World J Microbiol Biotechnol (2008) 24:27472755 DOI 10.

1007/s11274-008-9819-y

REVIEW

Mitigation of ruminant methane production: current strategies, constraints and future options
Muhammad Farooq Iqbal Yan-Fen Cheng Wei-Yun Zhu Basit Zeshan

Received: 24 March 2008 / Accepted: 9 July 2008 / Published online: 31 July 2008 Springer Science+Business Media B.V. 2008

Abstract Mitigating methane losses from cattle has economic as well as environmental benets. The aim of this paper is to review the current approaches in relation to associated advantages and disadvantages and future options to reduce enteric methane emission from cattle. Current technologies can be broadly grouped into those that increase productivity of the animal (improved nutrition strategies) so that less methane is produced per unit of meat or milk, and those that directly modify the rumen fermentation so that less methane is produced in total. Data suggest that many of these practices are not appropriate for long term mitigation of methane emissions in ruminants because of their constraints. So it is necessity to develop long term strategies in suppressing methane production. An integrated research investigating animal, plant, microbe and nutrient level strategies would offer a long term solution of methane production. Genetic selection of animals, vaccination, probiotics, prebiotics and plant improvement are the most promising options of all the future approaches discussed. These approaches will reduce enteric methane production without any hazard to animal or environment. Keywords Mitigation Ruminant methane Constraints Future options Introduction Global mean surface temperatures have increased between 1.0 and 1.7 F since 1850 (EPA 2008). Scientists expect that
M. F. Iqbal Y.-F. Cheng W.-Y. Zhu (&) B. Zeshan Laboratory of Gastrointestinal Microbiology, College of Animal Science and Technology, Nanjing Agricultural University, Nanjing 210095, Peoples Republic of China e-mail: zhuweiyunnjau@hotmail.com

global average temperatures could increase by as much as 5 C (9 F) by the middle of the 21st century. As a contributor to global warming, methane (CH4) is second only to carbon dioxide, and accounts for 16% of all greenhouse gas (GHG) emissions globally (Scheehle and Kruger 2006). The ability of methane to retain heat (global warming potential) is 21 times more than carbon dioxide (EPA 2008). Analyses of air trapped in polar ice show that over at least the past 450,000 years and four glacial cycles, the methane mixing ratio, while correlating strongly with temperature, has not exceeded *700 ppb (Delmotte et al. 2004). Etheridge et al. (1998) have documented a relatively stable atmospheric methane of 693 10 (1 SD) ppb over 10101700 AD, followed by a steadily rising mixing ratio reaching 1,750 ppb in 2000 (Fig. 1). There is no doubt that this 2.5-fold increase in atmospheric methane over three centuries is caused by human activities, with agriculture, most notably livestock and rice farming, prominent among them. Methane production from ruminants has been identied as the single largest source of anthropogenic CH4 (Mathison et al. 1998). Livestock emit methane as part of their natural digestive processes. The rumen is the home to billions of microbes, including bacteria, methanogens, protozoa and fungi. These microbes breakdown feed to produce volatile fatty acids (VFAs), carbon dioxide, ammonia and methane. The VFAs are used by animal as energy source whereas gases are removed by eructation. According to FAO, the world livestock population in 2004 comprised of 1,365 million cattle 172 million buffaloes 1,059.8 million sheep, 790 million goats and 943.8 million pigs. The estimated CH4 emission from enteric fermentation is 1730% of global production. According to Johnson and Johnson (1995), cattle can produce 250500 l of methane per day per animal. Cattle

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Fig. 1 A reconstruction of the global-mean methane mixing ratio in the troposphere (lower atmosphere) over three centuries based on measurements in air trapped in Antarctic ice and rn (Etheridge et al. 1998) and on contemporary measurements based on the global

NOAA/CMDL monitoring network (Dlugokencky et al. 1998, 2003). The atmospheric record for the centuries preceding 1700 AD is relatively featureless (Etheridge et al. 1998). 1 ppb = 1 nmol/mol dry air, measured against 1983 NOAA/CMDL gas standards

typically lose 215% of their ingested energy as eructated methane (Giger-Reverdin and Sauvant 2000). Thus, mitigating methane losses from cattle has two important benets. Firstly, less methane means a lower concentration of greenhouse gases (GHGs) in the atmosphere. Secondly, less methane means increased efciency of livestock production and increased income for farmers.

Current mitigation strategies Improved nutrition strategies Concentrates The proportion of concentrate within the diet has been reported to be negatively correlated with methane emissions (Holter and Young 1992; Yan et al. 2000; Fig. 2). Studies have been conducted to determine the effect of starch-based and ber-based concentrates on enteric CH4

production. Lovett et al. (2005) demonstrated that increased ber-based concentrate use at pasture reduced enteric methane per kilogram of animal product (19.26 and 16.02 g of CH4/kg of fat-corrected milk). A positive response to high levels of starch-based concentrate (grains) on methane reduction has also been reported by others (Beauchemin and McGinn 2005). An increased proportion of starch in the diet changes ruminal volatile fatty acid (VFAs) concentrations in such a way that less acetate and more propionate is formed, and the supply of hydrogen for methanogenesis is limited. Also, the pH decreases as the proportion of propionate increases, which reduces methanogenic activity (Walichnowski and Lawrence 1982). Further, concentrate feeding has shown to reduce methane output by reducing the protozoal population (Van Soest 1982). However, increased levels of concentrates may result in health problems e.g. acidosis. High cost of production in the case of concentrate feeding limits their use in low cost systems. Forage type and quality Benchaar et al. (2001) used a modelling approach to assess the effectiveness of different existing nutritional strategies to reduce methane production from ruminants. Simulated strategies included: dry matter intake (DMI), forage-toconcentrate ratio, nature of concentrate (brous versus starchy concentrate), type of starch (slowly versus rapidly degraded), forage species (legume versus grass), forage maturity, forage preservation method (dried versus ensiled), forage processing, and upgrading and supplementation of poor quality forages (straw). Methane production was reduced (-7% and -40%) by increasing DMI and the proportion of concentrate in the diet. The use

Fig. 2 The effects of increasing the proportion of concentrates in the diet of methane output when total digestible energy intake, dry matter intake and feeding level are kept constant (adopted from Yan et al. 2000)

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of more digestible forage (less mature and processed forage) resulted in a reduction of methane production (-15% and -21%). Methane production was lower with legume than with grass forage (-28%), Legumes generally have higher dry matter intakes and produce more milk solids. This reduces methane emissions per unit of milk or meat production. Hegarty (1999b) found that methane production per gram of live weight gain was reduced signicantly when animals were shifted from low digestible pasture to high digestible pasture. Methane emissions per unit of feed intake can be reduced by use of grass cultivars selected for improved animal performance. The PGgRC (2005) annual report indicated that cultivars showed signicant differences at 12 h in % accumulated methane in vitro. However, the practices for producing high quality forages have not been tested under eld conditions. To implement mitigation procedures under eld conditions, it is very important that there be condence in the accuracy of the methane measurement technology. Although many methods have been employed to measure or calculate methane emissions from ruminants (e.g Flux gradient techniques, Boundary layer techniques), the method of choice for grazing animals is the sulphur hexauoride (SF6) tracer technique developed by Johnson et al. (1994a). The other methods have too high an error to be able to detect the small differences that are important in inventory, regulatory or animal science research. Further work is also needed on the SF6 technique: rigorous evaluation against the respiration calorimeter; conrmation of the proportion of methane that is excreted in the atus; and evaluation of the high variability of the technique. Furthermore, pasture improvement will be a valuable option only if accompanied with reduced animal numbers; otherwise, due to the rise in feed intake associated with improved digestibility of pasture, daily (or annual) methane emission by the individual will increase although methane cost of production (methane/kg of beef or milk) will decrease. Rumen modication strategies Defaunation The elimination of protozoa from the rumen is termed defaunation. Ciliate protozoa are not present at birth but normally start establishing in the gastrointestinal tract 3 weeks after birth (Fonty et al. 1988). If newborns are reared in isolation from other ruminants, no protozoa will establish (Ivan et al. 1986). Defaunation techniques used in research have been reviewed by Hegarty (1999a). These defaunation approaches include dietary manipulation (virginiamycin, milk fat); synthetic chemicals including copper sulfate, calcium peroxide, dioctylsodium sulfosuccinate and detergents; and natural compounds like vitamin

A, non-protein amino acids and ecdysones, the steroidal hormones which cause skin shedding in insects. Defaunation results in reduced methane production due to (1) reduced ber digestion (Machmuller et al. 2003), (2) reduced methanogen population associated with protozoa (Machmuller et al. 2000), (3) reduced hydrogen transfer (Finlay and Fenchel 1993), and (4) increased partial pressure of oxygen on the rumen (Eckard 2001). Dohme et al. (1999) showed that methane production decreased by 61% in defaunated rumen uid. Newbold et al. (1995a, b) calculated that 925% of methanogenesis is due to methanogens living in a symbiotic relationship with protozoa. Hegarty (1999a) concluded from reviewed data that this symbiotic association results in 40% of methanogenesis in rumen uid. The results of in vivo trials are quite variable and the response may be subject to strong dietary effect. For example, in some studies, defaunation of cattle fed a high concentrate diet decreased methane production by approx. 50% (Kreuzer et al. 1986). On the other hand, Machmuller et al. (2003) found no reduction in methane production in sheep fed diets based on maize silage, hay and concentrate. Some problems are associated with defaunation. Digestion will be negatively affected if animals are def aunated completely (Machmuller et al. 2003). Some reports indicate that the reduction of methane production by defaunation is only temporary (Ranilla et al. 2004). This also limits the use of the above discussed option. Ionophores Ionophores are polyether antibiotics produced by soil microorganisms that modulate the movement of cations such as sodium, potassium and calcium across cell membranes. Monensin and lasolasid are two ionophores which have been extensively used to manipulate ruminal fermentation. Ionophores affect methane production in four ways: rst, they increase feed conversion efciency (Goodrich et al. 1984); second, they selectively reduce acetate production (Slyter 1979); third, they inhibit the release of H2 from formate (Van Nevel and Demeyer 1979); fourth, they depress ciliate protozoa population (Guan et al. no date). In relation to feed conversion efciency, dry matter intake is reduced and feed conversion efciency is increased as reported by Raun (1990). In a particular trial OKelly and Spiers (1992) attributed 55% reduction in methane to anorectic effect (due to reduced feed intake) and 45% to the specic rumen activity effect. In vitro study by Fuller and Johnson (1981) indicated that monensin and lasalocid additions to the high grain substrate reduced CH4 loss to as little as 53% of that in the control Callaway et al. (1997) claimed that maximum methane inhibition by

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monensin was 18% in vitro. In one study monensin added at 24 ppm in the diet of dairy cows decreased CH4 by 28% (Kinsman et al. 1997). McGinn et al. (2004) reported that loss of gross energy (GE) to methane decreased 9% using monensin. Van Nevel and Demeyer (1996) reviewed data from in vitro trials and found that percentage inhibition showed a wide range (076%). However, the methane-suppressing effect of monensin is not maintained for repeated applications (Omer 2004). Cattle studies have shown that ionophore-induced suppression of enteric CH4 production is short lived (Johnson et al. 1994b; McCaughey et al. 1997). Omer (2004) found that rumen microbes adapted ionophores within 45 days. Secondly, the use of ionophores in grazing ruminant has not been assessed. Furthermore, there is increasingly consumer resistance to the routine use of ionophores as they are a type of antibiotics. From 1 January 2006, monensin, the only EU-approved antibiotic growth promoter for use in beef cattle, had been prohibited in EU. Oils

et al. (2000) found a high efcacy of coconut oil against methane release in the intensive type diet (lower structural carbohydrate content) as compared to extensive diet (high structural carbohydrate content). The efciency of different oils is also different. Machmuller et al. (1998) identied coconut oil as more effective inhibitor than rape seed, sunower seed, and linseed oil. McGinn et al. (2004) reported a 22% decrease in methane emissions by addition of sunower oil to the diet of cattle. Chiquette and Benchaar (2005) showed that essential oils reduced methane production by up to 46%. However, fats and oils can have many negative effects on the animals. McGinn et al. (2004) reported that oil addition reduced ber digestibility signicantly. Jordan et al. (2006) calculated that nishing animals on the copra meal containing coconut oil would require a longer time to reach a common carcass weight and would lessen the effects on total CH4 emissions. High cost and the negative effect on milk fat concentration (Zheng et al. 2005) are other constraints of oil addition. Dicarboxylic acids

Addition of oils to ruminant diets may decrease methane emission by up to 80% in vitro (Fievez et al. 2003) and about 25% in vivo (Machmuller et al. 2000). Addition of unsaponied fats or oils to the diets of ruminants suppresses methane production by two mechanisms. The indirect mechanism includes protozoal inhibition, reduction of double bonds in unsaturated fatty acids, increased productivity and enhanced propionate production. Fatty acid toxicity to methanogens is the direct mechanism. Indirect effects of oils, which ultimately result in methane reduction, have been reported by many workers. Zheng et al. (2005) observed increased milk production using different vegetable oils. Czerkawski et al. (1966) reported the suppressive effect of long chain unsaturated fatty acids on methanogenesis and demonstrated the competition between methanogenesis and biohydrogenation process. McGinn et al. (2004) observed a lower acetate concentrations, higher propionate concentrations and lower acetate-topropionate ratio when sunower oil was added to the diets of cattle. The detrimental effects of certain oils on rumen protozoa were reported by Machmuller et al. (1998) and as a result methane production was reduced. When canola oil was added in the diets of sheep at 0%, 3.5% or 7%, the number of rumen protozoa was reduced by 8897% (Machmuller et al. 1998). Direct toxic effects of unsaturated fatty acids have also been demonstrated (Henderson 1973). Machmuller et al. (2003) suggested that coconut oil directly inhibited rumen methanogens, probably by changing their metabolic activity and composition. The effects of different oils on ruminal methane fer mentation are related to diet composition. Machmuller

Dicarboxylic organic acids (malate, fumarate) are potential precursors of propionate which stimulate H2 utilization for reduction of fumarate to succinate during propionate synthesis at the expense of enteric methane. In vitro studies conducted by Mohammed et al. (2004) demonstrated that addition of fumarate increased propionate production and reduced methane output. Malate has a similar effect (Jalc and Ceresnakova 2002). In vivo studies also demonstrated the potential effects of dicarboxylic acids on methane output (Bayaru et al. 2001). In contrasts, some scientists reported that fumaric acid was not effective in reducing CH4 losses in vivo (McGinn et al. 2004). The following constraints are associated with the use of dicarboxylic acids. First, they are expensive chemicals. Second, they are not suitable for grazing animals as they have to be fed daily. Third, their efciency is reduced when concentrates are fed, as evident from in vitro trial in which the efciency was only 4.8% (Carro and Ranilla 2003). Inhibitors and analogues Targeted inhibitors have strong negative effect on methanogenesis. Chloroform, a chlorinated methane analogue, inhibits methanogenesis through inhibition of methyl-CoM reductase (Gunsalus and Wolfe 1978) but is obviously not suitable for eld application. Mathers and Miller (1982) found signicant reduction in methane production by chloral hydrate, but prolonged feeding led to liver damage (Lanigan et al. 1978). Bromochloromethane appeared to be a potent methane inhibitors (Van Nevel and Demeyer

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2751 Table 1 Estimates of methane emissions (kg/year and kg/kg milk in parentheses) from dairy cows as effected by annual milk yield and body weight (Kirchgessner et al. 1995) Body weight (kg) Milk yield (kg/year)a 4,000 500 600 700
a

1996; Mathison et al. 1998) but its effect on ruminal methanogenesis was transient (Sawyer et al. 1974; Van Nevel and Demeyer 1996). However, the combination of bromochloromethane (BCM) and a-cyclodextrin reduced methane emissions for 28 days in cattle, indicating the potency of compound. 2-Bromoethanesulfonate (BES), a bromine analogue of coenzyme M involved in methyl group transfer during methanogenesis, is a potentially strong inhibitor of methane production but unfortunately its effects only lasted 3 days in sheep (Dong et al. 1997). Tomkins and Hunter (2003) reported that loss of GE to methane decreased from 3.9% to 0.6% in steers using a compound containing BCM. Microbial adaptation, decreased ber digestion and toxicity to the animal are drawbacks of these inhibitors. Recent studies have shown that p-aminobenzoic acid (pABA) derivatives bearing n-propyl, isopropyl and isobutyryl nitrogen substituents strongly inhibit RFA-P synthase (Dumitru et al. 2003). RFA-P synthase catalyzes the rst step in the synthesis of tetrahydromethanopterin, a key cofactor required for methane formation. Results with an articial rumen indicated that none of these RFA-P inhibitors affect bacterial metabolism adversely (Dumitru et al. 2003). However, much more work needs to be done before their commercial use.

5,000 100 (0.02) 108 (0.0216) 116 (0.0232)

6,000 105 (0.0175) 113 (0.0183) 121 (0.0202)

95 (0.0238) 103 (0.0258) 111 (0.0278)

310 days of lactation combined with a 55 day dry period

Large variations exist (40) in methane emissions between animals at the same level of performance and diet. In trials with Friesian Jersey crossbred herds, signicant variation was found between cows for this phenotype (PGgRC 2004). Goopy and Hegarty (2004) identied some steers as high and low emitters on identical feed and feed intakes. Work has not been done to determine whether these differences are related to intake behavior, or to potential anatomical and physiological differences in the gastrointestinal tract of cattle or the heritability of this trait. The assumed factors responsible for such differences are the rate of passage, microbial activity, fermentation conditions and grazing behavior. The degree of variability suggests that breeding animals with low methane emissions but uncompromised performance would be a helpful way for methane mitigation in future. Microbe level

Future options Animal level Probiotics Genetic selection of animals Animal production efciency can be increased by selection of animals with improved genetic merit (the ratio of an animals performance with the group average). As a result of increased productivity, methane production per unit of milk or meat will decrease. Kirchgessner et al. (1995) suggested that increasing milk production of dairy cows from 4,000 to 5,000 kg/cow/year increases annual methane emissions but will decrease emissions per kg of milk by 0.16 for a 600 kg cow. A further increase to 6,000 kg/year would decrease emissions per kg of milk by a further 0.128 (Table 1). Genetic variation in feed intake also exists, independent of liveweight and average daily gain and this variation provides a basis for genetic selection for feed-use efciency of cattle (Arthur et al. 2001). Cattle that eat less than their peers of equivalent liveweight and average daily gain have a low residual feed intake (RFI) and are more feedefcient, as shown by lines of cattle divergently selected for RFI (Arthur et al. 1996). So selection for reduced RFI will lead to substantial and lasting methane abatement. Probiotics are microbial feed additives that inuence rumen fermentation directly resulting in improved animal productivity. The most widely used probiotics are yeast and Aspergillus oryzae. Some available products guarantee high numbers of live yeast cells and are sold as live yeast while other products are sold as yeast cultures containing both yeast cells and the media on which they are grown. Although not yet clear, it is assumed that yeast cultures reduce methane production in four ways: (1) by increasing butyrate or propionate production (Lila et al. 2004); (2) by reducing protozoan numbers (Newbold et al. 1998); (3) by promoting acetogenesis (Chaucheyras et al. 1995); and (4) by improving animal productivity (Bruno et al. 2005). In a review of in vitro studies, Wallace and Newbold (1993) found that probiotics improved productivity by 7 8% resulting in reduced methane production per unit of product (milk or meat) in dairy cows and growing cattle. Mwenya et al. (2004) reported that sheep produced significantly less methane when 0.4% yeast culture was included in a basal hay and 30% concentrate diet. On the other hand, McGinn et al. (2004) reported that reduction in methane

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was small and not statistically signicant. It might however be argued that short term incubations are inappropriate to study the effects of yeast on rumen fermentation. The effects of S. cerevisiae are mediated through an effect on the numbers and activity of microbes in the rumen (Newbold 2003) and periods longer that 2448 h may be required to fully realise the effects of the yeast. Rumen-simulating fermentors provide a suitable tool to study the longer term effects of additives on ruminal fermentation and can be used to screen strains of S. cerevisiae for their effects in the rumen. Studies have also shown that methane suppression effects of proboitics are not consistent (Newbold et al. 1995a, b). There is a need to identify the dietary and management situation in which probiotics can give consistent results. Since probiotics are fed daily, they may only be suitable for dairy and intensive beef systems. Immunization One possible future pathway to reduce CH4 output is to immunize animals against their own methanogens and protozoa. Shu et al. (1999) demonstrated the reduction in the numbers of Streptococcus bovis in the presence of antiprotozoal antiserum. Scientists in Australia have claimed to invoke an immune response to rumen protozoa by administering an immunogenic preparation (Baker et al. 1997). This will indirectly affect the activity of rumen methanogens as they have a commensal relationship with rumen protozoa. The development of an antimethanogenic vaccine is also in progress. It is anticipated that vaccination could reduce CH4 output up to 70%. However, Wright et al. (2004) found a non-signicant reduction of 6% in the sheep after 4 weeks of vaccination with three methanogen mixtures and a signicant 7.7% reduction in methane production per kg DM intake after revaccination. Much more work is needed to make this technique effective, as there are multiple strains of Archaea in the rumen. If this proves successful, the vaccination would be a valuable tool for methane reductions as it could be applied to a whole ruminant population. Nutrient level Hexose partitioning Methanogenesis is an important sink for hydrogen. Beever (1993) suggested that hydrogen production and methanogenesis could be altered by increasing the quantity of microbial cells leaving the rumen per unit of carbohydrate consumed. This may be possible through hexose partitioning. Hexose partitioning alters the pattern of methanogenesis by changes in the diet, which manipulate the amount of feed carbohydrate going directly into

microbial growth as opposed to fermentation (Meeks and Bates 1999). Formulating diets with increasing water-soluble carbohydrate (WSC) to cell wall (NDF) ratio will decrease methane production because the partitioning of hexose into fermentation end-products (including methane) and microbial biomass in the rumen (in vivo) is inuenced by the dietary carbohydrate source as observed in vitro (Moss and Newbold 2000). They found that methane production was signicantly and negatively correlated to the water soluble carbohydrate content as a proportion of the neutral-detergent ber content of the diet. In their study, The use of 15N in vitro was shown to be a suitable marker of microbial growth for diets with a high proportion of concentrate. The scientists also found signicant reduction in methane production in vivo (Moss et al. 2001). Their results showed that increasing the proportion of WSC in the ration from 0.089 to 0.234 decreased methane production (l kg-1 rumen degradable OM) by 27.7%. However, further experimental research is required to conrm the inuence of diet on hexose partitioning and to investigate carbohydrate sources that provide improved hexose partitioning. Prebiotics Prebiotics are non-digestible dietary supplements which enhance the benecial intestinal organisms. Galacto-oligosaccharides (GOS), non-digestible carbohydrates in nonruminants, have encouraging results as a prebiotic in cattle nutrition. Mwenya et al. (2004) reported that methane production reduced by 10% in sheep fed basal hay and 30% concentrate diet supplemented with 20 g GOS. In contrast, Santoso et al. (2003) found that 2% supplementation of orchardgrass silage or mixed orchard grass/alfalfa silage diets did not reduce methane output. Nevertheless, further research is needed to conrm their effects and to understand mode of their action. Plant level There are some studies about the effects of plants and plant extracts on methane production both in vitro and in vivo. Puchala et al. (2005) reported that methane production expressed as both, quantity per day or relative to DMI, was lower for Sericea lespendeza than for crabgrass/tall fescue (7.4 vs. 10.6 g/day and 6.9 vs. 16.2 g/kg DMI). White clover results in reduced methane production per unit feed intake by upto 20% (PGgRC 2004). These effects of legumes on methane production are generally attributed to increased dry matter intakes. However, research at Dexel indicated that at least some of the measured decrease in methane production is the result of direct effect of condensed tannins in legumes forages (such as sullah and

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birdsfoot trefoil) on rumen fermentation (Woodward 2003). Hess et al. (2004) reported that in vitro methane production decreased when Calliandra tannins supplemented a tropical grass substrate. Using higher doses of tea saponin (8 mg), Hu et al. (2005) noted a 79% decrease in protozoan counts and 26% decline in in vitro methane release. Alfalfa contains from 2% to 8% malate in its dry matter (Callaway et al. 1997) which deprives methanogens of hydrogen. However, a limited number of studies have investigated the direct and indirect effects of plants on methane production in animals, and it is difcult to provide a comprehensive assessment at this stage about the size of decrease that might be realistically expected in vivo. Teferedegne et al. (1999) claimed that foliage of Sesbania sesban (a saponin-containing subtropical tree) defaunated the rumen of sheep. Under some conditions, severe reductions in methane emissions in sheep and dairy cows grazing kikuya grass (Pennisetum clandestinum) have also been reported (Ulyatt et al. 2002). This indicates the presence of yet unidentied suppressing compounds. Thus, selection and breeding of forages for the above discussed characteristics in combination with investigation of unidentied methane inhibitors is a promising way for the future.

These, if proven successful, will mitigate methane emissions without any hazard to animal or environment. Hexose portioning in the rumen is an interesting concept, but its efciency needs to be veried in vitro as well as in vivo. Improvement and breeding of plants is another helpful way to control of methanogenesis, but the estimation of time required must be realistic.
Acknowledgments The authors would like to thank the Natural Science Foundation of China (Grant No. 30530560) for support.

References
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Conclusion Since methane conversion efciency depends on feed quality and efciency of the rumen, so the current approaches which offer abatement in enteric methane production include improved-nutrition strategies and rumen modication strategies. Many of these strategies e.g. increased grain feeding, forage processing and pelleting, dicarboxylic acid supplementation, are not suitable to low cost production industry because of their high cost. Oils and defaunation have a remarkable ability to reduce methane emissions, but ber digestibility is also reduced signicantly. Addition of ionophores and halogenated methane analogues to ruminant diet offers the opportunity to reduce methane production in feed lot cattle but rumen microbes can adapt to them. The use of other currently available strategies (e.g. ionophores, inhibitors and analogues) is not a permanent solution of ruminant methane due to microbial adaptation. Thus, it is a necessity to focus on new approaches to reduce methane emissions. Integrated research investigating animal, plant, microbe and nutrient level strategies might offer a long term solution of methane production. At the animal level, genetic selection is the area of research with the best chance of nding a solution. However, such research will be expensive and long term. At the microbe level, vaccination and probiotics are the promising approaches for future research.

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