Digestion de Lipidos (Ovinos)

Bauman, D. E., J. W. Perfield II, M. J. de Veth, and A. L. Lock. 2003.
New perspectives on lipid digestion and metabolism in ruminants. Proc. Cornell Nutr. Conf. pp. 175-189.
NEW PERSPECTIVES ON LIPID DIGESTION AND METABOLISM IN RUMINANTS
D. E. Bauman, J. W. Perfield II, M. J. de Veth, and A. L. Lock

Department of Animal Science
Cornell University
INTRODUCTION
Fat is an important energy component in the diet of ruminants and over the last
decade fat supplementation has become a common practice to increase the energy
density of the diet for high producing dairy cows. Recent advances have renewed the
interest in dietary fats and lipid metabolism. One development is improved analytical
procedures which have revealed the complex pattern of fatty acids that are produced in
the rumen and incorporated into ruminant meat and milk fat. The old adage what you
eat is what you get is clearly incorrect in the case of dietary lipids and ruminant
animals, and this is of special importance given recent efforts to include lipid
metabolism in dynamic models of ruminant digestion. Following absorption, a major fate
of fatty acids is their oxidation for energy. However, a second recent development is the
recognition that specific fatty acids produced in the rumen also play a critical role as
signaling molecules involved in the expression of specific genes and the regulation of
metabolic processes. Finally, recent discoveries in the functional foods area indicate
that specific fatty acids produced in the rumen may have beneficial effects on human
health, and there is a keen interest in the possibility of designing natural food products
with enhanced levels of these fatty acids.
Given these recent advances, the CNC Organizing Committee asked us to review
digestion and metabolism of lipids in the ruminant. This is a challenging task and a
comprehensive review is well beyond the scope of our presentation. Thus, in the
following sections we will provide an overview of lipid digestion and metabolism, giving
special emphasis to recent developments that provide new insight and impact dairy
cattle nutrition. We will also make a conscientious effort to refer readers to reviews
where appropriate.
DIETARY SUPPLY
The diet of lactating dairy cows typically contains 4 to 5% fat. Higher levels may
adversely effect rumen microbial fermentation so the general recommendation is that
total dietary fat should not exceed 6 to 7% of dietary dry matter (Jenkins, 1993; Doreau
et al., 1997; NRC, 2001). Primary sources of lipid in the ruminant diet are forages and
concentrates, although the lipid content can be increased by the use of fat supplements.
Forages typically contain 4 to 6% of the dry weight of the leaf tissue as lipid, of which
the major lipid class is glycolipids (Harfoot, 1981). The lipid content of concentrates is
usually higher than that of forages, and the majority is present in the form of
triglycerides. Fat supplements that are by-products of rendering and vegetable oil
refining industries contain the lipid predominantly as triglycerides whereas rumen-

protected supplements of Ca-salts are comprised of free fatty acids.
Determining the lipid content of the diet is one of the first requirements for diet
formulation and this presents some challenges. Readers are referred to an excellent
recent review by Palmquist and Jenkins (2003) that compares methods used to
determine dietary lipid and discusses some of their practical limitations. The most
common analytical procedure is the Soxhlet ether extraction (EE) method. However, EE
methods also extract non-fatty acid material that has no nutritive value, and as a
consequence EE overestimates the lipid content of the diet. The degree of
overestimation depends on the type of feedstuff. Nutritive value is primarily associated
with the fatty acids and their content can be as high as 80% of the EE for concentrates
to less than 50% for forages (Palmquist, 1988). To account for this in feeding systems,
Allen (2000) used the limited published literature and developed an equation to predict
total fatty acid content from dietary values of ether extract. In turn, NRC (2001) adopted
a modified version of this equation where: Total fatty acids = EE 1. Clearly, the
accuracy of predicting fatty acid content from EE values will vary by diet and dietary
component.
The fatty acid profile of dietary components also varies, and this may affect the
characteristics of the diet and can be important in assessing nutritional value. The fatty
acid composition of representative feed components and fat supplements is provided in
Table 1. Variation of the fatty acid composition within a specific dietary component is
minor when compared to the variation among feedstuffs.
Table 1. Representative fatty acid composition of some typical feedstuffs and fat
supplementsa
Feedstuff
Fatty acid, % Pasture grass Corn Tallow RP Fatb RP Fat/Reprob
14:0 1 --- 3 1 2
16:0 16 16 25 43 40
16:1 2 --- 6 --- ---
18:0 2 2 18 4 4
18:1 3 30 39 34 29
18:2 13 47 5 8 7
18:3 61 2 1 --- ---
20:5 --- --- --- --- 2
22:6 --- --- --- --- 2
a
Adapted in part from Palmquist 1988.
b
Representative rumen protected (RP) fat supplements are Ca-salts of palm fatty acid
ditillate (EnerGII and EnerGII Repro). Fatty acid values for these provided by D.
Luchini (Bioproducts Inc.).
The double bonds in naturally occurring unsaturated fatty acids are almost
exclusively in the cis configuration; generally, forage sources contain a higher
concentration of linolenic acid (18:3; cis-9, cis-12, cis-15), whereas linoleic acid (18:2;
cis-9, cis-12) is the predominant fatty acid in cereal grains and seeds (Table 1).
Unsaturated fatty acids such as these are susceptible to alteration during processing
and storage. The lipids in forages can be altered during anaerobic fermentations that
occur during ensiling, and unsaturated fatty acids present in forages as well as
concentrates will also decline over time as a result of slow oxidation (Van Soest, 1982).
DIGESTION IN THE RUMEN
An extensive metabolism of lipid occurs in the rumen and this has a major impact on
the profile of fatty acids available for absorption and tissue utilization. Davis (1990)
provided an excellent schematic that illustrates the two major processes that occur in
the rumen, hydrolysis of ester linkages in lipids and biohydrogenation of the unsaturated
fatty acids (Figure 1).
Figure 1. Fat digestion in the rumen. Reproduced from Davis (1990).
Hydrolysis
When dietary lipids enter the rumen, the initial step in lipid metabolism is the
hydrolysis of the ester linkages found in triglycerides, phospholipids, and glycolipids.
Hydrolysis of dietary lipids is predominantly due to rumen bacteria with little evidence for
a significant role by rumen protozoa and fungi, or salivary and plant lipases. The
hydrolysis of lipids is extracellular, and the glycerol and sugars that are liberated are
readily metabolized by the rumen bacteria. Rumen hydrolysis has been most
extensively characterized in Anaerovibrio lipolytica which hydrolyzes triglycerides and
Butyrivibrio fibrisolvens which hydrolyzes phospholipids and glycolipids (Harfoot and
Hazlewood, 1997). Although the extent of hydrolysis is generally high (>85%), a number
of factors that affect the rate and extent of hydrolysis have been identified (see reviews
by Harfoot, 1981 and Doreau et al., 1997). For example, the extent of hydrolysis is
reduced as the dietary level of fat is increased (Beam et al., 2000) or when factors such
as low rumen pH and ionophores inhibit the activity and growth of bacteria (Van Nevel
and Demeyer, 1995; 1996b; Demeyer and Doreau, 1999).
Biohydrogenation
Biohydrogenation of unsaturated fatty acids is the second major transformation that

dietary lipids can undergo in the rumen. It requires a free fatty acid to proceed and as a
consequence rates are always less than those of hydrolysis, and factors that affect
hydrolysis also impact biohydrogenation (Figure 1). Most biohydrogenation (>80%)
occurs in association with the fine food particles and this has been attributed to
extracellular enzymes of bacteria either associated with the feed or free in suspension
(Harfoot and Hazlewood, 1997). The major substrates are linoleic and linolenic acids
(Table 1) and the rate of rumen biohydrogenation of fatty acids is typically faster with
increasing unsaturation. For most diets linoleic acid and linolenic acid are hydrogenated
to the extent of 70-95% and 85-100%, respectively (Doreau and Ferlay, 1994; Beam et
al., 2000). This extensive level of hydrogenation is reduced when diets high in
concentrates are fed, which can be attributed to inhibition of lipolysis at the low pH that
is typically observed on these diets (Van Nevel and Demeyer, 1995; 1996b).
Hydrogenation is also adversely affected when excessive unprotected lipid is present in
the diet. How fat interferes with microbial fermentation is not clear, but is believed to
result from either the coating of feed particles or a direct toxic effect on the rumen
microorganisms (Jenkins, 1993).
Fish oil contains two longer chain fatty acids, eicosapentaenoic acid (EPA; 20:5) and
docosahexaenoic acid (DHA; 22:6), that are often included in rumen-protected lipid
supplements designed to improve reproductive performance. The degree to which EPA
and DHA are biohydrogenated in the rumen is still not well understood. Studies
conducted in vitro suggest that there is little biohydrogenation of these two omega-3
fatty acids (Ashes et al., 1992; Gulati et al., 1999). However, in vivo studies involving
dietary supplements of fish oil indicate that much of the EPA and DHA is
biohydrogenated, although to a lesser extent than typically observed for linoleic and
linolenic acids (Chilliard et al., 2000; Wachira et al., 2000; Scollan et al., 2001).
Classical pathways of biohydrogenation were established using pure cultures of

rumen organisms and these are presented in Figure 2. The rumen bacteria involved in
biohydrogenation have been classified into two groups, A and B, based on their
metabolic pathways (Kemp and Lander, 1984). To obtain complete biohydrogenation of
polyunsaturated fatty acids (PUFA), bacteria from both groups are generally required
(Figure 2). Although Group A contains many bacteria that can hydrogenate PUFA to
trans 18:1 fatty acids, only a few species characterized as Group B can hydrogenate a
trans 18:1 fatty acid to stearic acid (Harfoot and Hazlewood, 1997). This feature of
biohydrogenation explains why increased feeding of PUFA simultaneously causes an
increase in the rumen concentration of monounsaturated fatty acids and a decrease in
the concentration of saturated fatty acids (Noble et al., 1974; Fellner et al., 1995).
The initial step in rumen biohydrogenation typically involves an isomerization of the

cis-12 double bond to a trans-11 configuration resulting in a conjugated di- or trienoic
fatty acid (Figure 2). Next is a reduction of the cis-9 double bond resulting in a trans-11
fatty acid. The final step is a further hydrogenation of the trans-11 double bond
producing stearic acid (linoleic and linolenic acid pathways) or trans-15 18:1 (linolenic
acid pathway). It is also possible to differentially affect steps in the biohydrogenation
process. For example, dietary supplements of fish oil appear to inhibit the last step of
biohydrogenation because they increase rumen outflow of trans 18:1 fatty acids and
reduce the outflow of stearic acid (Wachira et al., 2000; Shingfield et al., 2003). Thus,
supplements containing fish oils must primarily affect group B bacteria. Likewise, diets
that cause a low rumen pH and the feeding of ionophores inhibit the final step in
biohydrogenation resulting in an accumulation of trans 18:1 fatty acids. However, the
extent of the inhibition is much lower than their inhibition of hydrolysis (Van Nevel and
Demeyer, 1995, 1996b).
Figure 2. Biochemical pathways for the biohydrogenation of linoleic and linolenic acids
in the rumen (adapted from Harfoot and Hazlewood, 1997).
cis-9, cis-12 C18:2 cis-9, cis-12, cis-15 C18:3

Linoleic Acid D-Linolenic Acid
(Group A & B)
(Group A) cis-9, trans-11, cis-15 C18:3

(Group A & B)
cis-9, trans-11 C18:2 trans-11, cis-15 C18:2

(Group A) (Group A) (Group B)
trans-11 C18:1 trans-15 and cis-15 C18:1

Vaccenic Acid
(Group B)
C18:0
Stearic Acid
Two key biohydrogenation intermediates are trans-11 18:1 (vaccenic acid; VA)
formed from linoleic and linolenic acids and cis-9, trans-11 conjugated linoleic acid
(CLA) formed in the biohydrogenation of linoleic acid. These intermediates are present
in appreciable quantities in ruminant fat at a ratio of about 3:1, but in the rumen cis-9,
trans-11 CLA is only a transitory intermediate and instead it is VA that accumulates. The
difference is because most of the cis-9, trans-11 CLA found in ruminant fat originates in
the mammary gland and adipose tissue from endogenous synthesis involving the
enzyme delta-9 desaturase with rumen-derived VA as the substrate (see review by
Bauman et al., 2003). This discovery is of special importance in considerations of
designing foods because cis-9, trans-11 CLA is among the most potent naturally
occurring anti-carcinogens (see discussion in Lock and Bauman, 2003).
As analytical techniques improved, we have gained an appreciation of the complexity

of the biohydrogenation processes occurring in the rumen. In addition to major
pathways involving trans-11 18:1 and cis-9, trans-11 CLA as intermediates, there must
be many more pathways. A remarkable range of trans 18:1 and CLA isomers are
present and Table 2 presents their outflow from the rumen based on limited data from
growing cattle and lactating cows.
Table 2. Range of double bond positions of trans 18:1 and conjugated 18:2 fatty acids
and their ruminal outflow (g/day) in growing and lactating cattlea
Trans 18:1 Conjugated 18:2
Isomer Ruminal outflow Isomer Ruminal outflow
trans-4 0.5-0.7 trans-7, cis-9 <0.01
trans-5 0.4-0.6 trans-7, trans-9 <0.01-0.05
trans-6-8 0.4-6.7 trans-8, cis-10 0.01-0.02
trans-9 0.8-6.2 trans-8, trans-10 <0.01-0.10
trans-10 1.7-29.1 cis-9, cis-11 <0.01-0.01
trans-11 5.0-121.0 cis-9, trans-11 0.19-2.86
trans-12 0.5-9.5 trans-9, trans-11 0.22-0.55
trans-13 + 14 6.5-22.9 trans-10, cis-12 0.02-0.32
trans-15 3.2-8.5 trans-10, trans-12 0.05-0.06
trans-16 3.1-8.0 cis-11, trans-13 0.01-0.10
trans-11, cis-13 0.01-0.46
trans-11, trans-13 0.09-0.40
cis-12, trans-14 <0.01-0.05
trans-12, trans-14 0.08-0.19
a
Data derived from three studies where samples were collected from the
duodenum (Duckett et al., 2002; Piperova et al., 2002) or omasum (Shingfield et
al., 2003).
Clearly, many intermediates are formed in the ruminal biohydrogenation of PUFA

and these are absorbed and incorporated into ruminant fat. Some of the isomers may
be formed by chemical bond migration (Griinari and Bauman, 1999) and a portion of the
trans 18:1 isomers can be derived from oleic acid (cis-9 18:1; Mosley et al., 2002). We
also know that diet and changes in the rumen environment can shift the pathways of
biohydrogenation resulting in dramatic changes in the fatty acid intermediates. To date,
this has been best characterized for diets that cause milk fat depression where a
marked increase in trans-10, cis-12 CLA and trans-10 18:1 content of rumen fluid and
milk fat is generally observed (Bauman and Griinari, 2003). In fact, Kim et al. (2002)
recently identified a rumen bacterium, Megasphaera elsdenii, that produces significant
quantities of trans-10, cis-12 CLA. However, the extent to which the various pathways of
biohydrogenation are associated with specific enzymes and species of bacteria or
represent a lack of specificity of the enzymes involved in biohydrogenation is unknown.
Microbial Lipids
A portion of the fatty acids found in the rumen are phospholipid components of
microbial membranes. The rumen microorganisms derive these from de novo synthesis
(mainly 16:0 and 18:0) and the uptake of preformed fatty acids (mainly PUFA; Jenkins,
1993; Harfoot and Hazlewood, 1997). Thus, the level of fat supplementation and its
composition can affect the fatty acid composition of the rumen microorganisms
(Bauchart et al., 1990; OKelly and Spiers, 1991). Demeyer and Doreau (1999) pooled
data from five studies and estimated that the bacteria may contribute up to 17% of the
lipid flowing from the rumen when a corn silage diet is fed.
Fat Supplements
Fat supplements are used as a means to increase the energy density of the diet and
many of these are referred to as inert. In this case inertness simply means that the fat
or fatty acid supplement has minimal affects on rumen fermentation. Although deemed
inert at the level used, they can still be hydrolyzed, if a triglyceride, or hydrogenated, if
unsaturated. Often fat supplements are rumen-protected as a means of avoiding the
deleterious effects of a high fat diet on microbial fermentation. The requirement for
these fat supplements is that the method has efficacy in providing both rumen protection
and post-ruminal availability (Wu and Papas, 1997). Recent protection methods are pH-
dependent and take advantage of the transition occurring between rumen pH (~5.8 to
6.7) and pH of the abomasum (~2 to 4). These methods include formaldehyde-
treatment of a lipid-protein matrix (Ashes et al., 1979), microencapsulation with a water
insoluble lipid coating (Putnam et al., 2003), preparation of fatty acid amides (Fotouhi
and Jenkins, 1992), and formation of Ca salts of fatty acids (Jenkins and Palmquist,
1984). The first two methods can protect esterified or free fatty acids whereas the
protected lipid in the later two methods has to be free fatty acids. Of these, Ca-fatty acid
complexes are commercially used in the US and a typical fatty acid composition for
these supplements is presented in Table 1. The mechanism whereby Ca-salts provide
protection from rumen digestion is related to the inertness of the Ca-fatty acid complex
in the typical rumen environment. However, factors such as low rumen pH and
increased unsaturation of the fatty acid can lead to dissociation of the Ca-fatty acid
complex allowing biohydrogenation to occur (Demeyer and Doreau, 1999). Van Nevel
and Demeyer (1996a) found when free fatty acids derived from soybean oil or the Ca
salts of the same fatty acids were incubated at different pH, the protection from
hydrogenation (~30%) provided by the Ca-fatty acid complex at pH 6.9 was lost when
pH was below 6.0. Furthermore, degree of fatty acid saturation appears to be important,
with PUFA more easily dissociated from the calcium salts as compared to
monounsaturated fatty acids (Sukhija and Palmquist, 1990).
ABSORPTION IN THE SMALL INTESTINE
Biology of Absorption
Analysis of the lipid entering the small intestine has shown that it is virtually
identical to that leaving the rumen. Thus, there is no significant absorption or
modification of the long and medium chained fatty acids in the omasum or abomasum
(Noble, 1981). As a consequence of rumen metabolism as detailed in the previous
section, the lipid entering the small intestine consists of fatty acids that are highly
saturated, mainly palmitic and stearic acids. The total amount of lipid entering the
duodenum often exceeds lipid intake and the greatest increase typically occurs with
high forage based diets. Dietary lipid supplements can result in higher, similar or lower
post-ruminal flows relative to fatty acid intake, due to the range of effects they can have
on microbial lipid synthesis (Demeyer and Doreau, 1999; Lock and Shingfield, 2003).
Approximately 80-90% of the lipid entering the small intestine is as free fatty acids
attached to feed particles (Davis, 1990; Doreau and Chilliard, 1997). The remaining lipid
components are microbial phospholipids plus small amounts of triglycerides and
glycolipids from residual feed material, and these esterified fatty acids are hydrolyzed by
intestinal and pancreatic lipases (Doreau and Ferlay, 1994). Even when protected fats
are fed to dairy cows, increases in the flow of triglycerides will be observed only when
encapsulated lipids are fed. Calcium salts of free fatty acids are the predominant source
of protected lipids fed to ruminants and these dissociate to some extent in the rumen,
but dissociation is much more extensive under the highly acidic conditions of the
abomasum. Therefore, dietary lipid supplements protected by the use of Ca salts add to
the pool of free fatty acids that enters the small intestine.
Fat absorption in the small intestine occurs in the jejunum and micelle formation is
the key to efficient fatty acid absorption in all species (Davis, 1990). In non-ruminants,
monoacylglycerols are required for micelle formation (Doreau and Chilliard, 1997).
However, due to the nearly complete hydrolysis of dietary fatty acids in the rumen these
are absent from digesta present in the small intestine of the ruminant. This is
compensated for by the secretions of bile and pancreatic juice prior to the jejunum
(Figure 3; Demeyer and Doreau, 1999). Bile supplies bile salts and lecithin, and
pancreatic juice provides enzymes to convert lecithin to lysolecithins and bicarbonate to
raise the pH (Davis, 1990). Lysolecithins, together with bile salts, desorb fatty acids
from feed particles and bacteria, and this allows formation of the micelles. Once
micelles are formed they are taken up by the epithelial cells of the jejunum where the
fatty acids are re-esterified into triglycerides and then packaged into chylomicrons
(Demeyer and Doreau, 1999).
Figure 3. Fat digestion in the small intestine of ruminants. Reproduced from Davis
(1990).
Dynamics of Absorption
Doreau and Ferlay (1994) carried out an extensive review of the literature on lipid
digestion using data obtained from fatty acid disappearance between the duodenum
and ileum or between the duodenum and feces. Studies based on whole tract
disappearance of lipid were not used because they typically underestimate fatty acid
digestibility in the small intestine and lead to questionable results (Doreau and Ferlay,
1994). As discussed earlier, this is mainly due to inaccurate estimations of the dietary
intake of fatty acids and because the flow of fatty acids to the duodenum is generally
greater than dietary intake of fatty acids. Doreau and Ferley (1994) found a wide
variation in the reported values for fatty acid digestibility, ranging from 55 to 92%, but
this range was not related to fatty acid intake. In fact, the capacity of the dairy cow to
absorb long-chain fatty acids is very high and can exceed 1 kg/day (Doreau and
Chilliard, 1997).
In monogastrics, digestibility of individual fatty acids decreases when chain length

increases, and increases as the number of double bonds increase (Lessire et al., 1992).
Although similar patterns are observed in ruminants, the differences are modest (Ferlay
et al., 1993). Digestibility does not differ significantly between 16 and 18 carbon
saturated fatty acids and is lower for longer chain saturated fatty acids as compared to
PUFA. The review by Doreau and Ferley (1994) reported that mean digestabilities were
0.77, 0.85, 0.83 and 0.76 for 18 carbon fatty acids with zero, one, two and three double
bonds, respectively. Another approach to investigate fatty acid digestibility is abomasal
infusion of fatty acids and these studies have generally given similar estimates of
digestibility. The exception is the digestibility of linolenic acid where abomasal infusion
studies observe a slightly higher value. This is probably related to inaccuracies of
measuring 18:3 with unsupplemented diets because the flow of this fatty acid leaving
the rumen is minimal.
Avila et al. (2000) evaluated the effects of supplemental fat from sources varying in
proportions of unsaturated and saturated fatty acids on fatty acid digestibility. Increasing
the degree of unsaturation of abomasally infused fat source did not affect the fatty acid
digestibility of cis-9 18:1, 18:2 or 18:3 fatty acids which averaged 0.67, 0.64 and 0.76
respectively. Interestingly, digestibility of trans 18:1 fatty acids was greater as compared
to cis-9 18:1, a finding also reported by Enjalbert et al. (1997). Thus, there may also be
small differences in the absorption of different 18:1 isomers, although data are limited.
This may even be an evolutionary development since ruminants typically are exposed to
very low amounts of cis 18:1 isomers entering the small intestine compared with the
amount of trans 18:1 isomers. With the recent improvements in analytical techniques,
differences in the digestibilities of individual fatty acids and fatty acid isomers can be
more thoroughly examined, but application in feeding systems still requires accurate
information on the profile of fatty acids leaving the rumen.
The overall conclusion is that differences in digestibility among individual fatty acids
contribute very little to the extensive variation (~60-90%) in the digestibility of dietary
lipids. Rather, the majority of this variation reflects differences among individual
experiments, and thus relates to differences in diets and specific feed components
(Demeyer and Doreau, 1999). Consequently, the composition of absorbed fatty acids is
close to the composition of fatty acids leaving the rumen (Doreau et al., 1997).
The common use of rumen-protected fat in the diets of dairy cows has raised an
interest in the effects of Ca-salts of fatty acids on fat digestion and absorption. In an
attempt to model ruminal metabolism and intestinal digestion of fatty acids in ruminants
Chalupa et al. (2003) proposed that digestion coefficients for fatty acids derived from
Ca-salts were substantially greater than coefficients for free fatty acids entering the
small intestine. However, we find little support for this in published literature. The overall
pattern is that the use of Ca-salts has little or no effect on the apparent digestibility of
individual fatty acids in the small intestine (Moller, 1988; Wu et al., 1991; Ferlay et al.,
1993; Enjalbert et al., 1997). Where minor differences exist, these are related to
differences in the fatty acid profile of the supplements being compared, and the
existence of small differences in digestibility among individual fatty acids as discussed
above. As an example of the comparisons, Enjalbert et al. (1997) supplemented a
control diet based on maize silage and concentrate with Ca-salts of either palm or
rapeseed oil and found no differences from the control in the digestibility of total fatty
acids in the small intestine (average 78.2% for all dietary treatments) and no significant
differences in apparent digestibility for individual fatty acids across treatments (Table 3).
Likewise, Ferlay et al. (1993) compared rapeseed oil supplied either as a Ca-salt (free
fatty acids) or as the oil (fatty acids present in triglycerides) and observed no differences
between treatments for fatty acid digestibility in the small intestine. The lack of any
effect of Ca-salts is not surprising because the Ca-fatty acid complex would be
dissociated to give free fatty acids in the abomasum (Grummer and Rabelo, 1999).
Table 3. Apparent digestibility in the small intestine (proportion of duodenal flow;

adapted from Enjalbert et al., 1997)
Ca-salts Differences
Palm fatty Rapeseed Salts v. Rapeseed
Control acids fatty acids SEM control v. palm
16:0 0.69 0.76 0.76 0.09 NS NS

18:0 0.79 0.79 0.79 0.04 NS NS
cis-9 18:1 0.77 0.78 0.86 0.07 NS NS
trans-11 18:1 0.84 0.88 0.89 0.05 NS NS
18:2 0.60 0.66 0.73 0.08 NS NS
18:3 0.67 0.63 0.71 0.07 NS NS
Total 18 0.79 0.79 0.81 0.04 NS NS
CONCLUSIONS
The increasing use of fats in dairy cattle diets and efforts to include lipid metabolism
in nutrition models require an understanding of their fate. The digestion and metabolism
of dietary fats and fatty acids in ruminants is complex and in this paper we have
provided an overview that highlights current limitations and recent advances. Routine
methods of dietary lipid analysis, specifically ether extract, provide an estimate of the
fatty acid content that varies in accuracy among feedstuffs. Although lipid hydrolysis and
classical pathways of fatty acid biohydrogenation are well established, analytical
improvements have revealed the complexity of lipid metabolism in the rumen. Clearly,
many pathways of biohydrogenation exist and many factors related to diet and rumen
environment affect this process. As a consequence, there are numerous fatty acid
intermediates produced during rumen biohydrogentation. Recent research has
established that some of these unique fatty acids are signaling molecules that regulate
metabolic processes in the cow and that others can have beneficial health effects when
consumed in dairy products. The rumen outflow of lipids are predominantly free fatty
acids and differences in the digestibility of individual fatty acids in the small intestine are
negligible. Thus, the composition of fatty acids absorbed in the small intestine is similar
to the composition of fatty acids leaving the rumen.
The post-absorptive metabolism of fatty acids is also a complex area where

advances in understanding have occurred, and this will have to be the subject of a
future paper. The post-absorptive use of fatty acids as a source of energy and for
synthesis of milk fat and body fat has been extensively investigated. However, the role
of fatty acids as signaling molecules involved in the expression of specific genes and
their regulation of metabolic processes is an emerging area, especially as it applies to
ruminants. This includes recent developments relating specific biohydrogenation
intermediates to the regulation of fat synthesis and the role of polyunsaturated fatty
acids in the signaling pathways involved in immune function and reproduction. There is
also growing literature on the role of specific fatty acids in human health and the
prevention of diseases, and this knowledge may offer an exciting opportunity to design
the fatty acid composition of ruminant-derived food products. Obviously, our knowledge
of lipid digestion and metabolism is rapidly advancing and the opportunity and challenge
is to effectively apply this knowledge in the feeding and management of todays high
producing dairy cows.
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Bauman, D. E., J. W. Perfield II, M. J. de Veth, and A. L. Lock. 2003.

NEW PERSPECTIVES ON LIPID DIGESTION AND METABOLISM IN RUMINANTS

D. E. Bauman, J. W. Perfield II, M. J. de Veth, and A. L. Lock

refining industries contain the lipid predominantly as triglycerides whereas rumen-

DIGESTION IN THE RUMEN

Figure 1. Fat digestion in the rumen. Reproduced from Davis (1990).

Biohydrogenation of unsaturated fatty acids is the second major transformation that

Classical pathways of biohydrogenation were established using pure cultures of

The initial step in rumen biohydrogenation typically involves an isomerization of the

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

(Group A) cis-9, trans-11, cis-15 C18:3

cis-9, trans-11 C18:2 trans-11, cis-15 C18:2

trans-11 C18:1 trans-15 and cis-15 C18:1

As analytical techniques improved, we have gained an appreciation of the complexity

Clearly, many intermediates are formed in the ruminal biohydrogenation of PUFA

ABSORPTION IN THE SMALL INTESTINE

In monogastrics, digestibility of individual fatty acids decreases when chain length

Table 3. Apparent digestibility in the small intestine (proportion of duodenal flow;

16:0 0.69 0.76 0.76 0.09 NS NS

The post-absorptive metabolism of fatty acids is also a complex area where

Allen, M. S. 2000. Effects of diet on short-term regulation of feed intake by lactating

Wu, Z., O. A. Ohajuruka, and D. L. Palmquist. 1991. Ruminal synthesis,

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