反刍动物瘤胃产氢微生物构成及其对甲烷生成的调控

doi:10.3864/j.issn.0578-1752.2026.20.016

Abstract

Abstract:

Methane (CH₄) is the second most significant greenhouse gas globally, yet due to its potent short-term warming effect, it has a pronounced impact on near-term global climate warming. In recent years, atmospheric CH₄ concentrations have continued to rise, reaching 1.6 times pre-industrial levels by 2024. CH₄ emissions contribute to global warming, threatening human living environments and health, with the social cost of each ton of CH₄ emissions amounting to thousands of dollars. Ruminants represent one of the primary anthropogenic sources of CH₄ emissions in the agricultural sector, making their CH₄ mitigation crucial. The CH₄ emitted by ruminants is predominantly generated by rumen microorganisms through hydrogenotrophic, methylotrophic, and acetate-cleavage pathways, with the hydrogenotrophic pathway, where hydrogen (H₂) reduces carbon dioxide (CO₂) to form CH₄, which dominates the process. Consequently, H₂ serves as a critical precursor in rumen CH₄ production, and appropriately reducing or consuming H₂ production during rumen microbial fermentation represents the most direct strategy for CH₄ mitigation. H₂ is produced by various H₂-producing microorganisms (protozoa, bacteria, and fungi), with protozoa primarily consisting of holotrichs, bacteria dominated by Clostridia within the Firmicutes phylum, and certain fungi also capable of H₂ production. Protozoa, bacteria, and fungi generate H₂ during the fermentation of rumen carbohydrates to produce volatile fatty acids. The H₂-producing capacities vary among different H₂-producing microorganisms. Specifically, the H₂-producing ability of protozoa is related to factors, such as body length and the number of hydrogenosomes, with holotrich protozoa and large protozoa being high-H₂-producing groups. For bacteria, their H₂-producing capacity is associated with the H₂-producing pathways they mediate, with acetate- and H₂-producing bacteria showing relatively high H₂-producing abilities. There are relatively few reports on fungi. The aforementioned high-H₂-producing microorganisms can serve as important targets for reducing the transfer of H₂ to methanogenic pathways and promoting CH₄ mitigation. Although there are differences in the H₂-producing capacities of various H₂-producing microorganisms, most of their H₂-producing pathways require the participation of hydrogenases. Hydrogenases can utilize reduced nicotinamide adenine dinucleotide (NADH) and reduced ferredoxin (Fd_red) generated during rumen fermentation to reduce H⁺ and produce H₂. In addition, the H₂-producing pathways of H₂-producing microorganisms are influenced by the composition of the microbial community, the nutritional structure of the diet, and functional additives. These factors affect H₂ production efficiency by regulating the rumen H₂-producing fermentation pattern, hydrogen partial pressure, and the expression of hydrogenase in vivo, as well as the activity of H₂-producing microorganisms. Based on these hydrogen production characteristics, H₂ production by H₂-producing microorganisms can be reduced to promote CH₄ mitigation: the community composition of rumen H₂-producing microorganisms can be optimized to promote low-H₂-producing microorganisms dominating rumen fermentation, such as feeding Bacillus subtilis; the expression of hydrogenase in H₂-producing microorganisms can also be regulated, for example, through dietary nutrient regulation; additionally, the activity of H₂-producing microorganisms can be directly inhibited, such as by adding plant extracts like tannins or traditional Chinese medicine formulas to the diet. This review summarized the types and characteristics of H₂-producing microorganisms in the rumen, providing a theoretical basis for developing CH₄ mitigation strategies at the source.

Key words: ruminants, methane, rumen, hydrogen-producing microorganisms, methane mitigation

JIA Peng, XIA Dong, SU YanJing, JIANG XueYuan, WEI ZiHai, LEI HuLong, KANG JinGuo, JIAO Chong, LU NaiSheng. Composition of Hydrogen-Producing Microorganisms in the Rumen of Ruminants and Their Regulation on Methanogenesis[J].Scientia Agricultura Sinica, 2026, 59(20): 4642-4654.

Figures/Tables 2

Fig. 1

Table 1

Hydrogen-Producing Bacteria in the Rumen of Ruminants"

产氢途径 H₂ production pathways	门 Phylum	科 Family	产氢细菌 Hydrogen-producing bacteria	参考文献 References
产乙酸 Acetate production	Firmicutes	Eubacteriaceae	瘤胃内存在的Acetobacterium woodii与耗H₂的Methanobacterium strain AZ体外共同培养时，可产生H₂ Acetobacterium woodii resident in the rumen can produce H₂ when co-cultured in vitro with the H₂-utilising Methanobacterium strain AZ	Winter et al.^[33]
		Selenomonadaceae	瘤胃内Anaerovibrio robustus中，涉及产氢的乙酸合成途径的酶PF02550表达量高 The expression level of enzyme PF02550 involved in the hydrogen-producing acetate synthesis pathway in Anaerovibrio robustus is elevated in the rumen	Hagen et al.^[17]
		Oscillospiraceae	瘤胃内Ruminococcus albus和R. flavefaciens与乙酸生成有关，具备产氢能力，受抑制时可降低产甲烷菌对H₂的利用；Ruminococcus spp.的代谢途径与H₂的产量直接相关 Ruminal Ruminococcus albus and R. flavefaciens are associated with acetate production and possess hydrogenogenic capacity; when inhibited, they can reduce the utilisation of H₂ by methanogens. The metabolic pathways of Ruminococcus spp. are directly linked to the yield of H₂ in the rumen	Belanche et al.^[34] Danielsson et al.^[35]
	Bacteroidota	Prevotellaceae	瘤胃内P. ruminicola (BPl.162）、P. ruminicola (KHT3)、F. succinogens中涉及产氢的乙酸合成途径的酶PF01515表达量高 In the rumen, high expression of the enzyme PF01515, which participates in the acetogenic pathway linked to hydrogenogenesis, is observed in P. ruminicola (BPl.162), P. ruminicola (KHT3), and F. succinogenes	Hagen et al.^[17]
	Fibrobacteres	Fibrobacteraceae		Hagen et al.^[17]
	Actinobacteria	Bifidobacteriaceae	瘤胃中的Bifidobacterium可产生乙酸和乳酸，其代谢过程伴随H₂的生成 Ruminal Bifidobacterium can produce acetate and lactate, and its metabolic process is accompanied by the generation of H₂	Danielsson et al.^[35]
产丁酸 Butyrate production	Firmicutes	Acidaminococcaceae	瘤胃内Acidaminococcus fermentans与瘤胃液丁酸盐的比例有关，且早期研究表明此菌属具备产乙酸、丁酸及产氢的能力 In the rumen, Acidaminococcus fermentans is associated with the proportion of butyrate in rumen fluid, and early studies have indicated that this species possesses the capacity to produce acetate, butyrate, and hydrogen	Xu et al.^[36] Härtel et al.^[37]
产丁酸 Butyrate production	Firmicutes	Lachnospiraceae	瘤胃微生物基因组中参与CH₄代谢的基因主要由Lachnospiraceae（主要表达菌属为Butyrivibrio）和产甲烷菌表达，Butyrivibrio可通过碳水化合物发酵产生丁酸和H₂ In the rumen microbial genome, genes involved in CH₄ metabolism are mainly expressed by Lachnospiraceae (with Butyrivibrio being the predominant genus responsible for their expression) and by methanogens. Butyrivibrio can produce butyrate and H₂ through carbohydrate fermentation	Huws et al.^[38]
产乙酸和丁酸 Acetate and butyrate production	Firmicutes	Lachnospiraceae	瘤胃Lachnospiraceae分离菌株NK3A20在属水平上是瘤胃细菌丰度最高的十个属之一。其基因组中含有多种氢化酶基因，可利用葡萄糖产生乙酸、丁酸、甲酸和H₂ At the genus level, the rumen Lachnospiraceae isolate NK3A20 is one of the ten most abundant bacterial genera in the rumen. Its genome contains multiple hydrogenase genes, and it can utilise glucose to produce acetate, butyrate, formate and H₂	Kaminsky et al.^[39]
		Ruminococcaceae	瘤胃内Ruminococcaceae和Lachnospiraceae在高CH₄产量绵羊瘤胃中富集，可通过碳水化合物（如己糖）发酵产生H₂ Ruminococcaceae and Lachnospiraceae are enriched in the rumen of sheep exhibiting high CH₄ production, and are capable of generating H₂ via fermentation of carbohydrates such as hexoses	Kamke et al.^[14]
		Coprobacillaceae	此过程每摩尔己糖产生2摩尔H₂ In the rumen of low-methane-yielding sheep, Sharpea spp. first ferment carbohydrates to lactate, which is subsequently converted to butyrate by Megasphaera spp.; this overall process produces 2 mol of H₂ per mole of hexose
		Veillonellaceae
甲酸产氢 Hydrogen production from formate	Firmicutes	Lachnospiraceae	瘤胃内的Prevotella brevis、P. ruminicola BPI-162、P. ruminicola KHT3、Fibrobacter succinogenes、Butyrivibrio fibrisolvens、Lachnospiraceae的UBA1225和UBA1213均编码与甲酸产CO₂和H₂相关的酶PF01558 In the rumen, Prevotella brevis, P. ruminicola strains BPI-162 and KHT3, Fibrobacter succinogenes, Butyrivibrio fibrisolvens, and the Lachnospiraceae strains UBA1225 and UBA1213 all encode enzymes of the PF01558 family, which are involved in the production of CO₂ and H₂ from formate	Hagen et al.^[17]
	Bacteroidota	Prevotellaceae
	Fibrobacteres	Fibrobacteraceae
未明确报道 Not clearly reported	Firmicutes	/	反刍动物全消化道微生物中用于发酵产氢的酶主要分布于Firmicutes（占比72.7%），瘤胃Firmicutes编码电子分叉和发酵型[FeFe]-氢化酶；瘤胃微生物中用于发酵产氢的酶主要分布于Firmicutes的Clostridia ，少量分布于Bacilli 中；瘤胃微生物中的电子分叉[FeFe]-氢化酶主要分布于Clostridia，推断Clostridia发酵是瘤胃中H₂的主要来源	Xie et al.^[40] Lin et al.^[41] Li et al.^[22] Greening et al.^[42]
			Across the entire gastrointestinal microbiota of ruminants, the enzymes involved in fermentative hydrogen production are predominantly found in Firmicutes (accounting for 72.7%), and ruminal Firmicutes encode both electron-bifurcating and fermentative [FeFe]-hydrogenases.Within the ruminal microbiota, the enzymes responsible for fermentative H₂ production are mainly distributed among the Clostridia within Firmicutes, with a small proportion occurring in the Bacilli.The electron-bifurcating [FeFe]-hydrogenases in the ruminal microbiota are primarily distributed among the Clostridia, suggesting that Clostridial fermentation is the major source of H₂ in the rumen
		Lachnospiraceae	瘤胃中的Lachnospiraceae编码发酵型氢化酶；Lachnospiraceae的Butyrivibrio编码可催化产氢的Ech氢化酶（energy-converting hydrogenase，Ech）复合体 In the rumen, Lachnospiraceae encode fermentative hydrogenases; and Butyrivibrio, a genus within the Lachnospiraceae, encodes an Ech hydrogenase (energy-converting hydrogenase) complex that catalyses hydrogen production	Lin et al.^[41] Jiao et al.^[43]
		Oscillospiraceae	瘤胃中的Ruminococcus编码可催化产氢的Ech氢化酶复合体；瘤胃Ruminococcus albus为产氢细菌，可编码催化产氢的电子分叉氢化酶HydABC Ruminococcus in the rumen encodes the Ech hydrogenase complex, which catalyses hydrogen production; Ruminococcus albus, a hydrogen-producing ruminal bacterium, encodes the electron-bifurcating hydrogenase HydABC	Jiao et al.^[43] Xie et al.^[44] Ungerfeld^[15]
		Clostridiaceae	瘤胃内Clostridium的丰度与CH₄产量呈显著正相关，此细菌可通过营养物质发酵产氢 Ruminal Clostridium abundance shows a significant positive correlation with CH₄ yield, and this genus is capable of hydrogen production via the fermentation of substrates	Yang et al.^[45]
		Eubacteriaceae	瘤胃中的Eubacterium speies是高效的产氢细菌。Eubacterium_H编码Ech氢化酶复合体。Eubacterium sp.CAG-603通过发酵型[FeFe]-氢化酶催化产氢 In the rumen, Eubacterium species are efficient hydrogen-producing bacteria. Eubacterium_H encodes the Ech hydrogenase complex, and Eubacterium sp. CAG-603 catalyses hydrogen production via a fermentative [FeFe]-hydrogenase	Lan et al.^[32] Jiao et al.^[43] Li et al.^[22]
		Christensenellaceae	瘤胃中的Christensenellaceae R-7编码的氢化酶在瘤胃产氢酶中占比高，推断为瘤胃内重要的产氢微生物 The hydrogenase encoded by Christensenellaceae R-7 constitutes a significant proportion of ruminal hydrogenases, and it is therefore inferred to be a key hydrogen-producing microbe in the rumen	Leahy et al.^[46]
		/	瘤胃内少量存在的Acetogens HGM12587通过Ech氢化酶复合体产氢，此菌株多存在于盲肠 In the rumen, the acetogen HGM12587, which is present at low abundance, produces hydrogen via the Ech hydrogenase complex, and this strain is predominantly found in the caecum	Li et al.^[47]
	Spirochaetes	/	瘤胃中的Spirochaetes编码可催化产氢的发酵型氢化酶；瘤胃中的Treponema bryantii为产氢细菌 In the rumen, Spirochaetes encode fermentative hydrogenases that catalyse hydrogen production; Treponema bryantii is a hydrogen-producing bacterium	Lin et al.^[41] Xie et al.^[44]
	Proteobacteria	Enterobacteriaceae	瘤胃中存在的Klebsiella pneumoniae Y7-3具有高效的产氢能力 Klebsiella pneumoniae Y7-3, found in the rumen, is highly efficient at generating hydrogen	Liu et al.^[48]
	Proteobacteria	Vibrionaceae	瘤胃中的Vibrio cholerae RC9编码A3亚组[FeFe]-氢化酶的基因，具备产氢能力 Ruminal Vibrio cholerae RC9 encodes genes for an A3-subgroup [FeFe]-hydrogenase, thereby conferring hydrogen-producing capability	Lin et al.^[41]
	Synergistetes	Synergistaceae	瘤胃中的Synergistes有50%及以上的基因组编码Ech氢化酶复合体 In the rumen, 50% or more of the genomes of Synergistes encode the Ech hydrogenase complex	Jiao et al.^[43]

Table 1

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Composition of Hydrogen-Producing Microorganisms in the Rumen of Ruminants and Their Regulation on Methanogenesis

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