Ruminal Methanogenesis and Its Alternatives G Fonty, B Morvan
Total Page:16
File Type:pdf, Size:1020Kb
Ruminal methanogenesis and its alternatives G Fonty, B Morvan To cite this version: G Fonty, B Morvan. Ruminal methanogenesis and its alternatives. Annales de zootechnie, INRA/EDP Sciences, 1996, 45 (Suppl1), pp.313-138. hal-00889640 HAL Id: hal-00889640 https://hal.archives-ouvertes.fr/hal-00889640 Submitted on 1 Jan 1996 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Ruminal methanogenesis and its alternatives G Fonty, B Morvan INRA, Laboratoire de Microbiologie, C.R. de Clermont-Ferrand-Theix 63122 Saint-Genès-Champanelle, France Methanogenesis is the terminal step in carbon nucleotide sequences of archaea and bacteria flow in many anaerobic habitats including indicate early divergence of the two domains marine and freshwater sediments, marshes during evolution. The phylogenetic data based and swamps, rice paddies, geothermal on rRNA analysis and nucleic acid habitats, anaerobic sewage digestion systems hybridization, coupled with physiological, and animal gastrointestinal tracts. Biological immunological and biochemical characteristics methanogenesis plays a major role in the currently support the taxonomic organization of carbon cycle on earth. Methane escaping from methanogenic archaea into three orders anaerobic habitats can serve as a carbon and comprising six families. Nineteen genera energy source for aerobic methanotrophic containing more than 50 species have been bacteria or can escape to the atmosphere, described to date (Boone et al, 1993). where it is a major participant in atmospheric Methanogenic archaea are the dominant chemical reactions and is an important H2 utilizers in the rumen. However despite the greenhouse gas (Zinder, 1993). economic implication of methanogens for The rumen and the large intestine of agriculture, our knowledge of methanogen humans and monogastric animals, do not species in the rumen is still scarce. Our current completely convert organic matter to CH4 and information is based on a few isolates from a C02. They accumulate high concentrations of few animals. Hydrogenotrophic methanogens volatile fatty acids (c.a. 60 mM acetate, 20 mM ,which reduce C02 to CH4, are responsible for propionate and 10 mM butyrate) which are the major part of rumen methanogenesis. available to the animal host as an energy Methanobrevibacter ruminantium (formerly source. The predominant substrates for Methanobacterium ruminantium) was the first methanogens in these ecosystems are H2 and methanogen isolated from the bovine rumen C02. A major difference between intestinal (Smith and Hungate, 1958). It was present in ecosystems and complete bioconversion high numbers and formed CH4 from H2 and systems is the turnover time. Digestive C02- Strains of methanogenic archaea which ecosystems have turnover times of ap- had the morphological and physiological proximately 1 to 2 days, while in complete characteristics of Methanobrevibacter spp. but conversion systems the turnover times are which did not react with rabbit antisera raised weeks to months (Miller, 1991 Thus, the slow- against M. ruminantium were also isolated from growing fatty acids oxiders and acetogenic the bovine rumen (Lovley et al, 1984; Miller et methanogens cannot be sustained in intestinal al, 1986). Methanobrevibacter is considered to ecosystems. be the major methanogenic genus in the rumen (Miller, 1991). Paynter and Hungate (1968) reported the isolation of Methanomicrobium Rumen methanogens mobile from the rumen of a cow, and Methanosarcina species presumably growing Methanogens are a sub-group of the archaea on methylamines and methanol were found in domain (Woese et al, 1987). Bacteria and low numbers (104 ml- ) in a cow rumen methanogenic archaea differ in the (Patterson and Hespell, 1979). Strains composition of their cellular constituents. The ressembling Methanobrevibacter species have characteristic peptidoglycan polymer of the cell also been isolated from the rumen contents of walls of bacteria is absent from archaea. The young lambs (Morvan et al, unpublished data), lipids of methanogens are glycerol ethers but up to now there is no information on the rather than glycerol esters. Ribosomal RNA characteristics of the methanogens of the adult ovine rumen (Miller, 1994). In the rumen, some fermentation of feed. It would appear that the methanogens are associated with protozoa total methane emissions from ruminants (Stumm et al, 1982). Methanogens appear in increases by about 0.5 to 1 % per year. An the rumen of new-born animals very soon after estimation by Crutzen et al (1986) indicated birth since they were found as early as two or that over the last century, CH4 emission by three days of age in the lamb rumen (Fonty et ruminants has increased 4 fold and has almost al, 1987; Morvan et al, 1994). At the end of the doubled within the past 40 years. first week of life their numbers reached 106 Mi- The effects of environmental factors on 1 rumen contents and when the animals were methanogenesis in the rumen are not well two-weeks old the methanogenic population documented because it is difficult to design was close to that observed in adult sheep. experiments that include the effects of pH, toxic substances, composition of the diet etc. However some factors which influence Biochemistry of ruminal methane- methane emission by ruminants have been genesis investigated. The major factors are the followings : C02 is converted to CH4 by sequentially - differences between animal species: The reduced intermediates. The reactions, contributions of different animal species to enzymes and electron carriers involved in this global CH4 emissions are highly variable. Of process have been the subject of numerous the domestic species, cattle, with their large studies during the past decades, and the size and numbers, are the major contributions, results of these investigations on the producing 70°/ of the animal emissions. biochemistry of methanogenesis have been Smaller ruminant species, sheep and goats, intensively reviewed (Vogels et al, 1988; produce approximately 13% of animal Thauer et al, 1993) and will not be detailed in emissions, horses and mules appear to be the this paper. principal non ruminant methane emitters (2%), followed by pigs (1%). Crutzen et al (1986) estimate that wild ruminants globally produce Factors affecting methane production about 5% of total animal methane. Cattle from in the rumen developed countries produce more methane than the cattle from developing countries Owing to the lack of fatty acid oxidizers and (Crutzen et al, 1986; Seiler, 1984). acetotrophic methanogens, the rumen can be - digestibility of feed intake: Approximatively 4 considered as a truncated ecosystem when to 10 per cent of the gross energy ingested by compared to an anaerobic digestor and dairy and feed cattle is lost in the form of considerably less CH4 is produced per unit methane (Vermorel, 1995). The more carbon digested compared to the anaerobic digestible the feed intake, the greater the digestor. However, because of the large proportion of its gross energy that is released amounts of plant material ingested, the rumen in the form of methane. It must be noted, fermentations lead to the emission, by however, that the ingestion of highly digestible belching, of large volume of methane, feedstuffs results in a high level of animal approximately 200-400 liters per day from the performance (Sauvant, 1993). bovine rumen (Miller, 1991; Tyler, 1991) which - feeding level: Blaxter and Clapperton (1965) correspond to a production of 800 to 1600 liters demonstrated that the methane emission of hydrogen. A sheep eructates approximately expressed as the percentage of gross energy 50 liters of CH4 per day. According to Sauvant decreases significantly with an increase in the (1993) there are 65 liters of methane and 152 level of intake, i.e. it is affected by the level of liters of C02 produced in the rumen for each kg production and the rate of passage of the of organic matter fermented. particle in the digestive tract. This means that Ruminants are the second largest for similar diets, ruminants at a high level of anthropogenic methane source in the world production convert a smaller proportion of emitting 80 Tg/year (15% of the total) (Crutzen, ingested feed into methane. 1994). This represents a lost opportunity for - mode of production: According to Sauvant transforming more carbon into useful products, (1993), intensive animal production is not such as meat or milk during the natural associated with increased methanogenesis. In fact, the reverse occurs if methane production cellulose breakdown. The fungal metabolism is evaluated against the units of product shifts towards a greater production of acetate produced. at the expense of lactate and ethanol (Bauchop - nature of the feed: Generally, for equal and Mountfort, 1981; Fonty et al, 1988; Marvin- energy digestibility, low-fibre diets lead to a Sikkema et al, 1990). Fermentation by other lower methane production than high fibre diets. important microbes in the rumen, such as R. Fat supplementation of the feed