The Role of Microbes in Rumen Lipolysis and Biohyd

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The role of microbes in rumen lipolysis and biohydrogenation and their

manipulation
Article in animal · July 2010

DOI: 10.1017/S175173111000042X · Source: PubMed
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Animal (2010), 4:7, pp 1008–1023 & The Animal Consortium 2010
doi:10.1017/S175173111000042X
animal
The role of microbes in rumen lipolysis and biohydrogenation

and their manipulation
M. Lourenço1, E. Ramos-Morales2 and R. J. Wallace2-
1
Department of Animal Production, Ghent University, Laboratory for Animal Nutrition and Animal Product Quality, Proefhoevestraat 10, 9090 Melle, Belgium;
2
Gut Health Division, University of Aberdeen, Rowett Institute of Nutrition and Health, Greenburn Road, Bucksburn, UK
(Received 3 September 2009; Accepted 4 February 2010; First published online 23 March 2010)
Despite the fact that the ruminant diet is rich in polyunsaturated fatty acids (PUFA), ruminant products – meat, milk and
dairy – contain mainly saturated fatty acids (SFA) because of bacterial lipolysis and subsequent biohydrogenation of ingested PUFA
in the rumen. The link between SFA consumption by man and coronary heart disease is well established. In contrast, ruminant
products also contain fatty acids that are known to be beneficial to human health, namely conjugated linoleic acids (CLAs). The
aims of research in this field have been to understand the microbial ecology of lipolysis and biohydrogenation and to find ways of
manipulating ruminal microbes to increase the flow of PUFA and CLA from the rumen into meat and milk. This review describes
our present understanding of the microbial ecology of ruminal lipid metabolism, including some apparently anomalous and
paradoxical observations, and the status of how the metabolism may be manipulated and the possible consequential effects on
other aspects of ruminal digestion. Intuitively, it may appear that inhibiting the ruminal lipase would cause more dietary
PUFA to reach the mammary gland. However, lipolysis releases the non-esterified fatty acids that form the substrates for
biohydrogenation, but which can, if they accumulate, inhibit the whole process. Thus, increasing lipase activity could be beneficial
if the increased release of non-esterified PUFA inhibited the metabolism of CLA. Rumen ciliate protozoa do not carry out
biohydrogenation, yet protozoal lipids are much more highly enriched in CLA than bacterial lipids. How could this happen if
protozoa do not metabolise PUFA? The answer seems to lie in the ingestion of plant organelles, particularly chloroplasts, and the
partial metabolism of the fatty acids by contaminating bacteria. Bacteria related to Butyrivibrio fibrisolvens are by far the most
active and numerous biohydrogenating bacteria isolated from the rumen. But do we misunderstand the role of different bacterial
species in biohydrogenation because there are uncultivated species that we need to understand and include in the analysis?
Manipulation methods include dietary vegetable and fish oils and plant-derived chemicals. Their usefulness, efficacy and possible
effects on fatty acid metabolism and on ruminal microorganisms and other areas of their metabolism are described, and areas
of opportunity identified.
Keywords: biohydrogenation, cellulose digestion, lipase, microbial protein synthesis, rumen
Implications alter their activity. In this way, the healthiness of ruminant

products can be improved. Some methods may be doubly
The conversion of dietary polyunsaturated fatty acids (PUFA)
beneficial, by decreasing the environmentally damaging
to saturated fatty acids (SFA) by ruminants has important
methane and nitrogenous emissions from ruminant livestock
health implications for human health. However, ruminant
production.
products are also rich in conjugated linoleic acids (CLAs),
which have positive implications for human health. The
transformation from PUFA to SFA (biohydrogenation), and Introduction
the formation of CLA, is catalysed by the microorganisms
that inhabit the rumen. Researchers are finding ways to Human diets in industrialised countries are generally char-
manipulate biohydrogenation, by identifying the microorgan- acterised by high levels of saturated fat, n-6 fatty acids (FA)
isms responsible and by finding feed additives/ingredients that and trans-FA, and low levels of n-3 FA (Simopoulos, 2004).
World Health Organisation (WHO) (WHO, 2003) has recom-
mended that total fat, SFA and trans-FA should contribute
-
E-mail: [email protected] ,15% to 30%; ,10% and ,1% of total energy intake,
1008
Ruminal lipid metabolism
respectively. These recommendations arise from the fact that health-promoting CLA for human consumption (Givens and
SFA increase the risk of cardiovascular disease in humans, as Shingfield, 2004). The trans-11-18:1 FA, vaccenic acid (VA), is
well as increased plasma cholesterol levels. In contrast, also desirable as a product flowing from the rumen because
unsaturated FA (UFA) are known to decrease plasma VA acts as a substrate for the formation of cis-9,trans-11-18:2
cholesterol and low density lipoprotein-cholesterol (Givens, in the animal’s own tissues (Griinari et al., 2000).
2005). Ruminant products are characterised by high con- Despite rumen microbial activity, increasing dietary PUFA
centrations of SFA and low concentrations of UFA, compared intake enhances the PUFA content of ruminant meat and
with non-ruminants. Hence they are often regarded as milk (Dewhurst et al., 2006; Scollan et al., 2006). Different
detrimental to human health. However, the FA composition nutritional strategies have been used, such as forage feed-
of ruminant products can be improved to meet the WHO ing, supply of vegetable oils or oilseeds, marine products or
recommendations. The saturation of fats is a direct con- protected fat sources. Antimicrobial feed additives such as
sequence of the reduction (biohydrogenation) of UFA by monensin may be effective via their influence on the com-
ruminal microorganisms. Thus, improvement of the FA profile position of the microbial community. In fact, dietary strate-
of ruminant products can be achieved by two distinct gies that involve altering the FA composition of the diet
approaches: (i) modification of the FA profile during meat or often combine the benefits of bypass and manipulation of
milk processing or (ii) modification through the changes in biohydrogenating bacteria. UFA have a stronger anti-
animal diet. The latter might simply result in greater bypass microbial effect than saturated ones (Harfoot and Hazle-
of dietary FA from the rumen, or might be a consequence of wood, 1997) and different PUFA have been reported to
altered microbial metabolic activity. These will be discussed have differential toxicity toward rumen microorgani-
later in this review. sms (Maia et al., 2007; Zhang et al., 2008). Therefore, lipid
Ruminant products also contain potentially health- supplementation can lead to a shift in the rumen
promoting CLA (mainly cis-9,trans-11-18:2). Dietary CLA microbial population. For any dietary strategy to be useful,
have been shown in many animal studies to contribute to it must not compromise rumen fermentation and,
cancer prevention, decreased atherosclerosis, improved concomitantly, dry matter intake and animal production
immune response and altered protein/energy metabolism and/or performance. Often, studies report the effect of
(Whigham et al., 2000; Pariza, 2004; Palmquist et al., 2005). different nutritional strategies on lipid metabolism, but
CLA are present at higher concentrations in ruminant pro- no further information is given on the impact of a parti-
ducts than in corresponding meats from non-ruminants or in cular strategy on the whole ruminal function and processes
vegetable oils (Chin et al., 1992; Givens and Shingfield, (Figure 1). Here, we attempt to review the interactions
2004), as they are produced from the partial biohydrogena- between lipid metabolism, other aspects of rumen meta-
tion of linoleic acid (LA) and linolenic acid (LNA) in the rumen bolic function and the ruminal microbial community,
by ruminal microorganisms (Chilliard et al., 2007). The particularly the consequences of strategies that aim to alter
cis-9,trans-11 isomer is generally considered to be the main biohydrogenation.
Lipids Fibre
Lipid supplements
Lipolysis Plant secondary metabolites Pentoses, hexoses
Botanical diversity
UFA Pyruvate
CO2 + H2
Microbial
structure/activities
Biohydrogenation
CH4
SFA Acetate Propionate

Butyrate
NH3 + VFA
Aminoacids Peptides Protein
Microbial protein
Figure 1 Interventions to manipulate lipid metabolism in the rumen inevitably lead to effects on other processes. Sometimes the target organisms have
several functions, in other cases the metabolic pathways are linked, for example by the availability of H2. UFA 5 unsaturated fatty acid; SFA 5 saturated fatty
acid; VFA 5 volatile FA.
1009
Lourenço, Ramos-Morales and Wallace
Ruminal microbial processes in monitoring other effects. The secondary effect may not
necessarily be detrimental, however. As will be reviewed
The rumen evolved to slow down the passage of fibre- below, some of the feed composition changes that lead to a
containing foodstuffs through the gut, which enables better FA flow from the rumen also decrease methanogen-
microbial enzymes time to digest the constituent digestion- esis, which benefits the animal through improved energy
resistant polymers, mainly cellulose and xylan (Hungate, retention and benefits the environment by lowering the
1966). Mammalian enzymes cannot break down cellulose or emission of a greenhouse gas.
xylan. To achieve this digestion, the microbes must ferment
the released sugars to release ATP, which in turn fuels their
growth. In this anaerobic environment, the main products of Lipase activity in the rumen
the metabolic pathways that generate ATP (Figure 1) are
volatile FA (VFA), such as acetate, propionate, butyrate, and Most research on ruminal lipolysis was carried out many
gases such as CO2 and CH4 (Russell and Wallace, 1997). CH4 years ago. Recently, interest has revived because of the
is now well recognised as a greenhouse gas, a significant implications of ruminal lipolysis on subsequent biohy-
contributor to global warming (Lassey, 2008). Also essential drogenation of PUFA and generation of CLA.
for microbial growth is nitrogen, mainly in the form of pro-
tein in plants that form the bulk of the diet (Figure 1). Thus, Plant and microbial lipases
proteolysis and amino acid transport are essential for Lipolysis occurs rapidly in ruminal digesta (Garton et al., 1958;
microbial growth. In fact, microbial proteolysis is generally Dawson and Hemington, 1974; Dawson et al., 1977). In ani-
considered wasteful as far as the animal is concerned, mals receiving cereals and plant oils, the most abundant lipids
because its activity generally exceeds the capability of the would be in the form of triacylglycerol. Hydrolysis of tria-
microbes to utilise the products of hydrolysis (Walker et al., cylglycerol in these diets occurs predominantly by microbial
2005). The excessive catabolism leads to one of the major lipases (Dawson et al., 1977). The forage consumed by grazing
nutritional losses (and pollutants) from animal agriculture, ruminants in contrast contains little triacylglycerol, comprising
namely N excretion (Pfeffer and Hristov, 2007). mainly galacto-, sulfo- and phospholipids (Harfoot, 1981).
The metabolism of dietary lipid is not, in contrast, seen as Forage plant tissues are rich in galacto- and phospholipases.
an activity essential to provide nutrients to ruminal micro- These lipases remain active once ingested into the rumen for
organisms. The microorganisms are capable of synthesising up to 5 h, suggesting that the plant material itself may con-
their own FA and indeed do so extensively in the mixed tribute to ruminal lipolysis in grazing animals (Omar Faruque et
community (Garton, 1977). They also cannot derive energy al., 1974). Dawson et al. (1977) challenged this idea and
from b-oxidation, which does not occur anaerobically. Lipid concluded that microbial lipases were more important than
metabolism may, therefore, seem to be somewhat peripheral plant enzymes. The experiments of Dawson et al. (1977) used
to the main growth-generating activities of the rumen autoclaved grass as substrate, which is not ideal because of
microbial community. Nevertheless, it is extremely important the many effects that autoclaving has other than enzyme
from the microrganisms’ perspective in that it enables some denaturation, a conclusion made by the authors themselves.
of them to survive what would otherwise be a toxic chal- More recently, this issue has been revisited by Lee et al. (2002),
lenge. It is also fundamental in influencing the nutritional reporting increased free FA and decreased polar lipids after 6 h
quality of ruminant products. incubation of fresh ryegrass leaves in buffer, confirming that
The main members of the microbial community are bac- plant catalysed lipolysis occurred. Further, Van Ranst et al.
teria, archaea, protozoa and fungi. Bacteria are the most (2009) reported plant lipolysis up to 60% after 8 h incubation
abundant, followed by archaea (the CH4 producers), ciliate of fresh red clover leaves. Both papers suggested the observed
protozoa and in lower numbers the anaerobic fungi. Differ- lipolysis to be due to active plant lipases that could have some
ent species have different roles, which interact and are contribution to the overall ruminal lipolysis, though neither
essential for sustaining the microbial community and its compared plant activity with that of ruminal microorganisms.
collective activity. Furthermore, with the exception of CH4 We believe it is time for this issue to be revisited, given the
formation, individual genera or species seldom have a single importance of these esterases in the overall lipid metabolism
role. For example, the Gram-positive bacterium, Butyrivibrio and its implications for product quality and human health. How
fibrisolvens, is a key player in fibre digestion, but many important are plant lipases in comparison with microbial lipa-
strains are also highly proteolytic (Stewart et al., 1997). ses in ruminal metabolism? It may be a useful objective to
B. fibrisolvens also dominates the community in the biohy- breed forage plants low in lipase activity.
drogenation of FA. Thus, targeting one microbial activity for
manipulation always has consequences for others. Also, the Lipolytic ruminal microorganisms
way in which the manipulation is implemented seldom has Among the various types of ruminal microorganisms, the
influence on only the intended target. Cellulolysis is parti- bacteria are considered to be most active in lipolysis. This
cularly vulnerable to disruption, for example (Weimer, 1996; view was based on experiments with ciliate-free sheep, in
Chesson and Forsberg, 1997). Thus, any attempt to alter one which hydrolysis of phosphatidyl choline remained high
aspect of ruminal activity must be accompanied by diligence (Dawson and Kemp, 1969), and on fractionations carried out
1010
during the PhD studies of Hazlewood (Harfoot and Hazle- reported to liberate galactose from galactolipids, suggesting
wood, 1997). Again, we believe that it is important to carry galactosidase activity, although lipase activity was not
out further studies with up-to-date methods to assess again demonstrated (Bailey and Howard, 1963). Another protozoal
the possible role of protozoa in lipolysis, given their impor- species, Entodinium caudatum, was shown to have phos-
tance in the flow of PUFA, including CLA, from the rumen pholipase activity (Coleman et al., 1971), but it is most likely
(Devillard et al., 2006; Yáñez-Ruiz et al., 2006). As for the that this activity was more relevant to the internal economy
anaerobic fungi, their existence was not known when the of the protozoa than to the digestion of dietary lipids. The
vast majority of early studies were carried out. They merit earlier studies to determine the contribution of protozoa to
further investigation. the lipolytic activity in the rumen were carried out with
fractionated rumen fluid, with the possibility that lipolytic
Lipolytic ruminal bacteria and their lipases activity in protozoal fractions was more because of the
The most active bacterial species isolated selectively using activity of bacteria that the protozoa had ingested than that
triacylglycerol as substrate was Anaerovibrio lipolytica of the protozoa themselves.
(Hobson and Mann, 1961; Henderson, 1968). Its lipase
activity was investigated in some detail by methods avail- Lipases from the rumen microbial metagenome
able at the time, the 1970s (Henderson, 1971; Henderson Now we take a huge step forward in time to the present day
and Hodgkiss, 1973). A. lipolytica would be expected to of pyrosequencing, metagenomics, sequence databases and
dominate ruminal lipase activity in animals receiving mainly analogous technologies. Liu et al. (2009) screened a meta-
concentrate feeds, but, because A. lipolytica lacks the ability genomic library from the rumen of grazing Holstein cows for
to hydrolyse galacto- and phospholipids, other lipolytic spe- lipase activity, using trioleoylglycerol as substrate. Out of
cies would be expected to predominate in grazing animals. 15 360 bacterial artificial chromosome (BAC) clones, two
Nevertheless, Prins et al. (1975) found that A. lipolytica was were found with high lipase activity. This number seems
present at around 107/ml in grazing animals, reflecting per- surprisingly small, nevertheless the sequences provided
haps other activities, such as the fermentation of glycerol, some interesting information. The genes identified and
that may provide this species with a selective niche (Rattray expressed in Escherichia coli coded for esterases with a
and Craig, 2007). Several molecular microbial ecology stu- preference for long-chain FA, indicating that they were
dies have enumerated A. lipolytica in the rumen under dif- genuine lipases, but no further substrate specificity was
ferent dietary conditions (Tajima et al., 2001; Koike et al., tested. The likely origin of the genes was investigated, based
2007), with similar conclusions. Phospho- and galactolipids on the other open reading frames present in the BAC clones.
seem to be hydrolysed by Butyrivibrio-like species (Hazlewood It was impossible to tell the host for the first gene, Rlip1,
and Dawson, 1975 and 1979). A. lipolytica did not break down but the second, Rlip2, gave most similar homologues from
phospho- and galactolipids (Henderson, 1971), while the Thermosinus carboxydivorans, which has not previously
Butyrivibrio spp. did not break down triacylglycerols. The been associated with the ruminal ecosystem. Thus, once
Butyrivibrio spp. appeared to contain all the phospholipase A, again, the importance of these lipases in the overall com-
phospholipase C, lysophospholipase and phosphodiesterase munity is far from certain. Clearly many more lipase
activities typical of the mixed rumen contents (Harfoot and sequences must be analysed from the metagenome to
Hazlewood, 1997). The fact that these bacteria also possess understand the true nature of lipolytic enzymes in the rumen.
the ability to biohydrogenate UFA suggests that the two The potential for new discovery seems high, however,
properties are linked in a way that benefits their ‘biochemical because the Rlip1 lipase sequence clustered separately from
economy’ (Harfoot and Hazlewood, 1997). Specifically, the eight main families of lipase as identified by Arpigny and
because Butyrivibrio S2 was a FA auxotroph, it would require Jaeger (1999) and may be a member of a new family of
lipase to grow. The toxicity of non-esterifed PUFA (Maia lipolytic enzymes. One might argue that going to the meta-
et al., 2007) released by the lipase would then have to be genome before interrogating the genome of known lipolytic
removed by biohydrogenation. The importance of these species is premature, and that the pure culture studies
various species vis-à-vis other species that may not yet have should be carried out at least alongside metagenomics.
been isolated remains to be quantified. For example, two
lipolytic isolates were obtained from young deer, but they
Biohydrogenating activity in the rumen
were derived from such low dilutions that their population
size may be very small and their importance is uncertain The main FA substrate for biohydrogenation in grazing ani-
(Jarvis et al., 1998). mals is LNA (cis-9,cis-12,cis-15-18:3), because it is the most
abundant FA present in glycolipids and phospholipids of
Lipolytic ruminal protozoa and their lipases grass and other forages, whereas for animals receiving
Evidence of lipolytic activity in protozoa is not very con- dietary lipid supplements, LA (cis-9,cis-12-18:2) in the form
sistent, partly because there have been few recent studies of triacylglycerols will usually be the main substrate for
that investigate protozoal lipolysis. Wright (1961) suggested biohydrogenation. LA metabolism in the rumen involves the
Epidinium spp. to be responsible for 30% to 40% of the transient formation of CLA, mainly cis-9,trans-11-18:2 or
lipolytic activity in the rumen. Epidinium ecaudatum was rumenic acid, which is then converted to VA, and finally
1011
c9c12c15-18:3
c9t11c15-18:3
c9c12-18:2
8,10-CLA 10,12-CLA c9,t11 other other t11c15 c9t13 11,13-CLA

9,11-CLA 9,12-18:2
t10-18:1 c12-18:1 t11-18:1 t4-t9-18:1 t15+c15-18:1 t13+14-18:1
18:0
Figure 2 Biohydrogenation pathways in the rumen. LA – c9c12-18:2; LNA – c9c12c15-18:3; VA – t11-18:1. CLA 5 conjugated linoleic acid; LA 5 linoleic
acid; LNA 5 linolenic acid; VA 5 vaccenic acid. Adapted from Chilliard et al. (2007).
stearic acid (18:0; Figure 2). LNA is metabolised in a similar pathways of LA metabolism, including hydration and
way (Figure 2), though as there are three double bonds to be chain elongation or shortening, which may increase in
reduced the pathway is slightly more complicated (Harfoot importance depending on the diet. In all these aspects of FA
and Hazlewood, 1997; Jenkins et al., 2008). The main 18:3 metabolism it is important to understand the role of different
intermediate arising from isomerisation of LNA in mixed microbial species.
ruminal digesta was shown recently (Wa˛sowska et al., 2006)
to be the conjugated triene, cis-9,trans-11,cis-15-18:3, one Role of ciliate protozoa in biohydrogenation
of the possible conformations proposed by Wilde and Daw- Up to half of the rumen microbial biomass may be protozoal
son (1966). Other intermediates identified included trans- in origin (Williams and Coleman, 1992) and about three
9,trans-11,cis-15-18:3 and trans-11,cis-15-18:2. Conjugated quarters of the microbial FA present in the rumen may be in
trienes may have just as important health implications as protozoa (Keeney, 1970). Thus, protozoa could represent a
CLA (Tsuzuki et al., 2004) although much less work has been very important source of CLA and VA. Wright (1959 and
done on the trienoic than the dienoic fatty C-18 FA. 1960) concluded that both protozoa and bacteria were
cis-9,trans-11-18:2 is usually the predominant CLA isomer involved in biohydrogenation, but the extensive ingestion of
found in the rumen and in milk, but many others are present, bacteria by protozoa was considered by Dawson and Kemp
with trans-9,trans-11-18:2 usually most abundant of the (1969) to cast doubt on this conclusion. Biohydrogenation in
others (Shingfield et al., 2003; Palmquist et al., 2005; Chil- ruminal digesta was only slightly decreased after defauna-
liard et al., 2007). There are, however, times when the trans- tion and the presence of protozoa was not necessary for
10,cis-12 isomer becomes a major intermediate. This can be biohydrogenation to occur (Dawson and Kemp, 1969). Girard
precipitated by high-starch feeding or by fish or vegetable oil and Hawke (1978) and Singh and Hawke (1979) also sug-
supplementation (Bauman and Griinari, 2001; Shingfield and gested that the minor contribution of protozoa to the bio-
Griinari, 2007). High concentrations of trans-10-18:1 occur in hydrogenation process was due to the activity of ingested or
digesta and consequently in the FA flowing to animal tissues associated bacteria. It has been known for a long time that
(Offer et al., 2001; Daniel et al., 2004; Shingfield and Grii- protozoal lipids contain proportionally more UFA than the
nari, 2007). Under these circumstances, milk fat depression bacterial fraction (Katz and Keeney, 1966; Harfoot and
occurs, with other consequences to the animal, including Hazlewood, 1997). Recently, it was established that these
lower intake and decreased fibre digestion (Bauman and UFA include CLA and VA (Devillard et al., 2006), further
Griinari, 2001). Post-ruminal infusion experiments first indi- increasing the possible significance of protozoa in the
cated that trans-10,cis-12-18:2 exerts anti-lipogenic effects delivery of health-promoting FA from the rumen. Different
in the lactating cow (Baumgard et al., 2000; Saebø et al., protozoal species had different composition, with larger
2005; Lock et al., 2006). Recent studies suggest that it may species including Ophryoscolex caudatus containing more
actually be trans-10-18:1 rather than the trans-10,cis-12 CLA than 10 times higher concentrations of CLA and VA than
that decreases mammary lipogenesis (Shingfield et al., some small species, such as Entodinium nannelum (Devillard
2009). It is important, therefore, to understand how these et al., 2006). Isotricha prostoma, a large species and the only
isomers are formed. There are, in addition, other possible holotrich examined, had low concentrations of CLA and VA.
1012
In incubations with fractionated ruminal digesta, LA meta- to be 12% to 15% protozoal in origin, whereas in terms of FA
bolism was very similar in strained ruminal fluid and its flow, protozoa accounted for between 30% and 43% of the
derived bacterial fraction, while its mixed protozoal fraction CLA and 40% of the VA reaching the duodenum (Yáñez-Ruiz
had much lower activity. The opposite direction of reaction, et al., 2006). The contribution of protozoa to the flows of
namely desaturation, also did not occur in the protozoal 16:0 and 18:0 to the duodenum was less than 20% and
fraction. Radioactivity from 14C-stearate was not incorpo- 10%, respectively. Thus, even if protozoa do not themselves
rated into CLA or VA by protozoa. No genes with sequence produce CLA and VA by their own metabolism, nevertheless
similarity to FA desaturase genes from other organisms were they might be expected to have a significant influence on
found in cDNA libraries from ruminal protozoa (E. Devillard, CLA and VA available to the host animal.
personal communication). Thus, the protozoa are rich in
CLA and VA, yet they appear not to synthesise these two FA Biohydrogenating ruminal bacteria
from LA or stearate, confirming the opinion of Dawson and Bacteria play the main role in FA biohydrogenation (Jenkins
Kemp (1969). et al., 2008). In early microbiological studies (Polan et al.,
It might be argued that the high-UFA content of protozoa 1964), B. fibrisolvens was identified to undertake biohy-
results from the ingestion of plant particles, especially drogenation of FA and to form CLA and VA as intermediates
chloroplasts (Wright, 1959; Stern et al., 1977). A recent during the biohydrogenation of LA (Polan et al., 1964; Kepler
study by Huws et al. (2009a) showed that the engulfment of et al., 1966). Stearic acid was not formed from LA by B.
chloroplasts is a major contributor to the high LNA con- fibrisolvens, however. Later studies (Kemp et al., 1975;
centration of protozoa. This cannot explain the high con- Hazlewood et al., 1976) identified other bacteria that were
centration of CLA and VA in protozoa, however, as these FA capable of biohydrogenation, but they did not provide much
are absent from the plant material. Biohydrogenating activ- information about relative activities: the method used radio-
ity must be involved. Our investigations suggest that the labelled substrates and was highly sensitive but the con-
most likely explanation is that protozoa preferentially incor- centrations of labelled acids used as substrates were very
porate CLA and VA formed by ingested bacteria. The lower low (2 mg/ml), making comparing the activity and quantita-
conversion to stearate perhaps occurs because the bacteria tive significance of different bacteria difficult. Bacteria car-
responsible for conversion of monoenoic acids to SFA are rying out stearate formation were identified as Fusocillus
more vulnerable to protozoal digestive activities. A simpler spp. (Kemp et al., 1975). Van de Vossenberg and Joblin
explanation is that the reactions carrying out the early stages (2003) isolated from a grazing cow a bacterium, which could
of biohydrogenation are much more active than the last one, also form stearate from linoleate. It was phenotypically
and that if both are decreased by, say, 95%, by protozoal similar to ‘Fusocillus’ and their analysis indicated that it was
digestive activities, this may leave remaining enzymic activ- phylogenetically close to Butyrivibrio hungatei. Subse-
ity sufficient to form CLA and VA from LA, for example, but quently, a species named Clostridium proteoclasticum was
not enough to form significant amounts of stearate. We identified as a stearate producer with morphological and
believe that the antibiotic treatment described by Or-Rashid metabolic properties that were indistinguishable from those
et al. (2008), which resulted in increased accumulation of reported for Fusocillus (Wallace et al., 2006). Moon et al.
CLA in washed protozoa, reflects a similar phenomenon, and (2008) renamed C. proteoclasticum as Butyrivibrio proteo-
is not caused by genuine protozoal activity. Until the genes clasticus from its 16S rRNA gene sequence. For many years,
encoding the enzymes involved in both bacteria and proto- the bacteria that are involved in the different steps of the
zoa are identified, it will be difficult to resolve the issue biohydrogenation process were classified as group A and B
unambiguously, however. A recent paper by Boeckaert et al. (Harfoot and Hazlewood, 1997). Group A bacteria hydro-
(2009) showed that I. prostoma did not hydrogenate LA, but genated LA and LNA to VA, whereas group B bacteria con-
in view of its low CLA and VA concentrations, this observa- vert the same FA to stearic acid. We propose that it is now
tion can probably not be extrapolated to other species, par- much more appropriate to describe the bacteria based on
ticularly entodiniomorphs. their correct taxonomy (Figure 3). B. hungatei and B. pro-
These findings imply that the availability of PUFA teoclasticus groups are much more sensitive to the toxic
(including CLA) and VA, for absorption by the host animal effects of UFA than the rest of the Butyrivibrio/Pseudobu-
could depend more on the flow of protozoa rather than tyrivibrio cluster, such that their isolation from media con-
bacteria from the rumen. Some ciliate protozoa are retained taining UFA is made very difficult (Wallace et al., 2006;
selectively within the rumen by a migration/sequestration Paillard et al., 2007). They can also be distinguished based
mechanism that depends on chemotaxis (Abe et al., 1981; on the mechanism by which they form butyrate, their main
Ankrah et al., 1990). As a consequence, protozoal biomass fermentation product after acetate. B. hungatei and B. pro-
reaching the duodenum is proportionally less than would be teoclasticus had a butyrate kinase activity .600 U/mg pro-
expected if they were to flow with the rest of ruminal digesta tein, while the others had much lower activity (Figure 3;
(Hungate et al., 1971; Weller and Pilgrim, 1974). It might be Paillard et al., 2007). The butyrate kinase gene was present
imagined that this selective retention would be detrimental in B. hungatei and B. proteoclasticus but not in the other
to the flow of CLA and VA into meat and milk. The flow of group. It is tempting to suggest that the different sensiti-
microbial N at the duodenum of steers was recently shown vities to the toxic effects of UFA may be linked to the enzymic
1013
Butyrivibrio Produce
Growth
proteoclasticus 18:0
inhibited by
<200 µg/ml LA;
butyrate kinase
Butyrivibrio >600 U/mg
hungatei
CLA, TVA
formed
from LA
Growth not
Butyrivibrio
inhibited by
fibrisolvens and
<200 µg/ml LA;
Pseudobutyrivibrio
butyrate kinase
spp.
<40 U/mg
Figure 3 Relation between phylogenetic position, formation of biohydrogenation products from LA, sensitivity to growth inhibition and mechanism of
butyrate formation in the Butyrivibrio phylogenetic tree. From Wallace et al. (2006), Maia et al. (2007) and Paillard et al. (2007). CLA 5 conjugated linoleic
acid; VA 5 vaccenic acid; LA 5 linoleic acid.
mechanism by which butyrate is produced. Intracellular acyl- Anaerobic ruminal fungi

CoA concentrations, principally the precursors of butyrate, in Although ruminal fungi produce cis-9,trans-11-18:2 from LA
B. fibrisolvens seem to be particularly sensitive to the effects (Nam and Garnsworthy, 2007a and 2007b), their activity is very
of LA (Maia et al., 2010). low in comparison with B. fibrisolvens (Maia et al., 2007).
Metabolism of LA by Butyrivibrio results in the formation of
cis-9,trans-11 CLA and VA, but no trans-10,cis-12 CLA or trans- Correlating microbial ecology and biohydrogenation
10-18:1 is formed (McKain et al., 2010). The bacteria respon- Attempts to correlate the microbial community composition
sible for milk fat depression must therefore be different to the in vivo with the observed inhibition of biohydrogenation
Butyrivibrio spp. The formation of trans-10,cis-12 CLA also have had limited success. Kim et al. (2008) and Huws et al.
occurs by a different enzymic mechanism to that of cis-9,trans- (2009b) used group-specific PCR to analyse the bacterial
11 CLA (Wallace et al., 2007). Enrichment cultures with starch communities in cattle in which biohydrogenation and the FA
carried out by Kim et al. (2002) contained abundant large cocci composition of digesta had been altered by incremental
identified as Megasphaera elsdenii that formed trans-10,cis-12 amounts of fish oil. Only weak correlations were found
CLA. Our studies indicate that Propionibacterium acnes may be between numbers of B. proteoclasticus and the duodenal
responsible for the formation of trans-10,cis-12-18:2 (Devillard flow of 18:0 in fish oil supplemented diets. Boeckaert et al.
and Wallace, 2006), having been unable to find comparable (2008) performed a similar analysis in dairy cows receiving
activity in M. elsdenii. Furthermore, when digesta samples docosahexaenoic acid (DHA)-containing microalgae. Their
from cows producing high amounts of trans-10,cis-12-18:2 dendrogram based on Butyrivibrio-related bacteria indicated
were analysed for M. elsdenii by qPCR of 16S rRNA genes, that changes occurred in a group from which there are no
numbers were ,103/g, while much larger numbers of P. acnes cultivated strains. These three reports indicate that other, as
were detectable (unpublished observation). P. acnes does not, yet uncultivated, microbial species may be involved in bio-
however, convert trans-10,cis-12 CLA to trans-10-18:1 (McKain hydrogenation, particularly the final step, during the con-
et al., 2010). This can be accomplished by the Butyrivibrio spp. version of VA to 18:0.
(McKain et al., 2010). Weimer et al. (2010) observed major
changes in the bacterial community associated with cows
suffering milk fat depression. A future challenge will be to
Manipulating ruminal biohydrogenation
associate with certainty the role of individual microbial species
with this disorder. Several factors influence the concentration of FA in ruminant
There are often significant differences between bacterial food products. The quantity and composition of dietary lipids
communities attached to solids and those ‘planktonic’ have a major effect because of the FA that escape ruminal
communities that inhabit the liquid phase of digesta. The metabolism. FA may also have a direct manipulating effect,
same is true for biohydrogenation, it appears, with the however, whereby they inhibit biohydrogenation. Biohy-
planktonic community carrying out LA metabolism only drogenation is affected indirectly too when other activities
as far as 18:1 FA while the solids-associated community are changed, because FA metabolism is inextricably linked to
formed 18:0 (Boeckaert et al., 2009). B. proteoclasticus was other areas of ruminal metabolism, through a common reli-
present only in the solids-associated community, at 12% of ance on H2 metabolism and/or the microbial species that are
Butyrivibrio-related clones. involved in multiple metabolic processes. FA metabolism
1014
in animal tissues, particularly the mammary gland, has by Franklin et al. (1999). Not only was DHA increased, but
also been targeted as a means of altering FA composition CLA and VA also increased. VA increased more when
in ruminant products. Enhancing the activity of mammary unprotected rather than rumen-protected microalgae were
D9-desaturase activity would increase the conversion of fed, indicating a ruminal effect whereby biohydrogenation of
trans-11-18:1 into cis-9, trans-11 CLA. We shall deal only VA was inhibited by the microalgae. Another DHA-contain-
with ruminal events here, focussing on effects on biohy- ing microalgal preparation inhibited biohydrogenation in
drogenation and microbial ecology. vitro (Boeckaert et al., 2007a) and increased the CLA and VA
content of milk (Boeckaert et al., 2008). Ciliate denaturing
gradient gel electrophoresis profiles (uses 18S rRNA genes
Lipid supplements to assess diversity in the mixed community) suggested a
Fats in ruminant diets are used for two main purposes: (i) to suppression of the protozoal community, with a decreased
increase the energy content of the diet because of their high abundance of the holotrichs, Isotricha prostoma and I.
caloric density or (ii) to manipulate ruminal fermentation intestinalis, and some entodiniomorphs, Epidinium cauda-
because of their antimicrobial effect. The meta-analysis by tum, Eudiplodinium maggii and Diplodinium dentatum, in
Glasser et al. (2008) provides the most comprehensive the rumen of algae-fed cows (Boeckaert et al., 2007b).
analysis of oilseed lipid supplements and their influence on Butyrivibrio-related bacterial community was also changed
milk fat composition. The antimicrobial effect of dietary significantly (Boeckaert et al., 2008).
lipids is associated with the degree of unsaturation of the FA Coconut and palm oils are usually considered separately
present (Szumacher-Strabel et al., 2004; Váradyová et al., from other oils. Coconut oil is rich in lauric (12:0) and myristic
2007; Zhang et al., 2008; Yang et al., 2009). PUFA are more (14:0) acids and palm oil in palmitic acid (16:0). Their main
toxic for biohydrogenating bacteria than di- or monoenoic FA application when supplemented to ruminant diets has been
(Maia et al., 2007 and 2010), thus oils containing PUFA to suppress methanogenesis. It has been suggested that
such as LNA would be expected to have a greater effect on coconut oil could be the natural alternative to ionophores,
rumen biohydrogenation and fermentation processes than which decrease ruminal methane and ammonia productions
those rich in LA or oleic acid. Unsaturated oilseed products, (Yabuuchi et al., 2006). Some of the SFA of coconut and palm
including linseed, soybean and sunflower seeds and oils oils pass into adipose, without affecting total SFA compared
(including Ca salt and amides), increased trans-18:1 FA, this with control (Jordan et al., 2006). Palm oil supplementation
increase being more pronounced with the oils, particularly increased milk total SFA, while decreasing total mono-
linseed and sunflower. cis-9,trans-11 CLA in milk was sig- unsaturated FA and cis-9,trans-11-18:2 concentrations
nificantly increased by dietary seeds and oils (Glasser et al., (Mosley et al., 2007). Thus, this group of oils probably do not
2008). Oils rich in LA (sunflower and soybean) have been benefit FA composition of ruminant products, despite the
more effective in enhancing milk CLA than oils rich in LNA inhibitory effect of lauric acid (12:0) on B. fibrisolvens
(linseed; Kennelly, 1996; Bu et al., 2007). The same meta- (Henderson, 1973). In contrast, Goel et al. (2009) found that
analysis showed that, regarding total 18:2 FA, increases capric acid (10:0) inhibited biohydrogenation in vitro, as well
were observed with linseed, soybean and sunflower, for both as lipolysis.
seeds and oils, whereas 18:3 FA increased with linseed and The effectiveness of lipid supplements in controlling bio-
soybean seeds (Glasser et al., 2008). hydrogenation and on nutrition in general varies with the
Fish oil, rich in long-chain FA, especially 20:5n-3 eicosa- composition of the basal diet, the physical form of the sup-
pentaenoic acid (EPA) and 22:6n-3 (DHA), has been particularly plement, and the inclusion rate. Negative effects of UFA on
successful in inhibiting the last step of biohydrogenation, ruminal digestion tend to be minimised, if the diet contains
the conversion of trans-11-18:1 to 18:0 (Lee et al., 2005; a high proportion of forage, because of the ability of forage
Wa˛sowska et al., 2006; Chilliard et al., 2007). EPA and DHA to promote normal rumen function for maximum biohy-
inhibit biohydrogenation directly, possibly by competitive inhi- drogenation (Palmquist, 1988). The dietary forage : con-
bition (AbuGhazaleh and Jenkins, 2004; Wa˛sowska et al., centrate ratio, as well as the type and physical state of the
2006). They also inhibit the growth of biohydrogenating bac- forage fed to the animals, will influence ruminal lipid meta-
teria and other bacteria in pure culture (Maia et al., 2010). As a bolism (French et al., 2000). Milk CLA content is increased to
consequence, bacterial communities are changed significantly a greater extent by feeding processed seeds compared with
in response to dietary fish oil (Kim et al., 2008; Huws et al., feeding intact vegetable oil seeds, in which access by
2009b). Protozoal numbers may be increased, further restricting microorganisms to the lipid is restricted (Kennelly, 1996;
biohydrogenation (Loor et al., 2005). In contrast, oil prepared Dhiman et al., 2000). The inclusion rate is crucial, as detailed
from catfish, a freshwater species, contains little DHA or EPA in meta-analyses (Glasser et al., 2008; Schmidely et al.,
because the fish do not consume marine algae (Amorocho 2008). Prolonged lipid supplementation also carries with it
et al., 2009). Catfish oil had no influence on the CLA or VA the danger that biohydrogenation shifts to the production
composition of milk or on protozoal numbers in the rumen. of trans-10,cis-12-CLA and trans-10-18:1, which in turn
Microalgae are an alternative, more sustainable, form of leads to milk fat depression (Bauman and Griinari, 2001;
additive that contains long-chain PUFA similar to fish oil. The Shingfield et al., 2006; Shingfield and Griinari, 2007), as
benefits of microalgae to milk fat composition were reported described above.
1015
Botanical feed additives/ingredients and phytochemicals ruminal biohydrogenation because of their antibacterial
In the aftermath of the ban on antimicrobial growth pro- activity. Nevertheless, the antimicrobial activity of essential
moters in the EU, much interest has turned to plants and oils might not always persist in the ruminal ecosystem
their constituent biologically active chemicals, evolved to because of their degradation (Broudiscou et al., 2007; Mal-
resist microbial, insect or animal attack. Biohydrogenation is ecky and Broudiscou, 2009; Malecky et al., 2009). More
a particularly appropriate target for these chemicals. studies are required, therefore, to determine if essential oils
Forage feeding has been described as one of the best or essential oil compounds have value in manipulating bio-
strategies to increase n-3 PUFA, VA and cis-9, trans-11-CLA hydrogenation.
in ruminant milk and meat, partly because forages are a The effects of saponins on ruminal ammonia concentra-
major source of LNA and to a lesser extent LA (Dewhurst tions and VFA production were reviewed by Wina et al.
et al., 2006; Elgersma et al., 2006; Scollan et al., 2006). (2005). A major component of their mode of action is their
Particular forages may provide added benefits. Lourenço suppression of ruminal protozoa (Makkar et al., 1998;
et al. (2008b) reported botanically diverse pastures to affect Terefedegne, 2000). Decreased methane production has
FA metabolism in the rumen, with higher rumen outflow of been observed (Lila et al., 2003; Hess et al., 2004; Pen et al.,
cis-9,trans-11 CLA and VA. It was suggested that changes in 2007), although not always (Sliwinski et al., 2002).
the rumen FA metabolism were associated with the presence Regarding their effects on biohydrogenation, N. McKain
of plant secondary metabolites in herbs of botanically (unpublished results) found that a range of saponins inhib-
diverse pastures. Red clover silages result in a higher transfer ited LA metabolism in mixed digesta in vitro. However,
efficiency of PUFA to the milk and meat of ruminants com- Lourenço et al. (2008a) reported no differences in the
pared with ryegrass silages (Lee et al., 2003). This has been apparent ruminal biohydrogenation of forage LNA when
related to protein-bound phenols formed by polyphenol quillaja saponins were added to continuous culture fermen-
oxidase (PPO) that may inhibit the activity of plant lipases ters. The different results might be explained by different
(Lee et al., 2004; Lourenço et al., 2005; Van Ranst et al., types of saponins used (Lourenço et al., 2008a). Wallace
2009). More recently, Lee et al. (2007) showed lower ruminal et al. (1994) observed that a saponin-rich Yucca schidigera
biohydrogenation of LNA to be associated with red clover extract inhibited the growth of B. fibrisolvens more than
clonal lines with high-PPO activity, although the latter was other bacteria, indicating that saponins might be useful in
not confirmed by Van Ranst et al. (2009). Chrysanthemum controlling FA biohydrogenation. Whether saponins would
coronarium is another plant that has been described to be useful in vivo is unclear. Fewer protozoa flowing from the
enhance the CLA composition of milk (Cabiddu et al., 2006). rumen might tend to cancel out any benefit from inhibited
In vitro experiments confirmed that C. coronarium inhibited bacterial biohydrogenation. Furthermore, the microbial
ruminal biohydrogenation, resulting in increased accumula- community is known to adapt fairly quickly to metabolise
tion of VA and decreased stearic acid when LA was incu- and detoxify saponins (Makkar and Becker, 1997; Teferedegne
bated with ruminal digesta (McKain et al., 2008). et al., 1999).
Essential oils are steam-volatile or organic solvent extracts Tannins are polyphenolic substances that bind to protein
of plants, usually herbs (Crozier et al., 2006). The effects of and that can therefore affect most digestive activities,
essential oils on ruminal fermentation have been variable, including biohydrogenation. Tannins may also provide other
depending on the nature of the compounds (Calsamiglia benefits, including increased milk production and growth
et al., 2007; Benchaar et al., 2008), their dose (Castillejos rate (Bhatta et al., 2000), higher propionate proportions and
et al., 2006 and 2008; Macheboeuf et al., 2008), the basal lower protozoal numbers in the presence of tannins (Makkar
diet (Castillejos et al., 2005; Benchaar et al., 2008), ruminal et al., 1995a and 1995b) and lower methane emissions
pH (Spanghero et al., 2008) and adaptation time of ruminal (Roth et al., 2001; Woodward et al., 2001 and 2002; Puchala
microorganisms to the presence of essential oils (Castillejos et al., 2005). The inhibitory effect of tannins on ruminal
et al., 2007; Benchaar et al., 2008). Essential oils benefit biohydrogenation has only recently been shown (Vasta et al.,
ruminal N metabolism by inhibiting selectively bacteria that 2009), where the hydrogenation of VA to 18:0 seems to be
ferment amino acids (McIntosh et al., 2003). As to the effect more affected than the conversion of LA to CLA. The main
of these compounds on ruminal biohydrogenation, Benchaar problem to overcome with tannins is their effects on feed
et al. (2006 and 2007) found no effects of essential oils on intake, driven partly by their astringent taste and partly by
milk FA profiles when supplementing dairy cows, whereas their suppression of fibre digestion (Makkar, 2003).
Lourenço et al. (2009), found essential oils rich in the
monoterpenes limonene and carvone to result in the ruminal Lipase as a regulator of biohydrogenation
accumulation of cis-9,trans-11 CLA, suggesting some effects Intuitively, it would be thought that inhibiting lipase activity
of the latter on the extent of ruminal biohydrogenation would be a strategy to deliver more PUFA from ruminal fer-
in vitro. In addition, Durmic et al. (2008) reported some plant mentation, and studies have being carried out with that in
extracts and essential oils to inhibit the growth and/or mind (Van Nevel and Demeyer, 1996; Krueger et al., 2009).
activity of important ruminal biohydrogenating bacteria such Low pH inhibits lipase more than biohydrogenation (Van
as B. fibrisolvens and B. proteoclasticus. These studies sug- Nevel and Demeyer, 1996), which may be a useful tool if
gest some essential oils to have the potential to manipulate decreased fibre degradation at low pH (Stewart, 1977) can
1016
be avoided. However, long-chain PUFA are inhibitory to the number of processes as well as biohydrogenation Inter-
biohydrogenation process itself (Wa˛sowska et al., 2006; species H2 transfer is vital to maintain ruminal fermentation
Fievez et al., 2007), and indeed the bacteria that carry out of dietary nutrients because an accumulation of H2 can
the last step in the conversion of monoenoic FA to fully inhibit the activity of cellulolytic bacteria (Wolin et al., 1997).
SFA are extraordinarily sensitive to the toxic effects of PUFA Indeed, H2 transfer between species fundamentally alters
(Wallace et al., 2006; Maia et al., 2007). Thus, if the con- the metabolic routes employed by these species (Wolin et al.,
centrations of PUFA can be increased, possibly by increasing 1997). Thus, changes that affect one category of H2-producing
lipase activity, VA metabolism may be inhibited, leading to or H2-utilizing microorganisms will inevitably lead to changes
more VA and CLA flowing from the rumen and an end result in the others.
of higher CLA concentrations in meat and milk. Methane and propionate are the two largest sinks for
ruminal H2 (Wolin et al., 1997). When methanogenesis falls,
Defaunation propionate proportions increase, as bacteria such as Sele-
Defaunation as a means of improving ruminal productivity nomonas ruminantium alter their metabolism to dispose
has been a controversial issue for many years. Most aspects of H2 (Latham and Wolin, 1977). Biohydrogenation was
of the impact of defaunation on ruminal productivity have originally proposed to be an alternative H2 sink to metha-
been investigated (Williams and Coleman, 1992). On the nogenesis or propionigenesis (Lennarz, 1966). It is now
basis of the aforementioned sequestration of UFA and high believed that the significance of biohydrogenation to the
CLA and VA concentrations in ciliates, one might predict overall H2 sink is small, in that only 1% to 2% of H2 produced
that defaunation would result in increased biohydrogenation during fermentation is consumed by biohydrogenation
and SFA in milk and meat. Early studies of plasma FA (Nagaraja et al., 1997). Amino acid deamination is another
concentrations in defaunated sheep indicated that the con- aspect of ruminal metabolism that depends on H2 metabo-
centration of SFA in the blood remained the same as in lism, because the disposal of reducing equivalents and the
conventional sheep, whereas the concentration of 18:2 and NADH/NAD ratio are important effectors of branched-chain
18:3 tended to increase when the protozoa were removed amino acid fermentation (Hino and Russell, 1985). Thus,
(Abaza et al., 1975; Klopfenstein et al., 1966). More recently, methanogenesis, propionigenesis, amino acid metabolism
Yáñez-Ruiz et al. (2007) found the predicted higher ratio of and biohydrogenation are all linked metabolically, although
SFA/PUFA in muscle of defaunated lambs. Also of possible the consequences of changing one of these activities on the
relevance to milk fat depression was an increased content of others are often not investigated.
trans-10,cis-12-CLA in the defaunates. The microbial species involved in biohydrogenation carry
out other functions as well. Removing microorganisms
Do we want less biohydrogenation, or more? involved in biohydrogenation might therefore be expected to
The theme of this review is generally that biohydrogenation affect other reactions, and vice-versa. It is often difficult to
should be inhibited, at least partly, to enhance the healthful- disentangle metabolic effects from effects on microbial
ness of ruminant products. It is important to note that this is ecology. For example, agents that inhibit other H2-utilizating
not a universal view. B. proteoclasticus has been viewed by processes may inhibit biohydrogenation as well, rather than
most workers in the field as a target for removal from the the increased H2 availability stimulating biohydrogenation.
rumen microbial population to enhance trans-11-18:1 and The effects of ionophores provide an illustration. In the
cis-9,trans-11-18:2 concentrations in ruminant meat and milk. mixed ruminal community, monensin inhibits methanogen-
However, paradoxically, it may be desirable to increase its esis and amino acid deamination, and propionigenesis is
presence in the rumen to lower the concentration of trans FA increased as would be predicted (Bergen and Bates, 1984),
in these foods, the consumption of which is a known risk factor but it also inhibits biohydrogenation (Fellner et al., 1997).
for cardiovascular disease (Mensink et al., 2003; Mozaffarian As a consequence, ruminants receiving dietary monensin
et al., 2006). Increasing B. proteoclasticus numbers could also have higher concentrations of CLA in adipose tissue and
help to avoid milk fat depression in dairy cows. milk (Marmer et al., 1985; Sauer et al., 1998; Da Silva et al.,
2007). The methanogenesis effect is a metabolic one:
methanogenic archaea are insensitive to monensin, while
H2-producing Gram-positive bacteria are inhibited (Chen and
Interdependence of FA biohydrogenation with other
Wolin, 1979). Monensin also inhibits the growth of Butyri-
areas of ruminal metabolism
vibrio spp. (Chen and Wolin, 1979), which is probably more
In general, most metabolic processes in the rumen are important to biohydrogenation than restricted H2 availability.
affected by others, because of common metabolic pathways Similarly, the effect of monensin on amino acid metabolism
or common microbial species. Biohydrogenation is particu- is due more to the suppression of ‘hyper-ammonia-produ-
larly dependent on other factors, as is its manipulation. cing’ bacteria than a metabolic effect (Russell et al., 1991).
Figure 1 illustrates the interactions between different areas Ciliate protozoa are significant producers of H2 and they host
of ruminal metabolism. The common factor of H2 metabolism specific methanogens both in the cytoplasm and on the cell
is an important feature of these interdependencies. H2 is surface (Williams and Coleman, 1997). The loss of these
produced by the fermentation of sugars, and used in a activities by defaunation, as described above, would be
1017
expected to impact biohydrogenation as well. This mixture of remain where candidate bacterial species have yet to be
direct and indirect influences means that the effect of mea- cultivated. Identification of the bacteria causing milk fat
sures that inhibit biohydrogenation will have on the overall depression remains only provisional. Major scope exists for
fermentation are typically difficult to predict. dietary manipulation of biohydrogenation. This has been
Some FA supplements may be doubly beneficial to rumi- achieved successfully with plant and marine oils, yet some
nant nutrition, lowering the environmental damage of problems remain, particularly where other fermentative
methane emissions and improving the supply of UFA in activities are affected. Nevertheless, there are strong links
ruminant products. There is substantial overlap between the between inhibited biohydrogenation and other benefits,
inhibitory effects of long chain length PUFA and medium including decreased methane emissions. Thus, solutions to
chain SFA (10:0 to 14:0) on biohydrogenation (Jenkins et al., improving the FA profile of milk and meat may also benefit
2008; Goel et al., 2009), methanogenesis (Martin et al., the environment. Botanicals and phytochemicals seem to be
2009) and protozoal growth (Matsumoto et al., 1991; promising candidates help to achieve these objectives.
Boeckaert et al., 2007b; Sato and Karitani, 2009). Henderson
(1973) discovered, in a pure culture study, that both a Acknowledgements
Butyrivibrio spp. and a ruminal methanogen were inhibited
The Rowett Institute of Nutrition and Health receives funding
by 10:0 to 14:0, while other Gram-negative species were
from the Rural and Environment Research and Analysis Direc-
not. Comparisons in which effects of FA on both biohydro-
torate of the Scottish Government. The stay of M. Lourenço at
genation and methanogenesis are measured, such as that
the Rowett Institute of Nutrition and Health was supported by
carried out by Goel et al. (2009), are rare, however. Goel
the Fund for Scientific Research – Flanders (Belgium). Post-
et al. (2009) found that capric acid (10:0) inhibited both
doctoral grant of M. Lourenço from the Special Research Fund,
processes. The concentration dependence of how capric acid
Ghent University, Belgium. E. Ramos Morales has a TALENTIA
inhibited biohydrogenation differed from that of methano-
scholarship from the Regional Ministry for Innovation, Science
genesis, indicating that there were separate sites of action of
and Enterprise, Andalusia, Spain.
capric acid, one being methanogenic archaea, the other
biohydrogenating bacteria. This is not always the case,
however. For example, myristic acid (14:0) inhibited metha- References
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The role of microbes in rumen lipolysis and biohydrogenation and their

Article in animal · July 2010

Marta Lourenço Eva Ramos-Morales

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The role of microbes in rumen lipolysis and biohydrogenation

Keywords: biohydrogenation, cellulose digestion, lipase, microbial protein synthesis, rumen

Implications alter their activity. In this way, the healthiness of ruminant

SFA Acetate Propionate

8,10-CLA 10,12-CLA c9,t11 other other t11c15 c9t13 11,13-CLA

t10-18:1 c12-18:1 t11-18:1 t4-t9-18:1 t15+c15-18:1 t13+14-18:1

mechanism by which butyrate is produced. Intracellular acyl- Anaerobic ruminal fungi

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