extent of intestinal di-
gestion. Intestinal FA digestion averages 77% in cattle (Table 3) [93–96], but
varies greatly, depending on the level of supplementation. Indeed, the level
of supplementation is the single most important factor affecting the energy
value of fat (Fig. 3, Table 4) [97].

Table 3
Postruminal digestibility reported for different types of fats
Level of
Type of fat supplementation (%) Digestibility (%) Reference
AVB 3.0 73.8 [39]
YG 3.0 76.2 [22]
YG 3.0 82.0 [91]
Tallow 3.5 77.1 [66]
YG 4.0 80.0 [12]
YG 4.0 79.1 [35]
TG 5.0 76.0 [93]
YG 5.0 75.5 [95]
YG 5.0 84.2 [33]
YG 5.0 80.2 [96]
Tallow 5.0 72.0 [50]
YG 5.0 84.2 [33]
Tallow 5.5 81.2 [94]
YG 6.0 77.5 [34]
AVBa 6.0 66.9 [39]
YG 6.0 75.6 [91]
YG 6.0 77.1 [12]
AVB 6.0 78.4 [12]
YG 6.0 79.5 [14]
YG 6.0 77.5 [34]
YG 6.0 75.6 [91]
YG 8.0 69.3 [12]
YG 9.0 70.1 [91]
Abbreviations: AVB, animal-vegetable blend; YG, yellow grease; TG ¼ triglycerides.
a
FA of.
SUPPLEMENTAL FATS USED IN FEEDLOT CATTLE DIETS 259

90

Y= 87.560 - 8.591X ; R2= 0.89, P< 0.001


85 3 5
7 4
Fatty acid digestion, %

7 3
6 7
80 5 7
2 4
6 7
75 1
7

4 4
70 6
2
7 2
1 7
2
65 2

60

55
0 0.5 1 1.5 2 2.5 3 3.5
Total fatty acid intake, g/kg BW

Fig. 3. Influence of level of FA intake (g/kg of BW) on FA intestinal digestion. (From Plascen-
cia A, Mendoza GD, Vasquez C, et al. Relationship between body weight and level of fat sup-
plementation on fatty acid digestion in feedlot cattle. J Anim Sci 2003;81:2653–9; with
permission.)

FA digestion ð%Þ ¼ 87:560  ð8:591  FA intake ½g=kg BWÞ;


R2 ¼ 0:89; n ¼ 25

Given that 1 g of digestible fat has a ME value of 9 Kcal (100% of its


physiologic fuel value) and the partial efficiency of use of ME from dietary
fat for BW gain is 67% [24,98,99], the expected NE for gain (NEg) value of
dietary fat is 6.03 Kcal/g digested fat. Thus, the actual NE value of a given
fat is dynamic, largely depending on the level of supplementation. For ex-
ample, if TFA intake is 0.75 g/kg BW (ie, 3.5% total dietary fat), then
the expected intestinal FA digestion is 81.1% (0.8756  [0.0859  0.75])
and the corresponding NEg value is 4.89 Mcal/kg (6.03  0.811). When
the level of FA intake is increased to 1.50 g/kg BW (ie, 7% total dietary
fat), the expected intestinal digestion of dietary fat decreases to 74.7%,
and the corresponding NEg value of fat decreases to 4.50 Mcal/kg. Corre-
sponding NE for maintenance (NEm) values for fat at the two levels of in-
take are 6.04 and 5.60 Mcal/kg, respectively, where NEm ¼ (NEg þ 0.41)/
0.877. An example of how ME and NE values for supplemental fat are ex-
pected to vary according to total fat intake in a 450-kg steer with a dry mat-
ter intake of 10 kg/d is shown in Fig. 4. To optimize the feeding value of
supplemental fat, it is recommended that total lipid intake in finishing diets
not exceed 01.0 g/kg BW, or 7% of dietary dry matter.

Supplemental fats in Holstein versus conventional beef breeds


The impact of FA intake on the feeding value of fat is particularly impor-
tant when comparing the relative responses in beef versus Holstein steers.
260 ZINN & JORQUERA

Table 4
Comparison of observed NEm and NEg content (Mcal/kg) of feed fat estimated by the replace-
ment technique in finishing diets for steers
Level of
NEm NEg diet supplementation (%) Reference
6.35 5.15 1.5 [22]
6.20 4.53 4.0 [35]
6.06 4.90 3.0 [22]
6.02 4.77 5.0 [13]
5.82 4.69 6.0 [14]
5.78 4.61 6.0 [9]
5.55 4.46 4.0 [50]
5.34 4.41 3.5 [10]
5.33 4.30 6.0 [19]
4.98 6.85 5.0 [37]
4.78 3.87 5.0 [96]
4.63 3.50 6.0 [97]
4.38 3.45 5.0 [18]
3.77 2.95 4.0 [31]
3.66 2.80 9.0 [19]

Holstein steers are large in frame size, having heavier mature weights for
a given placement weight (weight at which cattle enter the feedlot) than con-
ventional beef steers. Furthermore, they have a 9% greater NEm require-
ment than beef steers [5,100]. Consequently, at comparable initial weight
and days on feed, Holstein steers are expected to have greater dry matter in-
take and, hence, FA intake, than conventional beef steers. However,

Fig. 4. Estimated energy value of supplemental fat (Mcal/kg) at different levels of


supplementation.
SUPPLEMENTAL FATS USED IN FEEDLOT CATTLE DIETS 261

provided that the level of FA intake is taken into consideration, little evi-
dence suggests that the feeding value of fat is different for Holsteins than
for beef breeds. In a series of comparative slaughter trials (carcass-specific
gravity), Zinn [35] compared the feeding value of supplemental fat in a finish-
ing diet fed to crossbred steers and calf-fed Holstein steers. Some of the 88%
concentrate (steam-flaked, barley-based) finishing diet was supplemented
with 4% yellow grease, and some without. With both crossbred and Hol-
stein steers, fat supplementation increased empty BW gain (12.5% and
4.4%, respectively). However, with the crossbred steers, fat supplementation
did not affect carcass component gain, whereas with Holstein steers, fat sup-
plementation increased empty body fat and energy gain. With crossbred
steers, fat supplementation increased rib eye area (the expected result of in-
creased weight gain). In both crossbred and Holstein steers, fat supplemen-
tation increased the percentage of kidney-pelvic-heart fat (a consistent effect
of fat supplementation). Fat supplementation did not affect dressing per-
centage, marbling score, or retail yield. Using the replacement technique,
the NEm and NEg values of supplemental fat were 6.40 and 5.20 Mcal/kg,
respectively, for crossbred steers, and 6.00 and 4.85 Mcal/kg, respectively,
for Holstein steers. Based on FA intake (0.85 g/kg BW), observed NE values
for supplemental fat fed to Holsteins were consistent with what was ex-
pected (5.99 and 4.84 Mcal/kg, respectively). It is not certain what the basis
is for the higher NE value for supplemental fat when fed to crossbred steers.
Adjusting for the lower FA intake (0.74 g/kg BW), the NE value of supple-
mental fat should have been only slightly (1%) higher than tabular values.
In a 151-d finishing trial involving 72 Holstein steers (273 kg), Zinn and
colleagues [18] observed that the addition of 5% yellow grease did not affect
average daily gain (ADG), but decreased (6.3%) dry matter intake, and in-
creased feed efficiency (4.7%) and dietary NE (6%). The replacement NEm
and NEg values for supplemental fat were 5.00 and 3.97 Mcal/kg, respec-
tively (83% of the tabular values). These low NE values were actually in
close agreement with those expected (4.95 and 3.93 Mcal/kg, respectively)
because of low intestinal fat digestion (65.2%).
In a 144-d finishing trial, Plascencia and colleagues [37] evaluated the in-
fluence of FFA content of yellow grease on feedlot growth performance in
96 Holstein steers (375 kg). Dietary treatments consisted of an 88%-concen-
trate finishing diet supplemented with 0% or 5% supplemental fat. Fat sup-
plementation increased ADG (11%), feed efficiency (9%), and dietary NE
(6.4%). The replacement NEm and NEg values for supplemental fat aver-
aged 5.39 and 4.37 Mcal/kg, respectively. Again, observed NE values were
in good agreement with expected values (5.49 and 4.40 Mcal/kg, respec-
tively), based on level of FA intake and intestinal FA digestion (73%).
Fat supplementation increased dressing percentage (1.2%) and kidney-pel-
vic-heart fat (20.4%), but did not affect rib eye area or fat thickness.
Because of their heavy mature weights, Holstein steers have the genetic
potential of achieving and maintaining high rates of gain throughout the
262 ZINN & JORQUERA

growing and finishing phases. Zinn and colleagues [14] observed that in well-
managed, calf-fed Holstein steers (initial weight of 122 kg), one can expect
an ADG of greater than 1.6 and 1.7 kg/d during the initial 56 and 112 days
on feed, respectively. Even with conventional steam-flaked, corn-based, fat-
supplemented diets (as are characteristic of most southwestern feedlots) con-
taining an energy density of 2.21 Mcal/kg NEm (dry matter basis), the dry
matter intake required to achieve 1.6 kg ADG is 4.64 kg, or 2.78% of
BW. Removing supplemental fat (4%) from the diet lowers the NEm to
2.06 Mcal/kg and increases the required dry matter intake to 5.04 kg/d, or
3.02% of BW! A sustained DMI of 3% of BW may be untenable. Although
the authors are not aware of any studies evaluating the feeding value of sup-
plemental fat in light-weight, calf-fed Holstein steers (100–120 kg BW) dur-
ing the initial 120 days in the feedlot, fat supplementation may be the only
practical means of optimizing dietary energy density and rate of gain.

Bovine spongiform encephalopathy–related safety of tallow


In recent years, public concern about the safety of foods of animal origin
has heightened because of problems that have arisen with bovine spongi-
form encephalopathy (BSE). These problems have drawn attention to feed-
ing practices within the livestock industry and have prompted health
professionals and the feed industry to scrutinize closely food quality and
safety problems that may arise in foods of animal origin as a result of animal
feeding systems. Nonetheless, despite the magnitude of livestock produc-
tion, the frequency of health problems associated with this sector is very
low.
BSE is regarded widely as the cause of variant Creutzfeldt-Jakob disease,
a rare, but fatal, brain disease that has thus far killed more than 150 people.
The largest outbreak of BSE has been in the United Kingdom, with almost
200,000 cases reported, but cases have been reported in native cattle
throughout Western Europe and in some Central European countries (Slov-
enia, Slovakia, Poland, Czech Republic), Israel, and Japan. However, the
potential spread to other countries through exportation of meat-and-bone
meal feed supplements and live cattle has triggered renewed concerns about
the globalization of BSE.
BSE belongs to a rare group of fatal neurologic diseases in mammals
known as transmissible spongiform encephalopathies. It is an epidemic dis-
ease in dairy cows, presumably caused by feeding cows feedstuffs containing
rendered remains of scrapie-affected sheep and BSE-affected cattle. The ‘‘in-
fective’’ agents consist predominantly or totally of prions (small proteins
found in the membranes of nerve tissues and white cells). ‘‘Normal’’ prions
have a characteristic alpha helix, and are degraded easily by proteolytic en-
zymes. In contrast, infective prions, although containing an identical amino
acid profile, are oriented in a beta helix formation that renders them very
SUPPLEMENTAL FATS USED IN FEEDLOT CATTLE DIETS 263

resistant to enzymatic degradation. Indeed, they are even resistant to high


temperature sterilization [101]. The feeding of meat-and-bone meal to cattle
was identified as the most likely route by which the BSE epidemics initiated
and expanded during the mid-1980s [102]. Tallow, on the other hand, was
considered to have a very low BSE risk. Experimental infection of cattle
with tallow has not been investigated yet, and, hence, the possibility that
it could transmit BSE-infective agents should not be overlooked. Neverthe-
less, epidemiologic studies have failed to find any association between occur-
rence of BSE and consumption of tallow by cattle [103]. Furthermore, in
BSE-spiked rendering studies, no infectivity was detectable in resultant
crude, unfiltrated tallow [104]. More recently, however, the use of tallow
in milk replacers has been suggested as a possible cause of BSE in some
countries [105]. Evidence to support this contention was circumstantial.
Nevertheless, as stated previously, it is not realistic to assume that tallow
could never become contaminated with BSE-infective agents, particularly
when the precise nature of the infective agent has yet to be defined. Perhaps
for this reason, and for this reason alone, the FDA labels tallow a specific
risk material, preventing its importation from countries where there exists
the possibility of rendering of BSE-infected materials.
Taylor and Woodgate [106] observed that the increasingly stringent reg-
ulations placed on tallow in countries that have BSE have depressed tradi-
tional markets for tallow both inside and outside these countries.
Regulations adopted by countries, including the United States and Mexico,
that prohibit the importation of specific risk materials from regions consid-
ered to present an unacceptable risk for introducing BSE also have a nega-
tive impact on tallow marketing.

Summary
Independent of source and form of fat addition to the diet, the nutrient
value of fat assigned by the National Research Council [5] tends to be con-
sistent when total fat intake does not exceed the proportion of 0.96 g intake
fat/kg BW. When fat intake is higher than this proportion, the energy value
of fat is reduced linearly as a direct result of the reduction of intestinal FA
digestibility (mainly C18:0), possibly because of a limited bile production ca-
pacity. Intestinal FA digestibility, and the NE value of supplemental fats
used in feedlot diets, is a highly predictable function, based on the TFA in-
take per unit of BW.

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