Section 20 – Methane Emission for Genetic Evaluation: Difference between revisions

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== Introduction ==
== Introduction ==
Increases in milk production through management and genetics have substantially improved feed efficiency and decreased costs per unit of product over recent decades. However, dairy systems are also associated with environmental costs (Baskaran et al., 2009<ref>Baskaran, R., Cullen, R., and Colombo, S. 2009. Estimating values of environmental impacts of dairy farming in New Zealand, New Zealand J. Agric. Res. 52: 377-389, DOI: 10.1080/00288230909510520.</ref>), with methane (CH<sub>4</sub>) emissions associated with rumen microbial fermentation being both an important contributor to global greenhouse gas (GHG) emissions, as well as an avoidable loss of energy that could otherwise be directed into milk production. The livestock sector is responsible for 14.5% of the global GHG (Gerber et al., 2013<ref>Gerber, P.J., Steinfeld, H., Henderson, B., Mottet, A., Opio, C., Dijkman, J., Falcucci, A.  , and Tempio, G. 2013. Tackling climate change through livestock – A global assessment of emissions and mitigation opportunities. Food and Agriculture Organization of the United Nations (FAO), Rome.</ref>); dairy cattle account for 18.9% of these emissions, mainly in the form of enteric CH<sub>4</sub> emissions (van Middelaar et al., 2014<ref>Van Middelaar, C.E., Dijkstra. J., Berentsen. P.B.M.. and De Boer. I.J.M. 2014. Cost-effectiveness of feeding strategies to reduce greenhouse gas emissions from dairy farming. J. Dairy Sci. 97:2427–2439.</ref>). Methane is a greenhouse gas with a global warming potential 28 times that of CO<sub>2</sub> (Myhre et al., 2013<ref>Myhre, G., Shindell, D., Bréon, F., Collins, W., Fuglestvedt, J., Huang, J., Koch, D., Lamarque, J., Lee, D., Mendoza, B., and Nakajima ,T. 2013. Anthropogenic and Natural Radiative Forcing. T.F. Stocker, D. Qin, G.-K. Plattner, M. Tignor, S.K. Allen, J. Boschung, A. Nauels, Y. Xia, V. Bex, and P.M. Midgley, ed. Cambridge University Press, Cambridge, United Kingdom and New York, NY, USA.</ref>). Methane from ruminant livestock is generated during microbial fermentation in the rumen and hindgut (enteric CH<sub>4</sub>), and from decomposition of manure. Enteric CH<sub>4</sub> contributes 80% of CH<sub>4</sub> emissions by ruminants, and manure decomposition contributes 20%. Enteric CH<sub>4</sub> accounts for 17% of global CH<sub>4</sub> emissions and 3.3% of total global greenhouse gas emissions from human activities (Knapp et al., 2014<ref>Knapp, J.R., Laur, G.L., Vadas, P.A., Weis,s W.P., and Tricarico, J.M. 2014. Invited review: Enteric methane in dairy cattle production: Quantifying the opportunities and impact of reducing emissions. J. Dairy Sci. 97:3231-3261.</ref>). There is, therefore, a significant research interest to find ways to reduce enteric CH<sub>4</sub> emissions by ruminants. Ruminant animals have a digestive system to digest plant materials efficiently. Like most mammals, ruminants lack the cellulase enzyme required to break the beta-glucose linkages in cellulose, but they play host to diverse populations of rumen microbes that can digest cellulose and other plant constituents. When rumen bacteria, protozoa and fungi ferment carbohydrates and proteins of plant materials, they produce volatile fatty acids, principally acetate, propionate and butyrate. High fibre diets favour acetate synthesis. Synthesis of acetate and butyrate are accompanied by release of metabolic hydrogen, which, if allowed to accumulate in rumen fluid, has negative effects on microbial growth, and feed digestibility (Janssen, 2010<ref>Janssen, P.H. 2010. Influence of hydrogen on rumen methane formation and fermentation balances through microbial growth kinetics and fermentation thermodynamics. Anim. Feed Sci. Technol. 160:1–22. doi:10.1016/j.anifeedsci.2010.07.002.</ref>). Rumen Archaea are microorganisms that combine metabolic hydrogen with CO<sub>2</sub> to produce CH<sub>4</sub> and water. Archaea play a vital role, therefore, in protecting the rumen from excess metabolic hydrogen, and the CH<sub>4</sub> they produce is an inevitable product of rumen fermentation. A number of CH<sub>4</sub> phenotypes have been defined (Hellwing et al., 2012<ref>Hellwing, A.L.F., Lund, P., Weisbjerg, M.R., Brask, M., and Hvelplund. T. 2012. Technical note: test of a low-cost and animal-friendly system for measuring methane emissions from dairy cows. J. Dairy Sci. 95:6077–85. doi:10.3168/jds.2012-5505.</ref>); the most widely used is CH<sub>4</sub> production (MeP) in liters or grams per day. The CH<sub>4</sub> production trait is highly correlated with feed intake (Basarab et al., 2013<ref>Basarab, J., Beauchemin, K., Baron, V., Ominski, K., Guan, L., Miller, S., and Crowley, J. 2013. Reducing GHG emissions through genetic improvement for feed efficiency: Effects on economically important traits and enteric methane production. Animal, 7(S2):303-315. doi:10.1017/S1751731113000888</ref>; De Haas et al., 2017<ref name=":0">de Haas, Y., Pszczola, M., Soyeurt, H., Wall, E., and Lassen, J. 2017. Invited review: Phenotypes to genetically reduce greenhouse gas emissions in dairying. J. Dairy Sci. 100:855-870.</ref>) and, thereby, with the ultimate breeding goal trait: milk production in dairy cattle. The economic value of daily dry matter intake and associated methane emissions in dairy cattle showed that increasing the feed performance estimated breeding value by one unit (i.e. 1 kg of more efficiently converted DMI during the cow’s first lactation) translates to a total lifetime saving of 3.23 kg in DMI and 0.055 kg in methane (Richardson et al., 2019<ref>Richardson, C., Baes, C., Amer, P., Quinton, C., Martin, P., Osborne, V., Pryce, J.E., and Miglior, F. 2020. Determining the economic value of daily dry matter intake and associated methane emissions in dairy cattle. Animal 14:171-179. doi:10.1017/S175173111900154X</ref>). Feed Performance was defined as a 1 kg increase in more efficiently used feed in a first parity lactating cow. These results show not only the relation between DMI and CH<sub>4</sub> production, but also the economic relationship between these traits. Persistency of lactation was found to be positively associated with increased feed efficiency and decreased methane production and intensity. Feed efficiency was associated with lower methane intensity. Feed efficiency and methane emissions can be improved by selecting for dairy cattle that are smaller and have increased persistency of lactation. Efficiency and methane emissions can be further improved by improved management of body condition score and by extending lactations beyond the conventional 305-day length (Seymour, 2019<ref>Seymour, D.J. 2019. Feed Efficiency Dynamics in Relation to Lactation and Methane Emissions in Dairy Cattle. PhD thesis, The University of Guelph, Canada.</ref>). According to Ellis et al. (2007)<ref>Ellis, J.L., Kebreab, E., Odongo, N.E., McBride, B.W., Okine, E.K., and France, J. 2007. Prediction of methane production from dairy and beef cattle. J. Dairy Sci. 90:3456–3466.</ref>, DMI predicted MeP with an R<sup>2</sup> of 0.64, and ME intake (MJ/d) predicted MeP with an R<sup>2</sup> of 0.53 for dairy cattle. AlternativePhenotype definitions include CH<sub>4</sub> intensity (MeI), which is defined as liters or grams of CH<sub>4</sub> per kg of milk, and CH<sub>4</sub> yield (MeY), which is defined as liters or grams of CH<sub>4</sub> per kg of dry matter intake (DMI) (Moate et al., 2016<ref>Moate, P.J., Deighton, M.H., Williams, S.R.O., Pryce, J.E., Hayes, B.J., Jacobs, J.L., Eckard, R.J., Hannah, M.C. and Wales, W.J., 2016. Reducing the carbon footprint of Australian milk production by mitigation of enteric methane emissions. Anim. Prod. Sci. 56:1017-1034.</ref>). Residual CH<sub>4</sub> production (RMP) is calculated as observed minus predicted CH<sub>4</sub> production (Herd et al., 2014<ref>Herd, R.M., Arthur, P.F., Bird, S.H., Donoghue, K.A., and Hegarty, R.S. 2014. Genetic variation for methane traits in beef cattle. In: Proc. 10th World Conference on Genetic Applied to Livestock Production (WCGALP), 17-22 August, 2014. Vancouver, Canada.</ref>, Berry et al., 2015<ref>Berry, D.P., Lassen, J., and de Haas, Y. 2015. Residual feed intake and breeding approaches for enteric methane mitigation In: Livestock production and climate change. P.K. Malik, R. Bhatta, J. Takahashi, R.A. Kohn, and C.S. Prasad, ed. CABI, Oxfordshire, UK. . Pages 273-291</ref>), with predicted values based on factors such as milk production, body weight and feed intake. At the moment, it is not obvious which of these phenotypes to use; but, it is important to monitor associations between the chosen CH<sub>4</sub> phenotype and the other important traits in the breeding goal (e.g. production, fertility, longevity) to avoid unfavorable consequences. Berry and Crowley (2012)<ref>Berry, D.P., and Crowley, J.J. 2012. Residual intake and body weight gain: A new measure of efficiency in growing cattle, J. Anim. Sci. 90:109–115, <nowiki>https://doi.org/10.2527/jas.2011-4245</nowiki></ref> describe advantages and limitations of ration traits. For example, because feed efficiency traits are a linear combination of other traits it is not recommended to include them in an overall total merit index, which is a clear limitation. For all applications it is necessary to measure the CH<sub>4</sub> emission of each animal individually. These guidelines are intended to make the right choices for this. Whilst diet changes and feed additives can be effective mitigation strategies for CH<sub>4</sub> emissions (Beauchemin et al., 2009<ref>Beauchemin, K.A., McAllister, T.A., and McGinn, S.M. 2009. Dietary mitigation of enteric methane from cattle. CAB Rev.: Perspect. Agric., Vet. Sci., Nutr. Nat. Res. 4:1–18.</ref>; Martin et al., 2010<ref>Martin C., Morgavi, D.P., and Doreau, M. 2010. Methane mitigation in ruminants: from microbe to the farm scale. Animal 4:351–365.</ref>; Hristov et al., 2013<ref>Hristov, A.N., Oh, J., Firkins, J.L., Dijkstra, J., Kebreab, E., Waghorn, G., Makkar, H.P.S., Adesogan, A.T., Yan,g W., Lee, C., Gerber, P.J., Henderson, B., and Tricarico, J.M. 2013. Special topics - Mitigation of methane and nitrous oxide emissions from animal operations: I. A review of enteric methane mitigation options. J. Anim. Sci. 91:5045–5069.</ref>), their effects depend on the continued use of a particular diet or additive and there have been issues with the rumen microbiomes adapting to additives. Rumen bacterial communities are highly dynamic after a diet switch and did not stabilize within 5 wk of cows grazing pasture (Bainbridge et al., 2016<ref>Bainbridge, M.L., Saldinger, L.K., Barlow, J.W., Alvez, J.P., Roman, J. Kraft, J. 2016. 1609 Rumen bacterial communities continue to shift five weeks after switching diets from conserved forage to pasture. J. Anim. Sci. 94, suppl_5:783, <nowiki>https://doi.org/10.2527/jam2016-1609</nowiki> (abstr.)</ref>). In contrast, breeding for reduced CH<sub>4</sub> emissions should result in a permanent and cumulative reduction of emissions (Wall et al., 2010<ref>Wall, E., Simm, G., and Moran, D. 2010. Developing breeding schemes to assist mitigation of greenhouse gas emissions. Animal 4:366-376.</ref>). Several studies have shown that CH<sub>4</sub> emissions by ruminants have a genetic component, with heritability in the range 0.20 – 0.30 (de Haas et al., 2011<ref>de Haas, Y., Windig, J.J., Calus, M.P.L., Dijkstra, J., de Haan, M., Bannink, A., and Veerkamp, R F. 2011. Genetic parameters for predicted methane production and potential for reducing enteric emissions through genomic selection. J. Dairy Sci. 94:6122–6134.</ref>; Donoghue et al., 2013<ref>Donoghue K.A., Herd, R.M., Bird, S.H., Arthur, P.F., and Hegarty, R F. 2013. Preliminary genetic parameters for methane production in Australian beef cattle. In: Proceedings of the Association for the Advancement of Animal Breeding and Genetics, 20-23 October 2013, Napier, New Zealand, pp. 290–293.</ref>; Pinares-Patiño et al., 2013<ref>Pinares-Patiño C.S., Hickey, S.M., Young, E.A., Dodds, K.G., MacLean, S., Molano, G., Sandoval, E., Kjestrup, H., Harland, R., Pickering, N.K., and McEwan, J.C. 2013. Heritability estimates of methane emissions from sheep. Animal 7: 316–321.</ref>, Kandel et al., 2014A<ref>Kandel, P.B., Vanderick, S., Vanrobays, M.L., Vanlierde, A., Dehareng, F., Froidmont, E., Soyeurt, H., and Gengler, N. 2014A. Consequences of selection for environmental impact traits in dairy cows. Page 19. (<nowiki>http://orbi.ulg.ac.be/bitstream/2268/164402/164401/NSABS162014_poster_Purna_abstract.pdf</nowiki>) I:n Proc. 19th National symposium on applied biological sciences, Gembloux, Belgium.</ref>, B<ref>Kandel, P.B., Vanderick, S., Vanrobays, M.L., Vanlierde, A., Dehareng, F., Froidmont, E., Soyeur,t H., and Gengler, N. 2014B. Consequences of selection for environmental impact traits in dairy cows. In: 10th World Congress on Genetics Applied to Livestock Production (WCGALP), 17-22 August, 2014. Vancouver, Canada.</ref>; Lassen and Lovendahl, 2016<ref>Lassen, J., and Løvendahl, P. 2016. Heritability estimates for enteric methane emissions from Holstein cattle measured using noninvasive methods. J. Dairy Sci. 99:1959-1967.</ref>; López-Paredes et al. 2020<ref>Lopez-Paredes, J., Goiri, I., Atxaerandio, R., García-Rodríguez, A., Ugarte, E., Jiménez-Montero, J.A., Alenda, R and  González-Recio, O. 2020. Mitigation of greenhouse gases in dairy cattle via genetic selection (i): Genetic parameters of direct methane using non-invasive methods and its proxies. J. Dairy Sci. 103.</ref>). Breeding for reduced CH<sub>4</sub> emissions, alone or together with other mitigation strategies, could therefore be effective in reducing the environmental impact of cattle farming and, possibly, also in increasing feed efficiency. Such a breeding scheme would require, as a fundamental starting point, accurate measures of individual CH<sub>4</sub> emissions on a large scale. Several techniques have been developed for the measurement of CH<sub>4</sub> emissions from ruminants, with varying degrees of accuracy (see reviews by Cassandro et al., 2013<ref>Cassandro, M., Mele, M., Stefanon, B.. 2013. Genetic aspects of enteric methane emission in livestock ruminants. Italian J. Anim. Sci. 12:e73: 450-458.</ref> and Hammond et al., 2016A<ref>Hammond, K.J., Crompton, L.A., Bannink, A., Dijkstra, J., Yáñez-Ruiz, D.R., O’Kiely, P., Kebreab, E., Eugenè, M.A., Yu, Z., Shingfield, K.J., Schwarm, A., Hristov, A.N., and Reynolds, C.K. 2016A. Review of current in vivo measurement techniques for quantifying enteric methane emission from ruminants. Anim. Feed Sci. Technol. 219:13–30. doi:10.1016/j.anifeedsci.2016.05.018.</ref>), but routine individual measurements on a large scale (a requisite for genetic selection) have proven to be difficult to obtain and expensive to measure (Pickering et al., 2015<ref>Pickering, N.K., Oddy, V.H., Basarab, J.A., Cammack, K., Hayes, B J., Hegarty, R.S., McEwan, J.C., Miller, S., Pinares, C., and de Haas, Y. 2015. Invited review: Genetic possibilities to reduce enteric methane emissions from ruminants. Animal 9:1431-1440.</ref>; Negussie et al., 2016<ref>Negussie E., Lehtinen, J., Mäntysaari, P., Liinamo, A-E., Mäntysaari, E., and Lidauer, M.. 2016. Non-invasive individual methane measurements in dairy cows using photoacoustic infrared spectroscopy technique. 6th Greenhouse Gases Animal Agriculture Conference (GGAA2016) 14-18 February 2016. Melbourne, Australia. Abstract. p62.</ref>). Therefore, identifying proxies (i.e. indicators or indirect traits) that are correlated to CH<sub>4</sub> emissions, but which are easy and relatively lowcost to record on a large scale, would be a welcome alternative. Proxies might be less accurate but could be measured repeatedly to reduce random noise and in much larger populations. These guidelines are highly indebted to Garnsworthy et al. (2019)<ref name=":1">Garnsworthy, P.C. Difford, G.F. Bell, M.J. Bayat, A.R. Huhtanen, P. Kuhla, B. Lassen, J. Peiren, N. Pszczola, M; Sorg, D. Visker, M.H., and Yan, T. 2019 Comparison of Methods to Measure Methane for Use in Genetic Evaluation of Dairy Cattle. Animals 9:837, 12p.</ref>. In this paper the methods to measure CH<sub>4</sub> are compared with special emphasis to the genetic evaluation of dairy cattle.
Increases in milk production through management and genetics have substantially improved feed efficiency and decreased costs per unit of product over recent decades. However, dairy systems are also associated with environmental costs (Baskaran et al., 2009<ref>Baskaran, R., Cullen, R., and Colombo, S. 2009. Estimating values of environmental impacts of dairy farming in New Zealand, New Zealand J. Agric. Res. 52: 377-389, DOI: 10.1080/00288230909510520.</ref>), with methane (CH<sub>4</sub>) emissions associated with rumen microbial fermentation being both an important contributor to global greenhouse gas (GHG) emissions, as well as an avoidable loss of energy that could otherwise be directed into milk production. The livestock sector is responsible for 14.5% of the global GHG (Gerber et al., 2013<ref>Gerber, P.J., Steinfeld, H., Henderson, B., Mottet, A., Opio, C., Dijkman, J., Falcucci, A.  , and Tempio, G. 2013. Tackling climate change through livestock – A global assessment of emissions and mitigation opportunities. Food and Agriculture Organization of the United Nations (FAO), Rome.</ref>); dairy cattle account for 18.9% of these emissions, mainly in the form of enteric CH<sub>4</sub> emissions (van Middelaar et al., 2014<ref>Van Middelaar, C.E., Dijkstra. J., Berentsen. P.B.M.. and De Boer. I.J.M. 2014. Cost-effectiveness of feeding strategies to reduce greenhouse gas emissions from dairy farming. J. Dairy Sci. 97:2427–2439.</ref>). Methane is a greenhouse gas with a global warming potential 28 times that of CO<sub>2</sub> (Myhre et al., 2013<ref>Myhre, G., Shindell, D., Bréon, F., Collins, W., Fuglestvedt, J., Huang, J., Koch, D., Lamarque, J., Lee, D., Mendoza, B., and Nakajima ,T. 2013. Anthropogenic and Natural Radiative Forcing. T.F. Stocker, D. Qin, G.-K. Plattner, M. Tignor, S.K. Allen, J. Boschung, A. Nauels, Y. Xia, V. Bex, and P.M. Midgley, ed. Cambridge University Press, Cambridge, United Kingdom and New York, NY, USA.</ref>). Methane from ruminant livestock is generated during microbial fermentation in the rumen and hindgut (enteric CH<sub>4</sub>), and from decomposition of manure. Enteric CH<sub>4</sub> contributes 80% of CH<sub>4</sub> emissions by ruminants, and manure decomposition contributes 20%. Enteric CH<sub>4</sub> accounts for 17% of global CH<sub>4</sub> emissions and 3.3% of total global greenhouse gas emissions from human activities (Knapp et al., 2014<ref>Knapp, J.R., Laur, G.L., Vadas, P.A., Weis,s W.P., and Tricarico, J.M. 2014. Invited review: Enteric methane in dairy cattle production: Quantifying the opportunities and impact of reducing emissions. J. Dairy Sci. 97:3231-3261.</ref>). There is, therefore, a significant research interest to find ways to reduce enteric CH<sub>4</sub> emissions by ruminants. Ruminant animals have a digestive system to digest plant materials efficiently. Like most mammals, ruminants lack the cellulase enzyme required to break the beta-glucose linkages in cellulose, but they play host to diverse populations of rumen microbes that can digest cellulose and other plant constituents. When rumen bacteria, protozoa and fungi ferment carbohydrates and proteins of plant materials, they produce volatile fatty acids, principally acetate, propionate and butyrate. High fibre diets favour acetate synthesis. Synthesis of acetate and butyrate are accompanied by release of metabolic hydrogen, which, if allowed to accumulate in rumen fluid, has negative effects on microbial growth, and feed digestibility (Janssen, 2010<ref>Janssen, P.H. 2010. Influence of hydrogen on rumen methane formation and fermentation balances through microbial growth kinetics and fermentation thermodynamics. Anim. Feed Sci. Technol. 160:1–22. doi:10.1016/j.anifeedsci.2010.07.002.</ref>). Rumen Archaea are microorganisms that combine metabolic hydrogen with CO<sub>2</sub> to produce CH<sub>4</sub> and water. Archaea play a vital role, therefore, in protecting the rumen from excess metabolic hydrogen, and the CH<sub>4</sub> they produce is an inevitable product of rumen fermentation. A number of CH<sub>4</sub> phenotypes have been defined (Hellwing et al., 2012<ref>Hellwing, A.L.F., Lund, P., Weisbjerg, M.R., Brask, M., and Hvelplund. T. 2012. Technical note: test of a low-cost and animal-friendly system for measuring methane emissions from dairy cows. J. Dairy Sci. 95:6077–85. doi:10.3168/jds.2012-5505.</ref>); the most widely used is CH<sub>4</sub> production (MeP) in liters or grams per day. The CH<sub>4</sub> production trait is highly correlated with feed intake (Basarab et al., 2013<ref>Basarab, J., Beauchemin, K., Baron, V., Ominski, K., Guan, L., Miller, S., and Crowley, J. 2013. Reducing GHG emissions through genetic improvement for feed efficiency: Effects on economically important traits and enteric methane production. Animal, 7(S2):303-315. doi:10.1017/S1751731113000888</ref>; De Haas et al., 2017<ref name=":0">de Haas, Y., Pszczola, M., Soyeurt, H., Wall, E., and Lassen, J. 2017. Invited review: Phenotypes to genetically reduce greenhouse gas emissions in dairying. J. Dairy Sci. 100:855-870.</ref>) and, thereby, with the ultimate breeding goal trait: milk production in dairy cattle. The economic value of daily dry matter intake and associated methane emissions in dairy cattle showed that increasing the feed performance estimated breeding value by one unit (i.e. 1 kg of more efficiently converted DMI during the cow’s first lactation) translates to a total lifetime saving of 3.23 kg in DMI and 0.055 kg in methane (Richardson et al., 2019<ref>Richardson, C., Baes, C., Amer, P., Quinton, C., Martin, P., Osborne, V., Pryce, J.E., and Miglior, F. 2020. Determining the economic value of daily dry matter intake and associated methane emissions in dairy cattle. Animal 14:171-179. doi:10.1017/S175173111900154X</ref>). Feed Performance was defined as a 1 kg increase in more efficiently used feed in a first parity lactating cow. These results show not only the relation between DMI and CH<sub>4</sub> production, but also the economic relationship between these traits. Persistency of lactation was found to be positively associated with increased feed efficiency and decreased methane production and intensity. Feed efficiency was associated with lower methane intensity. Feed efficiency and methane emissions can be improved by selecting for dairy cattle that are smaller and have increased persistency of lactation. Efficiency and methane emissions can be further improved by improved management of body condition score and by extending lactations beyond the conventional 305-day length (Seymour, 2019<ref>Seymour, D.J. 2019. Feed Efficiency Dynamics in Relation to Lactation and Methane Emissions in Dairy Cattle. PhD thesis, The University of Guelph, Canada.</ref>). According to Ellis et al. (2007)<ref>Ellis, J.L., Kebreab, E., Odongo, N.E., McBride, B.W., Okine, E.K., and France, J. 2007. Prediction of methane production from dairy and beef cattle. J. Dairy Sci. 90:3456–3466.</ref>, DMI predicted MeP with an R<sup>2</sup> of 0.64, and ME intake (MJ/d) predicted MeP with an R<sup>2</sup> of 0.53 for dairy cattle. AlternativePhenotype definitions include CH<sub>4</sub> intensity (MeI), which is defined as liters or grams of CH<sub>4</sub> per kg of milk, and CH<sub>4</sub> yield (MeY), which is defined as liters or grams of CH<sub>4</sub> per kg of dry matter intake (DMI) (Moate et al., 2016<ref>Moate, P.J., Deighton, M.H., Williams, S.R.O., Pryce, J.E., Hayes, B.J., Jacobs, J.L., Eckard, R.J., Hannah, M.C. and Wales, W.J., 2016. Reducing the carbon footprint of Australian milk production by mitigation of enteric methane emissions. Anim. Prod. Sci. 56:1017-1034.</ref>). Residual CH<sub>4</sub> production (RMP) is calculated as observed minus predicted CH<sub>4</sub> production (Herd et al., 2014<ref>Herd, R.M., Arthur, P.F., Bird, S.H., Donoghue, K.A., and Hegarty, R.S. 2014. Genetic variation for methane traits in beef cattle. In: Proc. 10th World Conference on Genetic Applied to Livestock Production (WCGALP), 17-22 August, 2014. Vancouver, Canada.</ref>, Berry et al., 2015<ref>Berry, D.P., Lassen, J., and de Haas, Y. 2015. Residual feed intake and breeding approaches for enteric methane mitigation In: Livestock production and climate change. P.K. Malik, R. Bhatta, J. Takahashi, R.A. Kohn, and C.S. Prasad, ed. CABI, Oxfordshire, UK. . Pages 273-291</ref>), with predicted values based on factors such as milk production, body weight and feed intake. At the moment, it is not obvious which of these phenotypes to use; but, it is important to monitor associations between the chosen CH<sub>4</sub> phenotype and the other important traits in the breeding goal (e.g. production, fertility, longevity) to avoid unfavorable consequences. Berry and Crowley (2012)<ref>Berry, D.P., and Crowley, J.J. 2012. Residual intake and body weight gain: A new measure of efficiency in growing cattle, J. Anim. Sci. 90:109–115, <nowiki>https://doi.org/10.2527/jas.2011-4245</nowiki></ref> describe advantages and limitations of ration traits. For example, because feed efficiency traits are a linear combination of other traits it is not recommended to include them in an overall total merit index, which is a clear limitation. For all applications it is necessary to measure the CH<sub>4</sub> emission of each animal individually. These guidelines are intended to make the right choices for this. Whilst diet changes and feed additives can be effective mitigation strategies for CH<sub>4</sub> emissions (Beauchemin et al., 2009<ref>Beauchemin, K.A., McAllister, T.A., and McGinn, S.M. 2009. Dietary mitigation of enteric methane from cattle. CAB Rev.: Perspect. Agric., Vet. Sci., Nutr. Nat. Res. 4:1–18.</ref>; Martin et al., 2010<ref>Martin C., Morgavi, D.P., and Doreau, M. 2010. Methane mitigation in ruminants: from microbe to the farm scale. Animal 4:351–365.</ref>; Hristov et al., 2013<ref>Hristov, A.N., Oh, J., Firkins, J.L., Dijkstra, J., Kebreab, E., Waghorn, G., Makkar, H.P.S., Adesogan, A.T., Yan,g W., Lee, C., Gerber, P.J., Henderson, B., and Tricarico, J.M. 2013. Special topics - Mitigation of methane and nitrous oxide emissions from animal operations: I. A review of enteric methane mitigation options. J. Anim. Sci. 91:5045–5069.</ref>), their effects depend on the continued use of a particular diet or additive and there have been issues with the rumen microbiomes adapting to additives. Rumen bacterial communities are highly dynamic after a diet switch and did not stabilize within 5 wk of cows grazing pasture (Bainbridge et al., 2016<ref>Bainbridge, M.L., Saldinger, L.K., Barlow, J.W., Alvez, J.P., Roman, J. Kraft, J. 2016. 1609 Rumen bacterial communities continue to shift five weeks after switching diets from conserved forage to pasture. J. Anim. Sci. 94, suppl_5:783, <nowiki>https://doi.org/10.2527/jam2016-1609</nowiki> (abstr.)</ref>). In contrast, breeding for reduced CH<sub>4</sub> emissions should result in a permanent and cumulative reduction of emissions (Wall et al., 2010<ref>Wall, E., Simm, G., and Moran, D. 2010. Developing breeding schemes to assist mitigation of greenhouse gas emissions. Animal 4:366-376.</ref>). Several studies have shown that CH<sub>4</sub> emissions by ruminants have a genetic component, with heritability in the range 0.20 – 0.30 (de Haas et al., 2011<ref>de Haas, Y., Windig, J.J., Calus, M.P.L., Dijkstra, J., de Haan, M., Bannink, A., and Veerkamp, R F. 2011. Genetic parameters for predicted methane production and potential for reducing enteric emissions through genomic selection. J. Dairy Sci. 94:6122–6134.</ref>; Donoghue et al., 2013<ref>Donoghue K.A., Herd, R.M., Bird, S.H., Arthur, P.F., and Hegarty, R F. 2013. Preliminary genetic parameters for methane production in Australian beef cattle. In: Proceedings of the Association for the Advancement of Animal Breeding and Genetics, 20-23 October 2013, Napier, New Zealand, pp. 290–293.</ref>; Pinares-Patiño et al., 2013<ref>Pinares-Patiño C.S., Hickey, S.M., Young, E.A., Dodds, K.G., MacLean, S., Molano, G., Sandoval, E., Kjestrup, H., Harland, R., Pickering, N.K., and McEwan, J.C. 2013. Heritability estimates of methane emissions from sheep. Animal 7: 316–321.</ref>, Kandel et al., 2014A<ref>Kandel, P.B., Vanderick, S., Vanrobays, M.L., Vanlierde, A., Dehareng, F., Froidmont, E., Soyeurt, H., and Gengler, N. 2014A. Consequences of selection for environmental impact traits in dairy cows. Page 19. (<nowiki>http://orbi.ulg.ac.be/bitstream/2268/164402/164401/NSABS162014_poster_Purna_abstract.pdf</nowiki>) I:n Proc. 19th National symposium on applied biological sciences, Gembloux, Belgium.</ref>, B<ref>Kandel, P.B., Vanderick, S., Vanrobays, M.L., Vanlierde, A., Dehareng, F., Froidmont, E., Soyeur,t H., and Gengler, N. 2014B. Consequences of selection for environmental impact traits in dairy cows. In: 10th World Congress on Genetics Applied to Livestock Production (WCGALP), 17-22 August, 2014. Vancouver, Canada.</ref>; Lassen and Lovendahl, 2016<ref>Lassen, J., and Løvendahl, P. 2016. Heritability estimates for enteric methane emissions from Holstein cattle measured using noninvasive methods. J. Dairy Sci. 99:1959-1967.</ref>; López-Paredes et al. 2020<ref>Lopez-Paredes, J., Goiri, I., Atxaerandio, R., García-Rodríguez, A., Ugarte, E., Jiménez-Montero, J.A., Alenda, R and  González-Recio, O. 2020. Mitigation of greenhouse gases in dairy cattle via genetic selection (i): Genetic parameters of direct methane using non-invasive methods and its proxies. J. Dairy Sci. 103.</ref>). Breeding for reduced CH<sub>4</sub> emissions, alone or together with other mitigation strategies, could therefore be effective in reducing the environmental impact of cattle farming and, possibly, also in increasing feed efficiency. Such a breeding scheme would require, as a fundamental starting point, accurate measures of individual CH<sub>4</sub> emissions on a large scale. Several techniques have been developed for the measurement of CH<sub>4</sub> emissions from ruminants, with varying degrees of accuracy (see reviews by Cassandro et al., 2013<ref>Cassandro, M., Mele, M., Stefanon, B.. 2013. Genetic aspects of enteric methane emission in livestock ruminants. Italian J. Anim. Sci. 12:e73: 450-458.</ref> and Hammond et al., 2016A<ref>Hammond, K.J., Crompton, L.A., Bannink, A., Dijkstra, J., Yáñez-Ruiz, D.R., O’Kiely, P., Kebreab, E., Eugenè, M.A., Yu, Z., Shingfield, K.J., Schwarm, A., Hristov, A.N., and Reynolds, C.K. 2016A. Review of current in vivo measurement techniques for quantifying enteric methane emission from ruminants. Anim. Feed Sci. Technol. 219:13–30. doi:10.1016/j.anifeedsci.2016.05.018.</ref>), but routine individual measurements on a large scale (a requisite for genetic selection) have proven to be difficult to obtain and expensive to measure (Pickering et al., 2015<ref>Pickering, N.K., Oddy, V.H., Basarab, J.A., Cammack, K., Hayes, B J., Hegarty, R.S., McEwan, J.C., Miller, S., Pinares, C., and de Haas, Y. 2015. Invited review: Genetic possibilities to reduce enteric methane emissions from ruminants. Animal 9:1431-1440.</ref>; Negussie et al., 2016<ref>Negussie E., Lehtinen, J., Mäntysaari, P., Liinamo, A-E., Mäntysaari, E., and Lidauer, M.. 2016. Non-invasive individual methane measurements in dairy cows using photoacoustic infrared spectroscopy technique. 6th Greenhouse Gases Animal Agriculture Conference (GGAA2016) 14-18 February 2016. Melbourne, Australia. Abstract. p62.</ref>). Therefore, identifying proxies (i.e. indicators or indirect traits) that are correlated to CH<sub>4</sub> emissions, but which are easy and relatively lowcost to record on a large scale, would be a welcome alternative. Proxies might be less accurate but could be measured repeatedly to reduce random noise and in much larger populations. These guidelines are highly indebted to Garnsworthy et al. (2019)<ref name=":1">Garnsworthy, P.C. Difford, G.F. Bell, M.J. Bayat, A.R. Huhtanen, P. Kuhla, B. Lassen, J. Peiren, N. Pszczola, M; Sorg, D. Visker, M.H., and Yan, T. 2019 Comparison of Methods to Measure Methane for Use in Genetic Evaluation of Dairy Cattle. Animals 9:837, 12p.</ref>. In this paper the methods to measure CH<sub>4</sub> are compared with special emphasis to the genetic evaluation of dairy cattle.
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== Scope ==
== Scope ==
A variety of technologies are being developed and employed to measure CH<sub>4</sub> emissions of individual dairy cattle under various environmental conditions, as is evidenced by frequent reviews (Storm et al., 2012<ref>Storm, I.M., Hellwing, A.L.F., Nielsen, N.I., and Madsen, J. 2012. Methods for measuring and estimating methane emission from ruminants. Animals 2:160-183.</ref>; Cassandro et al., 2013<ref>Cassandro, M., Mele, M., Stefanon, B.. 2013. Genetic aspects of enteric methane emission in livestock ruminants. Italian J. Anim. Sci. 12:e73: 450-458.</ref>; Hammond et al., 2016A<ref>Hammond, K.J., Crompton, L.A., Bannink, A., Dijkstra, J., Yáñez-Ruiz, D.R., O’Kiely, P., Kebreab, E., Eugenè, M.A., Yu, Z., Shingfield, K.J., Schwarm, A., Hristov, A.N., and Reynolds, C.K. 2016A. Review of current in vivo measurement techniques for quantifying enteric methane emission from ruminants. Anim. Feed Sci. Technol. 219:13–30. doi:10.1016/j.anifeedsci.2016.05.018.</ref>; de Haas et al., 2017<ref>de Haas, Y., Pszczola, M., Soyeurt, H., Wall, E., and Lassen, J. 2017. Invited review: Phenotypes to genetically reduce greenhouse gas emissions in dairying. J. Dairy Sci. 100:855-870.</ref>). The first objective of the current guidelines is to review and compare the suitability of methods for large-scale measurements of CH<sub>4</sub> output of individual animals, which may be combined with other databases for genetic evaluations. Comparisons include assessing the accuracy, precision and correlation between methods. Combining datasets from different countries and research centres could be a successful strategy for making genetic progress in this difficult to measure trait if the methods are correlated (de Haas et al., 2017<ref name=":0" />). Accuracy and precision of methods are important. Data from different sources need to be appropriately weighted or adjusted when combined, so any methods can be combined if they are suitably correlated with the ‘true’ value. The second objective of the current guidelines, therefore, is to examine correlations among results obtained by different methods, ultimately leading to an estimate of confidence limits for selecting individual animals that are high or low emitters (see also Garnsworthy et al., 2019<ref name=":1" />).
A variety of technologies are being developed and employed to measure CH<sub>4</sub> emissions of individual dairy cattle under various environmental conditions, as is evidenced by frequent reviews (Storm et al., 2012<ref>Storm, I.M., Hellwing, A.L.F., Nielsen, N.I., and Madsen, J. 2012. Methods for measuring and estimating methane emission from ruminants. Animals 2:160-183.</ref>; Cassandro et al., 2013<ref>Cassandro, M., Mele, M., Stefanon, B.. 2013. Genetic aspects of enteric methane emission in livestock ruminants. Italian J. Anim. Sci. 12:e73: 450-458.</ref>; Hammond et al., 2016A<ref>Hammond, K.J., Crompton, L.A., Bannink, A., Dijkstra, J., Yáñez-Ruiz, D.R., O’Kiely, P., Kebreab, E., Eugenè, M.A., Yu, Z., Shingfield, K.J., Schwarm, A., Hristov, A.N., and Reynolds, C.K. 2016A. Review of current in vivo measurement techniques for quantifying enteric methane emission from ruminants. Anim. Feed Sci. Technol. 219:13–30. doi:10.1016/j.anifeedsci.2016.05.018.</ref>; de Haas et al., 2017<ref>de Haas, Y., Pszczola, M., Soyeurt, H., Wall, E., and Lassen, J. 2017. Invited review: Phenotypes to genetically reduce greenhouse gas emissions in dairying. J. Dairy Sci. 100:855-870.</ref>). The first objective of the current guidelines is to review and compare the suitability of methods for large-scale measurements of CH<sub>4</sub> output of individual animals, which may be combined with other databases for genetic evaluations. Comparisons include assessing the accuracy, precision and correlation between methods. Combining datasets from different countries and research centres could be a successful strategy for making genetic progress in this difficult to measure trait if the methods are correlated (de Haas et al., 2017<ref name=":0" />). Accuracy and precision of methods are important. Data from different sources need to be appropriately weighted or adjusted when combined, so any methods can be combined if they are suitably correlated with the ‘true’ value. The second objective of the current guidelines, therefore, is to examine correlations among results obtained by different methods, ultimately leading to an estimate of confidence limits for selecting individual animals that are high or low emitters (see also Garnsworthy et al., 2019<ref name=":1" />).
= Methane determining factors =
<div class="mw-collapsible mw-collapsed">
    <div>
== Diet and rumen microbiota ==
Table 1 contains a list of dietary or microbiota factors that determine CH<sub>4</sub> production.
{| class="wikitable"
|+
!Factors
!Reference
|-
|The main determinants of daily methane production are dry matter intake and diet composition: the more feed consumed, and/or the greater the fibre content of the diet, the more methane is produced per day. However, per unit of DMI, and per unit of fat+protein yield the grass diet produced less enteric CH<sub>4</sub> per cow than the TMR diet. Nutritional approaches for methane mitigation include reducing the forage to concentrate ratio of diets, increasing dietary oil content, and dietary inclusion of rumen modifiers and methane inhibitors.
|Beauchemin et al., 2009<ref>Beauchemin, K.A., McAllister, T.A., and McGinn, S.M. 2009. Dietary mitigation of enteric methane from cattle. CAB Rev.: Perspect. Agric., Vet. Sci., Nutr. Nat. Res. 4:1–18.</ref>;
Cottle et al., 2011<ref>Cottle, D.J., Nolan, J.V., and Wiedemann, S.G. 2011. Ruminant enteric methane mitigation: A review. Anim. Prod. Sci. 51:491–514. doi:10.1071/AN10163.</ref>; Knapp et al., 2014<ref>Knapp, J.R., Laur, G.L., Vadas, P.A., Weis,s W.P., and Tricarico, J.M. 2014. Invited review: Enteric methane in dairy cattle production: Quantifying the opportunities and impact of reducing emissions. J. Dairy Sci. 97:3231-3261.</ref>; O’Neill et al., 2011<ref>O’Neill, B.F., Deighton, M.H., O’Loughlin, B.M., Mulligan, F.J., Boland, T.M., O’Donovan, M., and Lewis, E. 2011. Effects of a perennial ryegrass diet or total mixed ration diet offered to spring-calving Holstein-Friesian dairy cows on methane emissions, dry matter intake, and milk production. J. Dairy Sci. 94:1941 – 1951</ref>; Sauvant et al., 2011<ref>Sauvant, D., Giger-Reverdin, S., Serment, A., and Broudiscou, L. 2011. Influences des régimeset de leur fermentation dans le rumen sur la production de méthane par les ruminants. INRA Prod. Anim. 24:433–446.</ref>
|-
|Methane output per kg of product is affected mainly by cow milk yield or growth rate, and by herd-level factors, such as fertility, disease incidence and replacement rate.
|Garnsworthy, 2004<ref>Garnsworthy, P.C. 2004. The environmental impact of fertility in dairy cows: a modelling approach to predict methane and ammonia emissions. Anim. Feed Sci. Technol. 112:211-223.</ref>
|-
|Methane output varies considerably between individual animals. For animals fed the same feed, the between-animal coefficient of variation (CV) in methane was 8.1%.
|Blaxter and Clapperton, 1965<ref>Blaxter, K.L., and Clapperton, J.L. 1965. Prediction of the amount of methane produced by ruminants. Br. J. Nutr.19:511–522.</ref>
|-
|The amount of digestible nutrients consumed especially of the carbohydrate fraction (starch, sugar, N-free residuals) is reliable to estimate CH<sub>4</sub> release with high precision. Furthermore, diets rich in fat reduced CH<sub>4</sub> formation in the rumen.
|Jentsch et al., 2007<ref>Jentsch, W., Schweigel, M., Weissbach, F., Scholze, H., Pitroff, W., and Derno, M. 2007. Methane production in cattle calculated by the nutrient composition of the diet. Arch. Anim. Nutr. 61:10-19.</ref>
|-
|DMI was also the most important determining factor, but there were different regression lines for maize silage and dried grass as the main roughage component respectively:
<math>CH4\ (g)=93+16.8\times DMI(kg)</math>
<math>CH4\ (g)=81+14.0\times DMI(kg)</math>
Methane release was particularly dependent on the intake of crude fiber (CF) and ether extract (EE):
<math>CH4\ (g)=63+80xCF\ (kg)+11xNFE\ (kg)+19xCP(kg)-195xEE\ (kg)</math>
|Kirchgessner et al., 1991<ref>Kirchgessner, M., Windisch, W., Müller, H. L., and Kreuzer, M. 1991. Release of methane and of carbon dioxide by dairy cattle. Agribiol. Res. 44:91-102.</ref>
|-
|Methane linearly increased with NDF intake <math>CH4\ (L)=59.4\times NDF[kg]+ 64.6</math> for cows together with their calves independent of the breed.
|Estermann et al., 2002<ref>Estermann, B.L., Sutter, F., Schlegel, P.O., Erdin, D., Wettstein, H.R., and Kreuzer, M. 2002. Effect of calf age and dam breed on intake, energy expenditure, and excretion of nitrogen, phosphorus, and methane of beef cows with calves. J. Anim. Sci. 80:1124-1134.</ref>
|-
|Enteric CH<sub>4</sub> could be predicted with the equation:
<math>CH4\ (g/d)=84+47\times cellulose(kg/d)+32\times starch(kg/d)+62\times sugars\ (kg/d)</math>
|Hindrichsen et al., 2005<ref>Hindrichsen, I.K., Wettstein, H.R., Machmüller, A., Jörg, B., and Kreuzer, M. 2005. Effect of the carbohydrate composition of feed concentratates on methane emission from dairy cows and their slurry. Environ. Monit. Assess., 107:329-350.</ref>
|-
|The higher the percentage concentrate the lower Ym.
|Zeitz et al., 2012<ref>Zeitz, J.O., Soliva, C.R., and Kreuzer, M. 2012. Swiss diet types for cattle: how accurately are they reflected by the Intergovernmental Panel on Climate Change default values? J. Int. Environ. Sci. 9(sup1):199-216.</ref>
|-
|Additives can sometimes have a methane reducing effect: higher dosages mitigate methane more. Saponins mitigate methanogenesis by reducing the number of protozoa, whereas condensed tannins act both by reducing the number of protozoa and by a direct toxic effect on methanogens.
|Beauchemin et al., 2008;<ref>Beauchemin, K.A., Kreuze,r M., O’Mara, F., and McAllister, T.A. 2008. Nutritional management for enteric methane abatement: A review. Aust. J. Exp. Agric. 48:21–27.</ref>
Jayanegara et al.<ref>Jayanegara, A., Leiber, F., and Kreuzer, M. 2012. Meta‐analysis of the relationship between dietary tannin level and methane formation in ruminants from in vivo and in vitro experiments. J. Anim. Physiol. Anim. Nutr. 96:365-375.</ref>, 2012; Zmora et al., 2012<ref>Zmora, P., Cieslak, A., Pers-Kamczyc, E., Nowak, A., Szczechowiak, J. and Szumacher-Strabel, M. 2012. Effect of Mentha piperita L. on in vitro rumen methanogenesis and fermentation, Acta Agr. Scan. Section A — Anim. Sci. 62:46-52, DOI: 10.1080/09064702.2012.703228.</ref>; Cieslak et al., 2013<ref>Cieslak, A., Szumacher-Strabel, M., Stochmal, A., nad Oleszek, W. 2013. Plant components with specific activities against rumen methanogens. Animal, 7(s2):253-265.</ref>; Guyader et al., 2014<ref>Guyader, J., Eugène, M., Nozière, P., Morgavi, D.P., Doreau, M., and Martin, C. 2014. Influence of rumen protozoa on methane emission in ruminants: a meta-analysis approach. Animal 8:1816-1825.</ref>
|-
|Plant essential oils have been shown as promising feed additives to mitigate CH<sub>4</sub> and ammonia emission, but results were inconsistent.
|Cobellis et al., 2016;<ref>Cobellis, G., Trabalza-Marinucci, M. and Yu, Z. 2016. Critical evaluation of essential oils as rumen modifiers in ruminant nutrition: A review. Sci. Total Environ. 545: 556-568.</ref>
Moate et al., 2011<ref>Moate, P.J., Deighton, M.H., Williams, S.R.O., Pryce, J.E., Hayes, B.J., Jacobs, J.L., Eckard, R.J., Hannah, M.C. and Wales, W.J., 2016. Reducing the carbon footprint of Australian milk production by mitigation of enteric methane emissions. Anim. Prod. Sci. 56:1017-1034.</ref>
|-
|Nitrate and sulphate addition decreased the enteric methane emissions negatively affecting diet digestibility and milk production. The effects of the salts are additive.
|van Zijderveld et al., 2010<ref>Van Zijderveld, S.M., Gerrits, W.J.J., Apajalahti, J.A., Newbold, J.R., Dijkstra, J., Leng, R A., and Perdok, H.B. 2010. Nitrate and sulfate: effective alternative hydrogen sinks for mitigation of ruminal methane production in sheep. J. Dairy Sci. 93:5856-5866.</ref>;
van Zijderveld et al., 2011<ref>Van Zijderveld, S.M., Gerrits, W.J.J., Dijkstra, J., Newbold, J.R., Hulshof, R.B.A., and Perdok, H.B. 2011. Persistency of methane mitigation by dietary nitrate supplementation in dairy cows. J. Dairy Sci. 94:4028-4038.</ref>
|-
|The methanogenesis in the rumen of calves is associated with the development of the ruminal protozoa population. The absence of protozoa in the rumen reduced both the CH<sub>4</sub> production and the digestibility of carbohydrates.
|Schönhusen et al., 2003<ref>Schönhusen, U., Zitnan, R., Kuhla, S., Jentsch, W., Derno, M., and Voigt, J. 2003. Effects of protozoa on methane production in rumen and hindgut of calves around time of weaning. Arch. Anim. Nutr. 57:279-295.</ref>
|-
|Implementing good grazing management reduced gross energy intake loss as CH<sub>4</sub> by 14%.
|Wims et al., 2010<ref>Wims, C.M., Deighton, M.H., Lewis, E., O’Loughlin, B., Delaby, L., Boland, T.M., and O’Donovan, M. 2010. Effect of pregrazing herbage mass on methane production, dry matter intake, and milk production of grazing dairy cows during the mid-season period. J. Dairy Sci. 93:4976 – 4985</ref>
|}Table 1. Methane determining factors related to diet and rumen microbiota.
== Host genetics, physiology and environment ==
A low-moderate proportion of variation in CH<sub>4</sub> emissions among ruminants is under genetic control. Heritability coefficients of MeY and RMPR were h²=0.22 and 0.19 respectively in a population of 1,043 Angus growing steers and heifers measured during 2 days in RC (Donoghue et al., 2016<ref>Donoghue, K.A., Bird-Gardiner, T., Arthur, P.F., Herd, R.M., and Hegarty, R.F. 2016. Genetic and phenotypic variance and covariance components for methane emission and postweaning traits in Angus cattle. J. Anim. Sci. 94:1438–1445. doi:10.2527/jas2015-0065.</ref>). The heritability coefficient of MeY was h²=0.13 in a population of1,225 dual-purpose growing sheep measured during 2 days in RC (Pinares-Patino et al., 2013<ref>Pinares-Patiño C.S., Hickey, S.M., Young, E.A., Dodds, K.G., MacLean, S., Molano, G., Sandoval, E., Kjestrup, H., Harland, R., Pickering, N.K., and McEwan, J.C. 2013. Heritability estimates of methane emissions from sheep. Animal 7: 316–321.</ref>). Table 2 contains information of heritability of traits related to CH<sub>4</sub> production.
{| class="wikitable"
|+
!Factors
!Reference
|-
|List with several h<sup>2</sup>
|Pickering et al., 2015<ref>Pickering, N.K., Oddy, V.H., Basarab, J.A., Cammack, K., Hayes, B J., Hegarty, R.S., McEwan, J.C., Miller, S., Pinares, C., and de Haas, Y. 2015. Invited review: Genetic possibilities to reduce enteric methane emissions from ruminants. Animal 9:1431-1440.</ref>
|-
|List with several h<sup>2</sup>
|MPWG White paper Dec 18<ref>MPWG White paper Dec 18. <nowiki>http://www.asggn.org/publications,listing,95,mpwg-white-paper.html</nowiki>. Pickering, N.K., de Haas, Y., Basarab, J., Cammack, K., Hayes, B., Hegarty, R.S., Lassen, J., McEwan, J.C., Miller, S., Pinares-Patiño, C.S., Shackell, G., Vercoe, P. and Oddy, V.H. 2013.</ref>
|-
|Methane emissions from individual cows during milking varied between individuals with the same milk yield and fed the same diet. Between-cow variation in MERm is greater than within-cow variation and ranking of cows for CH<sub>4</sub> emissions is consistent across time. Variation related to body weight, milk yield, parity, and week of lactation/days in milk. The monitored variation might offer opportunities for genetic selection.
|Garnsworthy et al., 2011A<ref>Garnsworthy, P.C., Craigon, J., Hernandez-Medrano, J.H. and Saunders, H. 2012A. On-farm methane measurements during milking correlate with total methane production by individual dairy cows. J. Dairy Sci. 95:3166-3180.</ref>;
Garnsworthy et al., 2011B<ref>Garnsworthy, P.C., Craigon, J., Hernandez-Medrano, J.H., and Saunders, N. 2012B. Variation among individual dairy cows in methane measurements made on farm during milking. J. Dairy Sci. 95:3181–3189.</ref>
|-
|Mechanistic modelling approach: potential for dietary intervention as a means of substantially reducing CH<sub>4</sub> emissions without adverse effects on dietary energy supply.
|Mills et al., 2001<ref>Mills, J.A.N., Dijkstra, J., Bannink, A., Cammell, S.B., Kebreab, E., and France, J. 2001. A mechanistic model of whole-tract digestion and methanogenesis in the lactating dairy cow: model development, evaluation, and application. J. Anim. Sci. 79:1584-1597.</ref>
|-
|The CH<sub>4</sub>-to-CO<sub>2</sub> ratio measured using the non-invasive portable air sampler and analyzer unit based on Fourier transform infrared (FTIR) detection method is an asset of the individual cow and may be useful in both management and genetic evaluations.
|Lassen et al., 2012<ref>Lassen, J., Lovendahl, P., and Madsen, J. 2012. Accuracy of noninvasive breath methane measurements using Fourier transform infrared methods on individual cows. J. Dairy Sci. 95:890-898.</ref>
|-
|The estimated heritability for CH<sub>4</sub> g/day and CH<sub>4</sub> g/kg of FPCM were lower than common production traits but would still be useful in breeding programs.
|Kandel et al., 2013<ref>Kandel, P.B., Vanrobays, M.L., Vanlierde, A., Dehareng, F., Froidmont, E., Dardenne, P., Lewis, E., Buckley, F., Deighton, M.H., McParland, S. and Gengler, N., 2013. Genetic parameters for methane emissions predicted from milk mid-infrared spectra in dairy cows. J. Dairy Sci. 95(E-1):p.388.</ref>
|-
|Genetic correlation between CH<sub>4</sub> intensity and milk yield (MY) was - 0.67 and with milk protein yield (PY) was -0.46 in Holstein cows.
|Kandel et al., 2014A, B<ref>Kandel, P.B., Vanderick, S., Vanrobays, M.L., Vanlierde, A., Dehareng, F., Froidmont, E., Soyeur,t H., and Gengler, N. 2014B. Consequences of selection for environmental impact traits in dairy cows. In: 10th World Congress on Genetics Applied to Livestock Production (WCGALP), 17-22 August, 2014. Vancouver, Canada.</ref>
|-
|Milk production and CH<sub>4</sub> emissions of dairy cows seemed to be influenced by the temperature humidity index.
|Vanrobays et al., 2013A<ref>Vanrobays, M.-L., Gengler, N., Kandel, P.B., Soyeurt, H., and Hammami, H. 2013A. Genetic effects of heat stress on milk yield and MIR predicted methane emissions of Holstein cows. 64th Annual meeting of the European Federation of Animal Science, p498</ref>
|-
|Estimate the heritability of the estimated methane emissions from 485 Polish Holstein-Friesian dairy cows at 2 commercial farms using FTIR spectroscopy during milking in an automated milking system by implementing the random regression method. The heritability level fluctuated over the course of lactation, starting at 0.23 (SE 0.12) and then increasing to its maximum value of 0.3 (SE 0.08) at 212 DIM and ending at the level of 0.27 ± 0.12. Average heritability was 0.27 ± 0.09.
|Pszczola et al., 2017<ref>Pszczola, M., Rzewuska, K., Mucha, S., and Strabel, T. 2017. Heritability of methane emissions from dairy cows over a lactation measured on commercial farms. J. Anim. Sci. 95:4813-4819. doi: 10.2527/jas2017.1842.</ref>
|-
|CH<sub>4</sub> measured with a portable air-sampler FTIR detection method on 3,121 Holstein dairy cows from 20 herds using automatic milking systems. The heritability of CH<sub>4</sub>_MILK was 0.21 ± 0.06. It was concluded that a high genetic potential for milk production will also mean a high genetic potential for CH<sub>4</sub> production. The results suggested that CH<sub>4</sub> emission is partly under genetic control, that it is possible to decrease CH<sub>4</sub> emission from dairy cattle through selection, and that selection for higher milk yield will lead to higher genetic merit for CH<sub>4</sub> emission/cow per day.
|Lassen and Løvendahl, 2016<ref>Lassen, J., and Løvendahl, P. 2016. Heritability estimates for enteric methane emissions from Holstein cattle measured using noninvasive methods. J. Dairy Sci. 99:1959-1967.</ref>
|-
|CH<sub>4</sub> production was measured of 184 Holstein-Friesian cows in. the milking robot with a in total 2,456 observations for CH<sub>4</sub> production. Heritability for CH<sub>4</sub> production ranged from 0.12 ± 0.16 to 0.45 ± 0.11, and genetic correlations with MY ranged from 0.49 ± 0.12 to 0.54 ± 0.26. The positive genetic correlation between CH<sub>4</sub> production and milk yield indicates that care needs to be taken when genetically selecting for lower CH<sub>4</sub> production, to avoid a decrease in MY at the animal level. However, this study shows that CH<sub>4</sub> production is moderately heritable and therefore progress through genetic selection is possible.
|Breider et al., 2019<ref>Breider, I.S., Wall, E., Garnsworthy, P.C. 2019. Short communication: Heritability of methane production and genetic correlations with milk yield and body weight in Holstein-Friesian dairy cows, J. Dairy Sci. 102: 7277-7281.</ref>
|-
|CH<sub>4</sub> concentration was measured with NDIR, and CH<sub>4</sub> production was estimated from CH<sub>4</sub> concentration and body weight. Heritability for CH<sub>4</sub> concentration was 0.11 ± 0.03 and for CH<sub>4</sub> production 0.12 ± 0.04. Positive genetic correlation was observed with MY (0.17-0.21), PY (0.22-0.31) and FY (0.27-0.29). Other type traits showed positive correlation with methane production (chest width=0.26, angularity =0.19, stature = 0.43 and capacity = 0.31) possibly associated to higher milk feed intake from these animals. Rumination time was negatively correlated to CH<sub>4</sub> production (-0.24) and CH<sub>4</sub> concentration (-0.43). However, larger CH<sub>4</sub> production and CH<sub>4</sub> concentration was associated with shorter days open.
|López-Paredes et al. (2020)<ref>Lopez-Paredes, J., Goiri, I., Atxaerandio, R., García-Rodríguez, A., Ugarte, E., Jiménez-Montero, J.A., Alenda, R and  González-Recio, O. 2020. Mitigation of greenhouse gases in dairy cattle via genetic selection (i): Genetic parameters of direct methane using non-invasive methods and its proxies. J. Dairy Sci. 103.</ref>
|-
|Genetic parameters of CH<sub>4</sub> emissions predicted from milk fatty acid profile (FA) and those of their predictors in 1,091 Brown Swiss cows reared on 85 farms showed that enteric CH<sub>4</sub> emissions of dairy cows can be estimated on the basis of milk fatty acid profile. Additive genetic variation of CH<sub>4</sub> traits was shown which could be exploited in breeding programmes.
|Bittante and Cecchinato, 2020<ref>Bittante, G., and Cecchinato, A. 2020. Heritability estimates of enteric methane emissions predicted from fatty acid profiles, and their relationships with milk composition, cheese-yield and body size and condition, It. J. An. Sci. 19:114-126, DOI: 10.1080/1828051X.2019.1698979</ref>
|-
|A total of 670 test day records were recorded on lactating Holstein Friesian cows reared in 10 commercial dairy herds. Predicted methane production (PMP) was estimated to be 15.33±1.52 MJ/d in dairy cows with 23.53±6.81 kg/d of milk yeild (MY) and 3.57±0.68% of fat content (FC). Heritability of MY was 0.09 with a posterior probability for values of h<sup>2</sup> greater than 0.10 of 44%. Estimates of heritability for FC and protein content (PC) were 0.17 and 0.34, respectively, with a posterior probability for values of h<sup>2</sup> greater than 0.10 of 77% and 99%. For somatic cell score (SCS), heritability was 0.13 with a posterior probability for values of h2 greater than 0.10 of 67%. Heritability for the trait PMP was moderate to low (0.12); however, posterior probability for values of h<sup>2</sup> greater than 0.10 was 60%. Medians of the posterior distributions of genetic correlations between PMP and milk production traits were: 0.92, 0.67, 0.14, and 0.14 between PMP and MY, PMP and FC, PMP and PC, and PMP and SCS, respectively. Reduction of PMP seems to be viable through selection strategies without affecting udder health and PC.
|Cassandro et al., 2010<ref>Cassandro, M., Cecchinato, A., Battagin, M., Penasa, M., 2010. Genetic parameters of predicted methane production in Holstein Friesian cowsIn: Proc. 9th World Congr. on Genetics Applied to Livestock Production, Leipzig, Germany. . Page 181</ref>
|-
|GWAS to study the genetic architecture of CH<sub>4</sub> production and detected genomic regions affecting CH<sub>4</sub> production. Detected regions explained only a small proportion of the heritable variance. Potential QTL regions affecting CH<sub>4</sub> production were located within QTLs related to feed efficiency, milk-related traits, body size and health status. Five candidate genes were found: CYP51A1 on BTA 4, PPP1R16B on BTA 13, and NTHL1, TSC2, and PKD1 on BTA 25. These candidate genes were involved in a number of metabolic processes that are possibly related to CH<sub>4</sub> production. One of the most promising candidate genes (PKD1) was related to the development of the digestive tract. The results indicate that CH<sub>4</sub> production is a highly polygenic trait.
|Pszczola et al., 2018<ref>Pszczola, M., Strabel, T., Mucha, S., and Sell-Kubiak, E. 2018. Genome-wide association identifies methane production level relation to genetic control of digestive tract development in dairy cows. Scientific Rep. 8 (1), 15164 <nowiki>https://doi.org/10.1038/s41598-018-33327-9</nowiki></ref>
|-
|A 1000-cow study across European countries revealed that the ruminant microbiomes can be controlled by the host animal. A 39- member subset of the core microbiome formed hubs in co-occurrence networks linking microbiome structure to host genetics and phenotype (CH<sub>4</sub> emissions, rumen and blood metabolites, and milk production efficiency).
|Wallace et al.,
2019<ref>Wallace, R.J., Sasson, G., Garnsworthy, P.C., Tapio, I., Gregson, E., Bani, P., Huhtanen, P., Bayat, A.R., Strozzi, F., Biscarini, F., Snelling, T.J., Saunders, N., Potterton, S.L., Craigon, J., Minuti, A., Trevisi, E., Callegari, M.L., Cappelli, F.P., Cabezas-Garcia, E.H., Vilkki, J., Pinares-Patino, C., Fliegerov, K.O., Mrazek, J., Sechovcova, H., Kope, J., Bonin, A., Boyer, F., Taberlet, P., Kokou, F., Halperin, E., Williams, J.L., Shingfield, K.J., and Mizrahi, I. 2019. A heritable subset of the core rumen microbiome dictates dairy cow productivity and emissions. Sci. Adv. 5:(7):eaav8391. doi 10.1126/sciadv.aav8391.</ref>
|}Table 2. Heritability information of methane-related traits and measurements.</div>
</div>
= Methane measurements methods =
= Discussion of methods =
= Comparison of methods to measure methane =
= Proxies =
= Proxies discussion =
= Merging and sharing data in genetic evaluations =
= Recommendations =
= Conclusions =


== Sub-sections ==
== Sub-sections ==
<div style="column-count:2">
<div style="column-count:2">
:[[Definition and Terminology]]
:[[Section 20: Definition and Terminology |Definition and Terminology]]
:[[Methane determining factors]]
:[[Section 20: Methane determining factors |Methane determining factors]]
:[[Methane measuring methods|Methane measurements methods]]
:[[Section 20: Methane measuring methods|Methane measurements methods]]
:[[Discussion of methods]]
:[[Section 20: Discussion of methods |Discussion of methods]]
:[[Comparison of methods to measure methane]]
:[[Section 20: Comparison of methods to measure methane |Comparison of methods to measure methane]]  
:[[Proxies]]
:[[Section 20: Proxies |Proxies]]
:[[Proxies Discussion|Proxies discussion]]
:[[Section 20: Proxies Discussion|Proxies discussion]]
:[[Merging and sharing data in genetic evaluations]]
:[[Section 20: Merging and sharing data in genetic evaluations |Merging and sharing data in genetic evaluations]]
:[[Recommendations|Reccomendations]]
:[[Section 20: Ongoing activities |Ongoing activities]]
:[[Conclusions]]
:[[Section 20: Recommendations|Recomendations]]
:[[Section 20: Conclusions |Conclusions]]
[[Completed Activities]]
</div>
</div>



Latest revision as of 10:06, 12 May 2025

NOTE: This version of Section 20 has been approved by the working group's Chair. Please be aware that further revisions may occur before final review and approval by the Board and ICAR members per the Approval of Page Process.

Introduction

Increases in milk production through management and genetics have substantially improved feed efficiency and decreased costs per unit of product over recent decades. However, dairy systems are also associated with environmental costs (Baskaran et al., 2009[1]), with methane (CH4) emissions associated with rumen microbial fermentation being both an important contributor to global greenhouse gas (GHG) emissions, as well as an avoidable loss of energy that could otherwise be directed into milk production. The livestock sector is responsible for 14.5% of the global GHG (Gerber et al., 2013[2]); dairy cattle account for 18.9% of these emissions, mainly in the form of enteric CH4 emissions (van Middelaar et al., 2014[3]). Methane is a greenhouse gas with a global warming potential 28 times that of CO2 (Myhre et al., 2013[4]). Methane from ruminant livestock is generated during microbial fermentation in the rumen and hindgut (enteric CH4), and from decomposition of manure. Enteric CH4 contributes 80% of CH4 emissions by ruminants, and manure decomposition contributes 20%. Enteric CH4 accounts for 17% of global CH4 emissions and 3.3% of total global greenhouse gas emissions from human activities (Knapp et al., 2014[5]). There is, therefore, a significant research interest to find ways to reduce enteric CH4 emissions by ruminants. Ruminant animals have a digestive system to digest plant materials efficiently. Like most mammals, ruminants lack the cellulase enzyme required to break the beta-glucose linkages in cellulose, but they play host to diverse populations of rumen microbes that can digest cellulose and other plant constituents. When rumen bacteria, protozoa and fungi ferment carbohydrates and proteins of plant materials, they produce volatile fatty acids, principally acetate, propionate and butyrate. High fibre diets favour acetate synthesis. Synthesis of acetate and butyrate are accompanied by release of metabolic hydrogen, which, if allowed to accumulate in rumen fluid, has negative effects on microbial growth, and feed digestibility (Janssen, 2010[6]). Rumen Archaea are microorganisms that combine metabolic hydrogen with CO2 to produce CH4 and water. Archaea play a vital role, therefore, in protecting the rumen from excess metabolic hydrogen, and the CH4 they produce is an inevitable product of rumen fermentation. A number of CH4 phenotypes have been defined (Hellwing et al., 2012[7]); the most widely used is CH4 production (MeP) in liters or grams per day. The CH4 production trait is highly correlated with feed intake (Basarab et al., 2013[8]; De Haas et al., 2017[9]) and, thereby, with the ultimate breeding goal trait: milk production in dairy cattle. The economic value of daily dry matter intake and associated methane emissions in dairy cattle showed that increasing the feed performance estimated breeding value by one unit (i.e. 1 kg of more efficiently converted DMI during the cow’s first lactation) translates to a total lifetime saving of 3.23 kg in DMI and 0.055 kg in methane (Richardson et al., 2019[10]). Feed Performance was defined as a 1 kg increase in more efficiently used feed in a first parity lactating cow. These results show not only the relation between DMI and CH4 production, but also the economic relationship between these traits. Persistency of lactation was found to be positively associated with increased feed efficiency and decreased methane production and intensity. Feed efficiency was associated with lower methane intensity. Feed efficiency and methane emissions can be improved by selecting for dairy cattle that are smaller and have increased persistency of lactation. Efficiency and methane emissions can be further improved by improved management of body condition score and by extending lactations beyond the conventional 305-day length (Seymour, 2019[11]). According to Ellis et al. (2007)[12], DMI predicted MeP with an R2 of 0.64, and ME intake (MJ/d) predicted MeP with an R2 of 0.53 for dairy cattle. AlternativePhenotype definitions include CH4 intensity (MeI), which is defined as liters or grams of CH4 per kg of milk, and CH4 yield (MeY), which is defined as liters or grams of CH4 per kg of dry matter intake (DMI) (Moate et al., 2016[13]). Residual CH4 production (RMP) is calculated as observed minus predicted CH4 production (Herd et al., 2014[14], Berry et al., 2015[15]), with predicted values based on factors such as milk production, body weight and feed intake. At the moment, it is not obvious which of these phenotypes to use; but, it is important to monitor associations between the chosen CH4 phenotype and the other important traits in the breeding goal (e.g. production, fertility, longevity) to avoid unfavorable consequences. Berry and Crowley (2012)[16] describe advantages and limitations of ration traits. For example, because feed efficiency traits are a linear combination of other traits it is not recommended to include them in an overall total merit index, which is a clear limitation. For all applications it is necessary to measure the CH4 emission of each animal individually. These guidelines are intended to make the right choices for this. Whilst diet changes and feed additives can be effective mitigation strategies for CH4 emissions (Beauchemin et al., 2009[17]; Martin et al., 2010[18]; Hristov et al., 2013[19]), their effects depend on the continued use of a particular diet or additive and there have been issues with the rumen microbiomes adapting to additives. Rumen bacterial communities are highly dynamic after a diet switch and did not stabilize within 5 wk of cows grazing pasture (Bainbridge et al., 2016[20]). In contrast, breeding for reduced CH4 emissions should result in a permanent and cumulative reduction of emissions (Wall et al., 2010[21]). Several studies have shown that CH4 emissions by ruminants have a genetic component, with heritability in the range 0.20 – 0.30 (de Haas et al., 2011[22]; Donoghue et al., 2013[23]; Pinares-Patiño et al., 2013[24], Kandel et al., 2014A[25], B[26]; Lassen and Lovendahl, 2016[27]; López-Paredes et al. 2020[28]). Breeding for reduced CH4 emissions, alone or together with other mitigation strategies, could therefore be effective in reducing the environmental impact of cattle farming and, possibly, also in increasing feed efficiency. Such a breeding scheme would require, as a fundamental starting point, accurate measures of individual CH4 emissions on a large scale. Several techniques have been developed for the measurement of CH4 emissions from ruminants, with varying degrees of accuracy (see reviews by Cassandro et al., 2013[29] and Hammond et al., 2016A[30]), but routine individual measurements on a large scale (a requisite for genetic selection) have proven to be difficult to obtain and expensive to measure (Pickering et al., 2015[31]; Negussie et al., 2016[32]). Therefore, identifying proxies (i.e. indicators or indirect traits) that are correlated to CH4 emissions, but which are easy and relatively lowcost to record on a large scale, would be a welcome alternative. Proxies might be less accurate but could be measured repeatedly to reduce random noise and in much larger populations. These guidelines are highly indebted to Garnsworthy et al. (2019)[33]. In this paper the methods to measure CH4 are compared with special emphasis to the genetic evaluation of dairy cattle.

Disclaimer

The fact that specific device manufacturers are mentioned in these guidelines is in no way an endorsement of the devices or their accuracy by ICAR.

Scope

A variety of technologies are being developed and employed to measure CH4 emissions of individual dairy cattle under various environmental conditions, as is evidenced by frequent reviews (Storm et al., 2012[34]; Cassandro et al., 2013[35]; Hammond et al., 2016A[36]; de Haas et al., 2017[37]). The first objective of the current guidelines is to review and compare the suitability of methods for large-scale measurements of CH4 output of individual animals, which may be combined with other databases for genetic evaluations. Comparisons include assessing the accuracy, precision and correlation between methods. Combining datasets from different countries and research centres could be a successful strategy for making genetic progress in this difficult to measure trait if the methods are correlated (de Haas et al., 2017[9]). Accuracy and precision of methods are important. Data from different sources need to be appropriately weighted or adjusted when combined, so any methods can be combined if they are suitably correlated with the ‘true’ value. The second objective of the current guidelines, therefore, is to examine correlations among results obtained by different methods, ultimately leading to an estimate of confidence limits for selecting individual animals that are high or low emitters (see also Garnsworthy et al., 2019[33]).

Sub-sections

Summary of Changes

Date of change Nature of change
March 2020 Draft from Feed & Gas WG put into standard template for ICAR Guidelines. Separate out EDGP database to become a standalone appendix.
April 2020 Edits and acknowledgements added by Feed & Gas WG.
May 2020 Approved by ICAR Board on 26th May subject to addition of disclaimer.

Disclaimer added as new chapter 2 - the fact specific device manufacturers are mentioned in these guidelines is in no way an endrosement of the devices or their accuracy by ICAR.

December 2023 Creation of Methane Emission for Genetic Evaluation Wiki Page.

References

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