反刍动物甲烷抑制剂的研究与应用评价

doi:10.3864/j.issn.0578-1752.2026.20.015

Abstract

Abstract:

After carbon dioxide, methane is a greenhouse gas that has the greatest impact on global warming. Methane produced through ruminal fermentation in ruminants not only contributes substantially to greenhouse gas emissions from the livestock sector, but also represents a loss of dietary energy, thereby reducing feed energy utilization efficiency and animal productivity. Therefore, elucidating the regulatory mechanisms underlying ruminal methane production and emission and establishing scientific, standardized mitigation assessment methods applicable to different production systems are of considerable importance for promoting the green and low-carbon development of the livestock industry and achieving the “Dual Carbon” goals. This paper systematically reviewed recent advances in the mechanisms and detection methods associated with methane mitigation in ruminants. First, according to the processes involved in ruminal methanogenesis and their regulatory points, methane inhibitors were classified into feed additives that regulated volatile fatty acid production and reduced hydrogen accumulation, inhibitors targeting methanogens and their key enzymes, and substances that promoted methane oxidation. The methane mitigation effects of 3-nitrooxypropanol (3-NOP), seaweed, nitrate, essential oils, tannins, lipids and combined additives were summarized, with particular emphasis on their effects on feed intake, average daily gain, milk yield and methane emission intensity in ruminants. Secondly, the effects of different physiological stages, dietary compositions, production systems and experimental durations on the assessment of mitigation efficacy were discussed in relation to Latin-square, crossover and continuous experimental designs. The importance of combining short-term trials with long-term monitoring was also emphasized. Thirdly, research approaches for elucidating the mechanisms of action of methane inhibitors were systematically reviewed from the perspectives of microbiology, cellular and molecular biology, ruminal fermentation and ecosystem functioning, animal metabolism, and integrated multidisciplinary research. These mechanisms included the regulation of methanogen communities, inhibition of methyl-coenzyme M reductase and coenzyme M methyltransferase activities, modification of hydrogen metabolism and volatile fatty acid production, alteration of electron acceptor utilization, and promotion of methane oxidation. In addition, this paper compared empirical screening with target-based design strategies for discovering bioactive substances, and introduced technologies including high-throughput screening, isolation and purification, structural identification using mass spectrometry and nuclear magnetic resonance spectroscopy, molecular docking and computational modelling. With regard to methane emission measurements, the principles, application conditions, advantages and limitations of respiration chambers, the sulphur hexafluoride tracer technique, the GreenFeed system, face masks, portable accumulation chambers, house-flux methods, open-path laser detection, unmanned aerial vehicles and satellite remote sensing were reviewed. The selection of an appropriate detection method should be based on the research objectives, animal production system, available equipment and the required measurement accuracy. Finally, the applications of empirical, mechanistic and machine-learning models in predicting methane emissions and mitigation potential were summarized. The need to strengthen the integration of high-quality data, standardize additive dosage and delivery methods, assess both short-term effects and long-term adaptation, and conduct integrated evaluations of mitigation efficacy, animal performance, product safety and life-cycle environmental impacts was highlighted. This review provided a theoretical basis and technical reference for the screening of methane inhibitors in ruminants, elucidation of their mechanisms of action, experimental design, methane emission measurement and prediction of mitigation potential.

Key words: methane, greenhouse gas, ruminants, feed additives, mitigation mechanisms, detection methods, model prediction

JIA Peng, TU Yan, ZHANG NaiFeng, DIAO QiYu. Research Methods and Application Evaluation of Methane Inhibitors for Ruminants[J].Scientia Agricultura Sinica, 2026, 59(20): 4630-4641.

Figures/Tables 4

Table 1

Table 2

Methods to study the mechanism of action of CH4 inhibitors"

研究层面 Research level	基础探索 Basic exploration	深入研究 In-depth research	未来研究方向 Future research direction
微生物学 Microbiology	用添加和不添加H₂的混合瘤胃微生物进行培养，初步了解CH₄抑制剂是否抑制产甲烷菌或产H₂微生物；使用定量PCR和高通量测序探索目标瘤胃微生物的变化；用与CH₄抑制共生的纯培养物验证直接作用 Cultivate mixed rumen microorganisms with and without added H₂ to gain an initial understanding of whether CH₄ inhibitors inhibit methanogens or H₂-producing microorganisms; use quantitative PCR and high-throughput sequencing to explore changes in the target rumen microorganisms; use pure cultures associated with CH₄ inhibition to verify direct effects	不同种类目标微生物纯培养物的剂量反应；研究目标微生物对CH₄抑制剂敏感性的差异；通过高通量测序对目标微生物和其他微生物进行长期（≥84天）体内监测；研究对非目标微生物的直接不良影响 Dose-response of pure cultures of different target microorganisms; investigate differences in the sensitivity of target microorganisms to CH₄ inhibitors; conduct long-term (≥84 days) in vivo monitoring of target and other microorganisms by high-throughput sequencing; investigate direct adverse effects on non- target microorganisms	通过培养组学研究评估CH₄抑制剂对目前不可培养微生物的影响 Assess the effects of CH₄ inhibitors on currently unculturable microorganisms through cultivation omics studies
细胞和分子 Cells and molecules	识别活性化合物 Identify active compounds	确定所涉及的机制类型：酶抑制、膜破坏、阻断DNA和蛋白质合成；通过纯酶-底物体系验证目标酶；展示抑制的具体机制，包括结合位点、反应中间体和机制；确定类型（竞争性或非竞争性）和动力学（即抑制50%酶活性的化合物浓度） Determine the types of mechanisms involved: enzyme inhibition, membrane disruption, and inhibition of DNA and protein synthesis; validate the target enzyme using a pure enzyme-substrate system; demonstrate the specific mechanism of inhibition, including the binding site, reaction intermediates and mechanism; determine the type (competitive or non-competitive) and kinetics (i.e. the concentration of the compound that inhibits 50% of enzyme activity)	研究目标微生物敏感性差异与获得耐药性的作用机制 Investigate the mechanisms underlying differences in target microorganism sensitivity and the acquisition of resistance
微生物发酵与生态系统 Microbial fermentation and ecosystems	调查总VFA浓度和单个VFA摩尔百分比的变化；测定体外H₂积累量和体内H₂排出量；当活性化合物或其代谢物并非天然存在于瘤胃中，或当降解途径已被明确且研究重点在于参数化速率时，分析化学方法可能足以用于研究降解速率；如果活性化合物或其代谢产物可能存在于瘤胃中，则进行体外放射性标记试验以了解活性化合物的瘤胃代谢 Investigate changes in total VFA concentration and the molar percentage of individual VFAs; measure in vitro H₂ accumulation and in vivo H₂ emissions; when the active compound or its metabolites are not naturally present in the rumen, or when the degradation pathway has been established and the research focus is on rate parameterization, analytical chemistry methods may be sufficient to study the degradation rate; if the active compound or its metabolites may be present in the rumen, conduct in vitro radiolabeling experiments to understand the ruminal metabolism of the active compound	测定溶解H₂、甲酸、琥珀酸、乳酸、甲醇和乙醇的浓度；进行宏基因组学和宏转录组学，特别关注氢化酶 Measure the concentrations of dissolved H₂, formate, succinate, lactate, methanol and ethanol; conduct metagenomics and metatranscriptomics, with particular attention to hydrogenases	通过代谢组学分析全面解析CH₄抑制剂对电子汇和代谢中间体的影响；宏蛋白质组学分析有助于更深入地了解代谢途径速率的变化；确定抑制CH₄生成对VFA产生的影响；分离与鉴定代谢活性化合物的微生物 Comprehensively analysis the effects of CH₄ inhibitors on electron sinks and metabolic intermediates through metabolomic analysis; metaproteomic analysis will help to gain a deeper understanding of changes in the rates of metabolic pathways; determine the effects of inhibiting CH₄ production on VFA production; isolate and identify microorganisms that metabolize active compounds
动物 Animals	了解活性化合物及其瘤胃代谢物在动物产品（奶类或肉类）和组织（瘤胃、肝脏、乳腺、血液和其他器官）中的吸收、代谢、排泄和沉积；活性化合物及其中间体和终产物代谢物的毒理学与致突变性检测；如果采食量下降，需检查是否存在适口性问题或炎症问题 Understand the absorption, metabolism, excretion and deposition of active compounds and their ruminal metabolites in animal products (milk or meat) and tissues (rumen, liver, mammary glands, blood and other organs); conduct toxicological and mutagenicity testing of active compounds and their intermediate and end-product metabolites; if feed intake decreases, check for palatability or inflammatory problems	对反刍动物和非反刍动物口服同位素标记化合物，然后在组织、粪便、尿液、奶样和呼气中进行定量检测；对瘤胃液和血浆进行代谢组学分析，以筛选潜在有毒化合物或可能影响采食量的化合物 Administer isotope-labelled compounds orally to ruminants and non-ruminants, followed by quantitative analysis in tissues, faeces, urine, milk samples and exhaled breath; conduct metabolomic analysis of rumen fluid and plasma to screen for potentially toxic compounds or compounds that may affect feed intake	确定对VFA吸收、糖异生和其他吸收后过程的影响；研究通过抑制瘤胃CH₄产生进而降低采食量的机制 Determine the effects on VFA absorption, gluconeogenesis and other post- absorptive processes; investigate the mechanisms by which inhibiting rumen CH₄ production subsequently reduces feed intake
交叉层面 Cross-cutting level	科研论文中方法涉及的动物物种、品种、生产类型、年龄、生产阶段和水平、饮食、饲养管理和行为、采食模式及其他信息的系统阐述 Systematic description in research papers of the animal species, breeds, production types, ages, production stages and levels, diets, husbandry management and behavior, feeding patterns and other information involved in the methods	描述活性化合物在瘤胃中浓度的每日动态变化；优化具有协同作用模式的各种CH₄抑制剂组合的设计 Describe the daily dynamics of active compound concentrations in the rumen; optimize the design of various combinations of CH₄ inhibitors with synergistic modes of action	在体内水平以及在混合和定向培养的对照试验中，研究不同添加剂的功效如何受到生理和微生物变量的影响 Investigate how the efficacy of different additives is affected by physiological and microbial variables in vivo and in controlled trials involving mixed and targeted cultures

Table 2

Table 3

Research methods of exploring bioactive substances that inhibit ruminal methanogenic"

研究方法 Research methods	优越性 Superiority	局限性 Limitation
经验法 Empirical approach	允许在短时间内获得和检验较多数量且多样化的候选物 Allows a large number and variety of candidates to be obtained and tested within a short period	母料中存在的CH₄抑制剂候选物是复杂化合物混合物的一部分，并且含有量可能较低，因此提取和检测所需的母料数量可能相当大且难以获取。尽管可以直接使用未经分离或纯化的母体材料，但其中其他化合物的存在可能会显著影响试验结果，并引入额外的干扰因素，这些因素会阻碍对CH₄抑制剂混合产物的有效性认定。因此，建议使用专用设备和适当的分馏和纯化方法进行高效加工 The candidate CH₄ inhibitors present in the feedstock are part of a mixture of complex compounds and may be present in low quantities; consequently, the quantity of feedstock required for extraction and detection may be considerable and difficult to obtain. Although the parent material can be used directly without separation or purification, the presence of other compounds may significantly affect the test results and introduce additional interfering factors that hinder the effective determination of the efficacy of the CH₄ inhibitor mixed product. Therefore, it is recommended that specialized equipment and appropriate fractionation and purification methods be used for efficient processing
机理法 Mechanistic approach	能够实现无限搜索，以相对快速、高通量的方式发现、分析和开发产甲烷菌的特异性抑制剂，并有望找到新型化合物 Enables unlimited screening to discover, analyse and develop specific inhibitors of methanogens in a relatively rapid, high-throughput manner, and is expected to identify novel compounds	需要精密的设备和特定的专业知识。潜在的CH₄抑制剂候选物可能面临以下挑战：天然来源中无法获取或尚未被发现；即使已知，仍需从母体材料中分离；或必须通过合成获得，部分化合物难以足量合成或成本高昂。此外，通过该途径筛选出的部分CH₄抑制剂候选物可能在细胞内生物利用率较低，且转运蛋白突变可能导致细胞对该候选物产生耐药性 Requires precise equipment and specific expertise. Potential CH₄ inhibitor candidates may face the following challenges: they may be unobtainable or undiscovered in natural sources; even when known, they may still need to be isolated from the parent material; or they may have to be obtained through synthesis, with some compounds being difficult or costly to synthesize in sufficient quantities. In addition, some CH₄ inhibitor candidates screened through this route may have low intracellular bioavailability, and mutations in transport proteins may result in cellular resistance to the candidate

Table 3

Table 4

Detection methods for CH4 emissions from ruminants and their application characteristics"

方法 Methods	优点 Advantages	缺点 Disadvantages
便携式蓄积室 Portable accumulation chamber，PAC	方法简单。对于绵羊，可以在拖车上放置10到12个PAC单位，以便在农场进行测量。每天可以测量到相对大量的动物个体（70—84只）。在一次试验中可设置多个处理 The method is simple. For sheep, 10 to 12 PAC units can be placed on a trailer to enable measurements to be conducted on the farm. A relatively large number of individual animals (70 to 84) can be measured each day. Multiple treatments can be established in a single trial	需要将动物从其生产环境中移出以进行测量，并且在测量过程中通常不喂食。无法测量采食量。劳动密集型。不覆盖整个喂养周期的抽样；需要密集的现场采样计划来提供准确的CH₄排放量测量。目前的设置和测量方法主要是对动物的排放进行排名，适宜于动物选育 Animals need to be removed from their production environment for measurement and are usually not fed during the measurement process. Feed intake cannot be measured. Labor-intensive. Sampling does not cover the entire feeding period; an intensive field sampling plan is required to provide accurate measurements of CH₄ emissions. The current setup and measurement methods mainly rank emissions from animals and are suitable for animal breeding
呼吸面罩 Face mask，FM	方法相对简单，可以获得准确的数据。该方法有移动版本 The method is relatively simple and can provide accurate data. The method has a mobile version	需要一个密集的现场采样计划以获得准确的CH₄产量测量。动物在测量过程中不能进食或饮水，需要动物配合才能进行测量。劳动密集型。可以同时测量的动物数量有限 An intensive field sampling plan is required to obtain accurate measurements of CH₄ production. Animals cannot eat or drink during the measurement process, and their co-operation is required for measurements to be conducted. Labour-intensive. The number of animals that can be measured simultaneously is limited
房屋通量 House flux	在散栏式畜舍中进行群体测量。能够测量包括粪便释放的气体排放量。对养殖场正常饲养管理干扰很小或没有干扰 Group measurements are conducted in loose- housing facilities. Gas emissions, including those released from faeces, can be measured. There is little or no interference with normal farm husbandry management	测量的通风率存在较大的不确定性。在自然通风的畜舍里很难获得准确的测量值。漏缝地板型畜舍的粪便量会对结果产生较大影响。捕获肠道CH₄排放量微小差异（5%—10%）的灵敏度较差。如果用示踪剂（sulfur hexafluoride，SF₆），SF₆是一种强效GHG。不建议用于测试CH₄抑制剂 There is considerable uncertainty in the measured ventilation rates. It is difficult to obtain accurate measurements in naturally ventilated housing. The quantity of faeces in slatted-floor housing can have a considerable effect on the results. It has poor sensitivity for detecting small differences (5% to 10%) in enteric CH₄ emissions. If a tracer is used (sulfur hexafluoride, SF₆), SF₆ is a potent GHG. It is not recommended for testing CH₄ inhibitors
结合模型确定流量的方法 Methods to determine fluxes combined with models	能够测量大群动物或整个养殖场（或地区），且对动物无干扰性。可在圈养条件下进行长期测量。可测量圈养环境下粪便和放牧条件下土壤的气体排放量，包括N₂O。对于CH₄主要来自反刍动物的地区，大规模设置（区域塔）有可能被用作自上而下的方法 Can measure large groups of animals or an entire farm (or region) without disturbing the animals. Long-term measurements can be conducted under housed conditions. Gas emissions from faeces under housed conditions and from soil under grazing conditions, including N₂O, can be measured. In regions where CH₄ comes mainly from ruminants, large-scale installations (regional towers) could potentially be used as a top-down approach	不适用于室内。测量位置需要有相对平坦的地形，有最少的障碍物（如建筑物、树木）和其他产生CH₄的来源（在测量动物群体时）。分析中包含的测量仅在适当的天气条件下进行（数据丢失率很高）。在大多数情况下没有统计复制。捕获肠道CH₄排放量微小差异（5%—10%）的灵敏度较差。不建议用于测试CH₄抑制剂。如果用示踪剂（nitrous oxide，N₂O），N₂O是一种强效GHG It is not suitable for indoor use. The measurement location needs to have relatively flat terrain, with minimal obstacles (such as buildings and trees) and other sources of CH₄ (when measuring a group of animals). Measurements included in the analysis are only conducted under suitable weather conditions (the data loss rate is very high). In most cases, there is no statistical replication. It has poor sensitivity for detecting small differences (5% to 10%) in enteric CH₄ emissions. It is not recommended for testing CH₄ inhibitors. If a tracer is used (nitrous oxide, N₂O), N₂O is a potent GHG
无人机载传感器 Unmanned aerial vehicle-mounted sensors	不干扰养殖场的正常饲养管理。获得整个养殖场的CH₄排放量 Does not interfere with normal farm husbandry management. Obtains the CH₄ emissions of the entire farm	短期和不频繁的测量活动无法提供动物排放量的代表性样本。很难对养殖场的烟羽排放进行具有代表性的捕获。在测量活动中，如果依赖区域气象站的数据，其变异性极高；而依靠农场气象站的数据进行测量，其变异性也很大（误差范围有时甚至包括零排放）。难以准确估算气流。无法区分农场的不同排放源（例如，肠道、粪便储存、幼龄动物、成年动物）。无法在受控条件下比较处理。通常需要养殖场数据（例如动物类别、动物数量、粪便的储存量、粪便储存系统等）才能理解这些数据。不建议用于测试CH₄抑制剂 Short-term and infrequent measurement activities cannot provide representative samples of animal emissions. It is difficult to capture representative measurements of the farm plume emissions. During measurement activities, if data from a regional weather station are relied upon, their variability is extremely high; while measurements relying on data from a farm weather station also have considerable variability (the error range sometimes even includes zero emissions). It is difficult to estimate airflow accurately. It is not possible to distinguish between different emission sources on the farm (e.g. enteric, faecal storage, young animals and adult animals). Treatments cannot be compared under controlled conditions. Farm data are usually required (e.g. animal category, number of animals, quantity of faeces stored, faecal storage system, etc.) to understand these data. It is not recommended for testing CH₄ inhibitors
卫星 Satellite	测量一个地区长时间序列（季节性和跨年）内的CH₄浓度。有可能确定CH₄浓度高的热点区域。有可能作为一种自上而下的方法来跟踪CH₄浓度热点是否随时间变化 Measures CH₄ concentrations in an area over a long time series (seasonal and interannual). It may be possible to identify hotspot areas with high CH₄ concentrations. It may be possible to use it as a top-down approach to track whether CH₄ concentration hotspots change over time	方法尚未完全开发。目前仅在阳光充足且无云的条件下进行粗分辨率的CH₄浓度测量。现阶段，尚不清楚卫星是否可以定位排放源并跟踪动物排放。无法区分一个地区不同的CH₄排放源（如动物排放、土壤排放、自然陆地排放、垃圾填埋场CH₄、化石燃料CH₄），也无法区分单个养殖场（可能是孤立的大型养殖场）。仅适用于以畜牧业为主的地区。只能验证一个地区由于实施了所有缓解措施而导致的总体CH₄减排情况，而不能针对特定的缓解措施（如单个农场使用CH₄抑制剂）进行验证。不建议用于测试CH₄抑制剂 The method has not yet been fully developed. At present, only coarse-resolution measurements of CH₄ concentrations are conducted under sunny and cloud-free conditions. At this stage, it is not yet clear whether satellites can locate emission sources and track animal emissions. It is not possible to distinguish between different CH₄ emission sources in an area (such as animal emissions, soil emissions, natural terrestrial emissions, landfill CH₄ and fossil-fuel CH₄), nor between individual farms (which may be isolated large farms). It is only suitable for regions dominated by livestock farming. It can only verify the overall reduction in CH₄ emissions in an area resulting from the implementation of all mitigation measures, and cannot verify specific mitigation measures (such as the use of CH₄ inhibitors on an individual farm). It is not recommended for testing CH₄ inhibitors

Table 4

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CH₄抑制剂 CH₄ inhibitor	添加剂量 Supplementation dosage	动物 Animal	平均CH₄产量 Average CH₄ production (g·d^-1)	CH₄排放量变化 Changes in CH₄ emissions				其他指标变化 Other indicator changes
CH₄抑制剂 CH₄ inhibitor	添加剂量 Supplementation dosage	动物 Animal	平均CH₄产量 Average CH₄ production (g·d^-1)	CH₄产量 CH₄ production (g·d^-1)	CH₄产率 CH₄ yield (g·kg^-1 DMI)	CH₄排放强度 CH₄ emission intensity (g·kg^-1 ECM or FPCM)	CH₄排放因子 CH₄ emission factor (% of GE)	干物质采食量 Dry matter intake (kg·d^-1)	日增重 Daily liveweight gain (g·d^-1)	料重比 Feed conversion ratio	产奶量 Milk yield (kg·d^-1)
3-NOP^[26]	70.5 mg·kg^-1 DM	奶牛 Dairy cattle	361	-32.7±1.5	-30.9±1.5	-30.9±1.5	—	—	—	—	—
海藻^[27] Seaweed	—	奶牛和肉牛 Dairy and beef cattle	—	—	-5.3±3.5	—	—	0.28 (-0.63 to 0.07; 95% CI)	-0.01 (-0.05 to 0.03; 95% CI)	-0.38 (-0.58 to -0.18; 95% CI)	1.35 (0.91 to 1.78; 95% CI)
硝酸盐^[28] Nitrate	16.6 g·kg^-1 DM	奶牛 Dairy cattle	339	-16.9±1.0	-15.5±1.2	—	—	—	—	—	—
硝酸盐^[28] Nitrate	16.8 g·kg^-1 DM	肉牛 Beef cattle	155	-12.2±1.3	-9.0±1.8	—	—	—	—	—	—
复合精油^[29] Blended essential oils	1 g·d^-1	奶牛 Dairy cow	321	0.954 (0.921 to 0.987; 95% CI)	0.982 (0.918 to 1.050; 95% CI)	0.925 (0.864 to 0.989; 95% CI)	—	1.003 (0.985 to 1.020; 95% CI)			1.020 (1.011 to 1.028; 95% CI)

Research Methods and Application Evaluation of Methane Inhibitors for Ruminants

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