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

中国农业科学 ›› 2026, Vol. 59 ›› Issue (20): 4642-4654.doi: 10.3864/j.issn.0578-1752.2026.20.016

• 专题：农食系统低碳转型与反刍动物甲烷减排的理论、技术与实践 • 上一篇下一篇

反刍动物瘤胃产氢微生物构成及其对甲烷生成的调控

贾鹏¹(), 夏东¹, 苏衍菁², 姜雪元¹, 韦子海², 雷胡龙¹, 康金国³, 焦崇¹(), 陆乃升¹()

¹ 上海市农业科学院畜牧兽医研究所，上海 201106
² 光明牧业有限公司，农业农村部南方奶牛遗传改良重点实验室，上海 200436
³ 上海市浦东新区农业广播电视学校，上海 201299

收稿日期:2025-08-06 接受日期:2025-10-30 出版日期:2026-10-16 发布日期:2026-10-21
通信作者:
焦崇，E-mail：jiaochong2017@163.com。
陆乃升，E-mail：lunaisheng@saas.sh.cn
联系方式: 贾鹏，E-mail：jiazhenpeng123123@163.com。
基金资助:
上海市农业科技创新项目(K2024-02-08-00-12-F00023); 上海市自然科学基金(24ZR1462000); 上海市农业科学院新进博士2025-2026年度揭榜挂帅项目(BS25017)

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

JIA Peng¹(), XIA Dong¹, SU YanJing², JIANG XueYuan¹, WEI ZiHai², LEI HuLong¹, KANG JinGuo³, JIAO Chong¹(), LU NaiSheng¹()

¹ Institute of Animal Science and Veterinary Medicine, Shanghai Academy of Agricultural Sciences, Shanghai 201106
² Bright Farming Co. Ltd./ Key Laboratory of Dairy Cattles Genetic Improvement in Southern China, Ministry of Agriculture and Rural Affairs, Shanghai 200436
³ Shanghai Pudong New Area Livestock and Aquaculture Products Technology Promotion Center, Shanghai 201299

Received:2025-08-06 Accepted:2025-10-30 Published:2026-10-16 Online:2026-10-21

摘要/Abstract

摘要：

甲烷（CH₄）为全球第二大温室气体，因其短期增温效应强，对全球气候快速变暖具有显著贡献。近年来，全球大气CH₄浓度持续攀升，2024年较工业革命前增长了1.6倍。CH₄排放导致的全球变暖，威胁了全人类的生存环境与身体健康，每吨CH₄排放的社会成本高达数千美元。反刍动物养殖是农业领域主要的人为CH₄排放源之一，其CH₄减排至关重要。反刍动物排放的CH₄主要由瘤胃微生物通过氢营养途径、甲基营养途径和乙酸裂解途径生成，其中利用氢气（H₂）还原二氧化碳（CO₂）生成CH₄的氢营养途径占主导地位。所以H₂是瘤胃产CH₄过程至关重要的前体物质，合理地减少或消耗瘤胃微生物发酵过程中的H₂产量，是最直接的CH₄减排策略。H₂由各类产氢微生物（原虫、细菌、真菌）生成，其中原虫以全毛类为主，细菌以Firmicutes门的Clostridia纲为优势菌群，部分真菌也可产氢。原虫、细菌、真菌通过瘤胃碳水化合物发酵生成挥发性脂肪酸的过程生成H₂，然而其产氢能力存在差异。具体而言，原虫的产氢能力与体长、体内氢酶体的数量等因素有关，其中全毛类原虫和大型原虫为高产氢菌群；细菌则与其介导的产氢途径有关，其中可产生乙酸及H₂的细菌产氢能力较高；真菌报道较少。上述高产氢微生物可作为减少H₂向产CH₄途径传递，促进CH₄减排的关键靶点。尽管产氢微生物的产氢能力存在差异，但多需体内氢化酶参与产氢途径，产氢微生物体内的氢化酶能利用瘤胃发酵过程中产生的还原型烟酰胺腺嘌呤二核苷酸（NADH）和还原型铁氧还蛋白（Fd_red），还原H⁺生成H₂。此外，产氢微生物的产氢途径受微生物群落组成、饲粮营养结构、功能性添加剂的影响，可分别通过调控瘤胃产氢发酵类型、氢分压及氢化酶的表达、产氢微生物的活性影响产氢效率。可基于此产氢特征，减少产氢微生物的H₂产量，促进CH₄减排：可优化瘤胃产氢微生物的群落组成，促进低产氢微生物主导瘤胃发酵，如饲喂枯草芽孢杆菌；也可调控产氢微生物体内氢化酶的表达，如通过饲粮营养调控实现；还可直接抑制产氢微生物的活性，如在饲粮中添加单宁、中药复方等。在维持瘤胃正常氢分压的基础上，减少用以产CH₄的H₂产量，是有效的CH₄减排策略。文章就瘤胃内产氢微生物的种类及产氢特征进行综述，为从源头上制定CH₄减排措施提供理论依据。

关键词: 反刍动物, 甲烷, 瘤胃, 产氢微生物, 甲烷减排

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

贾鹏, 夏东, 苏衍菁, 姜雪元, 韦子海, 雷胡龙, 康金国, 焦崇, 陆乃升. 反刍动物瘤胃产氢微生物构成及其对甲烷生成的调控[J]. 中国农业科学, 2026, 59(20): 4642-4654.

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.

图/表 2

图1

表1

反刍动物瘤胃内的产氢细菌"

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]

表1

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反刍动物瘤胃产氢微生物构成及其对甲烷生成的调控

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

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