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Article(id=1203053368513094598, tenantId=1146029695717560320, journalId=1189873630562394117, issueId=1203053366290113441, articleNumber=null, orderNo=null, doi=10.11855/j.issn.0577-7402.2023.03.0339, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1632931200000, receivedDateStr=2021-09-30, revisedDate=null, revisedDateStr=null, acceptedDate=1637251200000, acceptedDateStr=2021-11-19, onlineDate=1764759874704, onlineDateStr=2025-12-03, pubDate=1679932800000, pubDateStr=2023-03-28, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1764759874704, onlineIssueDateStr=2025-12-03, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1764759874704, creator=13701087609, updateTime=1764759874704, updator=13701087609, issue=Issue{id=1203053366290113441, tenantId=1146029695717560320, journalId=1189873630562394117, year='2023', volume='48', issue='3', pageStart='245', pageEnd='366', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=0, createTime=1764759874174, creator=13701087609, updateTime=1764810242575, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1203264626747220064, tenantId=1146029695717560320, journalId=1189873630562394117, issueId=1203053366290113441, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1203264626747220065, tenantId=1146029695717560320, journalId=1189873630562394117, issueId=1203053366290113441, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=339, endPage=344, ext={EN=ArticleExt(id=1203053368940913619, articleId=1203053368513094598, tenantId=1146029695717560320, journalId=1189873630562394117, language=EN, title=Recent advance in relation between trimethylamine N-oxide metabolized by intestinal microbes and cognitive impairment, columnId=1190243275882729994, journalTitle=Medical Journal of Chinese People’s Liberation Army, columnName=Review, runingTitle=null, highlight=null, articleAbstract=

Cognition is a process in which the human brain receives external information, processes it, and transforms it into internal psychological activities to obtain or apply knowledge. Cognitive impairment is the impairment of one or more functions in memory, language, visual space, execution, calculation, and understanding, affecting an individual's daily or social ability. With the aging of the population, the incidence rate of cognitive impairment is increasing. Currently, biomarkers related to the cognitive impairment have become a research focus. Gut microbiota has emerged as a potential player in pathophysiology of cognitive impairment. Trimethylamine N-oxide (TMAO), the metabolite produced by gut microbiota, has mechanistic relevance to cognitive impairment. Therefore, the main goal of the present review is to provide the reader with potential mechanisms of TMAO and cognitive impairment. Although a link between TMAO and cognitive impairment is far from definitive, this review will serve as a call for research into this new area.

, correspAuthors=Mei-Juan Gu, authorNote=null, correspAuthorsNote=
* E-mail:
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认知是人脑接受外界信息，经过加工处理转换成内在心理活动，从而获取或应用知识的过程。认知功能障碍是指记忆、语言、视空间、执行、计算和理解力等方面中的一项或多项功能受损，影响个体的日常或社会能力。随着我国人口老龄化程度的加剧，认知功能障碍的发病率逐渐增高，目前与认知功能障碍相关的生物学标志物已成为研究热点。近年来研究发现，肠道菌群紊乱与认知功能障碍关系密切。肠道微生物代谢产物氧化三甲胺（TMAO）在认知功能障碍疾病的发生发展中起着重要作用。本文就TMAO与认知功能障碍疾病关系的研究进展进行综述，以期为今后该领域的研究提供线索。

, correspAuthors=顾美娟, authorNote=null, correspAuthorsNote=
顾美娟，E-mail：
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周传彬，硕士研究生，主要从事神经系统退行性疾病的基础与临床研究

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周传彬，硕士研究生，主要从事神经系统退行性疾病的基础与临床研究

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周传彬，硕士研究生，主要从事神经系统退行性疾病的基础与临床研究

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Related therapeutics to improve cognition by modulating gut microbiota

, figureFileSmall=null, figureFileBig=null, tableContent=
治疗方法措施
调整饮食结构均衡膳食纤维、减少红肉摄入量[46]
菌群移植粪便微生物菌群移植[54]
益生菌双歧杆菌、乳杆菌、肠球菌、酪酸杆菌、复合制剂[49-50]
益生元菊粉、葡聚糖、低聚果糖、低聚半乳糖、乳果糖[55]
中药白藜芦醇[47]、小檗碱[48]、补肾天水方[52]、针灸[53]
海藻提取物GV-971(甘露特钠胶囊)[51]
), ArticleFig(id=1203053372531236971, tenantId=1146029695717560320, journalId=1189873630562394117, articleId=1203053368513094598, language=CN, label=表1, caption=

通过调节肠道菌群改善认知的相关治疗方法

, figureFileSmall=null, figureFileBig=null, tableContent=
治疗方法措施
调整饮食结构均衡膳食纤维、减少红肉摄入量[46]
菌群移植粪便微生物菌群移植[54]
益生菌双歧杆菌、乳杆菌、肠球菌、酪酸杆菌、复合制剂[49-50]
益生元菊粉、葡聚糖、低聚果糖、低聚半乳糖、乳果糖[55]
中药白藜芦醇[47]、小檗碱[48]、补肾天水方[52]、针灸[53]
海藻提取物GV-971(甘露特钠胶囊)[51]
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肠道菌群代谢产物氧化三甲胺与认知功能障碍疾病关系的研究进展
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周传彬 , 顾美娟 *
解放军医学杂志 | 综述 2023,48(3): 339-344
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解放军医学杂志 | 综述 2023, 48(3): 339-344
肠道菌群代谢产物氧化三甲胺与认知功能障碍疾病关系的研究进展
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周传彬, 顾美娟*
作者信息
  • 昆明医科大学第一附属医院老年神经内科，云南昆明 650000

    • 周传彬，硕士研究生，主要从事神经系统退行性疾病的基础与临床研究

通讯作者:

顾美娟，E-mail：
Recent advance in relation between trimethylamine N-oxide metabolized by intestinal microbes and cognitive impairment
Chuan-Bin Zhou, Mei-Juan Gu*
Affiliations
  • Department of Geriatric Neurology, the First Affiliated Hospital of Kunming Medical University, Kunming, Yunnan 650000, China
出版时间: 2023-03-28 doi: 10.11855/j.issn.0577-7402.2023.03.0339
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摘要
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认知是人脑接受外界信息，经过加工处理转换成内在心理活动，从而获取或应用知识的过程。认知功能障碍是指记忆、语言、视空间、执行、计算和理解力等方面中的一项或多项功能受损，影响个体的日常或社会能力。随着我国人口老龄化程度的加剧，认知功能障碍的发病率逐渐增高，目前与认知功能障碍相关的生物学标志物已成为研究热点。近年来研究发现，肠道菌群紊乱与认知功能障碍关系密切。肠道微生物代谢产物氧化三甲胺（TMAO）在认知功能障碍疾病的发生发展中起着重要作用。本文就TMAO与认知功能障碍疾病关系的研究进展进行综述，以期为今后该领域的研究提供线索。

肠道微生物  /  氧化三甲胺  /  认知功能障碍  /  痴呆

Cognition is a process in which the human brain receives external information, processes it, and transforms it into internal psychological activities to obtain or apply knowledge. Cognitive impairment is the impairment of one or more functions in memory, language, visual space, execution, calculation, and understanding, affecting an individual's daily or social ability. With the aging of the population, the incidence rate of cognitive impairment is increasing. Currently, biomarkers related to the cognitive impairment have become a research focus. Gut microbiota has emerged as a potential player in pathophysiology of cognitive impairment. Trimethylamine N-oxide (TMAO), the metabolite produced by gut microbiota, has mechanistic relevance to cognitive impairment. Therefore, the main goal of the present review is to provide the reader with potential mechanisms of TMAO and cognitive impairment. Although a link between TMAO and cognitive impairment is far from definitive, this review will serve as a call for research into this new area.

gut microbiota  /  trimethylamine N-oxide  /  cognitive impairment  /  dementia
周传彬, 顾美娟. 肠道菌群代谢产物氧化三甲胺与认知功能障碍疾病关系的研究进展. 解放军医学杂志, 2023 , 48 (3) : 339 -344 . DOI: 10.11855/j.issn.0577-7402.2023.03.0339
Chuan-Bin Zhou, Mei-Juan Gu. Recent advance in relation between trimethylamine N-oxide metabolized by intestinal microbes and cognitive impairment[J]. Medical Journal of Chinese People’s Liberation Army, 2023 , 48 (3) : 339 -344 . DOI: 10.11855/j.issn.0577-7402.2023.03.0339
正文
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认知功能障碍是指记忆、语言、视空间、执行、计算和理解判断等认知功能中的一项或多项受损[1]，当上述认知领域有2项及以上受累并影响个体的日常或社会能力时，可考虑为痴呆。痴呆包括变性病性痴呆和非变性病性痴呆两种。随着我国人口老龄化程度的加剧，认知功能障碍患者的数量逐渐增多，给患者及照料者造成了沉重的生活和经济负担。科学有效的预防和治疗方法对认知功能障碍疾病至关重要，除了传统的危险因素外，目前肠道菌群及其代谢产物已成为认知功能障碍疾病的研究热点。随着高通量测序技术的发展，多项研究表明，微生物群是人类健康和疾病的关键因素，其中研究最多的是分布于胃肠道的微生物菌群。肠道微生物群包括6个主要门类：拟杆菌门、厚壁菌门、变形菌门、放线菌门、疣微菌门和梭杆菌门，其中以拟杆菌门和厚壁菌门为主[2]。“微生物-肠-脑轴”的发现揭示了肠道中的微生物群及其代谢产物可通过迷走神经介导中枢神经系统发生变化[3]。故当宿主与微生物群的相互作用失衡时，微生物群可促进中枢神经系统疾病的发生和发展。来自日常饮食中的磷脂酰胆碱和左旋肉碱进入肠道后在肠道菌群的作用下可代谢为三甲胺（trimethylamine，TMA），后者可在肠道被吸收入血，然后在肝脏黄素单加氧酶3（flavin-containing monooxygenase 3，FMO3）的作用下氧化为氧化三甲胺（trimethylamine-N-oxide，TMAO）。研究发现，TMAO与神经系统疾病、心血管疾病、内分泌疾病及肾脏疾病等多种慢性疾病相关[4]。血浆中TMAO的浓度与FMO3的活性、饮食结构、年龄、性别、肠道菌群及肾功能等因素有关[5]。本文就肠道菌群代谢产物TMAO与认知功能障碍疾病关系的研究进展进行综述。
1 TMAO与认知功能障碍疾病的关系
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1.1 与阿尔茨海默病(Alzheimer's disease，AD)的关系
AD是一种中枢神经系统退行性疾病，是老年期认知功能障碍最常见的原因之一。AD的典型病理特征是β淀粉样物质在细胞外沉积形成神经炎性斑，过度磷酸化的Tau蛋白在神经细胞内聚集形成神经原纤维缠结，神经元缺失，以及胶质细胞增生[6-7]。
2016年Xu等[8]采用集成计算方法证实了AD与肠道微生物代谢产物之间的相关性，其中TMAO与AD密切相关。有研究发现，TMAO可穿过并破坏血脑屏障[9-11]；2017年Del Rio等[12]研究发现，TMAO存在于AD患者的脑脊液中；另有研究发现，FMO1和FMO3可在人脑中表达[13]。因此，脑脊液中的TMAO是由中枢神经系统自身合成，还是来源于外周血中的TMAO通过血脑屏障进入脑脊液，目前仍需进一步探索。Vogt等[14]评估了轻度认知功能障碍患者、AD患者和认知正常者的脑脊液TMAO浓度，结果显示，与认知正常者相比，AD患者和轻度认知功能障碍患者的脑脊液TMAO浓度较高，但AD患者与轻度认知功能障碍患者比较无明显差异；脑脊液TMAO浓度与AD生物标志物t-Tau、神经丝轻链（neurofilament light chain，NFL）、p-Tau及p-Tau/Aβ42呈正相关。由此推测，TMAO可能是AD发生病理变化的促进因素。
已经证实，TMAO能够导致小鼠大脑老化、神经退化和认知功能障碍[15]。Wang等[16]研究APP/PS1双转基因AD模型小鼠发现，喂食胆碱的小鼠血浆TMAO浓度升高、学习及记忆能力下降；ROC曲线及皮尔森相关分析显示，血浆TMAO浓度与小鼠认知功能障碍相关。通过肠道菌群重塑可抑制肠道微生物代谢产物TMA的分泌，降低TMAO浓度，从而阻止认知功能障碍的恶化[15-16]。此外，AD小鼠给予美金刚治疗并添加植物乳杆菌可抑制TMAO的合成，降低Aβ42的浓度，同时降低炎性因子白细胞介素-2（interleukin-2，IL-2）、IL-17、α干扰素（interferon-α，INF-α）水平，保护海马神经元，减轻神经炎症；与单独应用美金刚相比，添加植物乳杆菌缓解患者认知功能障碍的效果更显著。Li等[17]发现，TMAO可诱导海马CA3区神经元衰老，破坏海马CA1区的超微结构，其具体表现形式和作用机制如下：（1）TMAO组的突触囊泡更少，突触后区域减小，电子致密物质密度更低，突触裂隙更广，线粒体和内质网肿胀；（2）小鼠海马氧化应激水平增高，海马中过氧化氢（H2O2）浓度较高、总超氧化物歧化酶（T-SOD）活性较低；（3）通过干扰线粒体能量代谢并促进自由基产物的释放诱导神经元衰老；（4）减少突触可塑性相关蛋白突触素（synaptophysin，SYN）、突触后密度蛋白-95（postsynaptic density protein-95，PSD-95）和N-甲基-D-天门冬氨酸受体1（N-methyl-D-aspartate receptor 1，NMDAR1）的表达；（5）下调mTOR信号通路的活性。另有研究发现，TMAO也可能在Aβ聚集中起作用[18]，从而促进AD的发生和发展。总之，TMAO在AD发展中发挥着重要作用。
1.2 与脑卒中后认知功能障碍(post-stroke cognitive impairment，PSCI)的关系
脑卒中在我国发病率很高，卒中是导致认知功能障碍的常见原因之一，中老年卒中患者PSCI发病率为70%，严重影响患者的生活质量，增加了社会负担[19]。此外，PSCI与卒中发生的解剖部位、面积等因素有关[20]。
近年来国内外研究发现，TMAO可促进动脉硬化，在脑血管病的发生发展中起着重要作用。具体机制总结如下：（1）给予TMAO饮食的小鼠，其肝脏合成胆固醇7α-羟化酶和胆汁酸的能力降低，同时胆汁酸中牛磺酰脱氧胆酸（taurochenodeoxycholic acid，TCDCA）及牛磺酸脱氧胆酸（taurohyodeoxycholic acid，THDCA）的浓度升高，后两者可通过上调法尼醇X受体（farnesoid X receptor，FXR）的表达诱导小二聚体伴侣的表达，从而抑制肝脏胆固醇7α-羟化酶的合成，进而抑制胆汁酸的合成[21]。（2）TMAO可通过上调巨噬细胞CD36的表达参与动脉粥样硬化的发生发展[22]。此外，TMAO可显著上调巨噬细胞表面清道夫受体的数量，促进泡沫细胞的形成[23]。（3）TMAO可抑制胆固醇的逆向转运，影响脂蛋白及胆固醇的代谢，并改变肝脏、肠道、动脉壁等多种脏器的胆固醇代谢通路，促进动脉粥样斑块的形成[24]。（4）血管内皮细胞具有生物合成、降解血管活性介质、抗凝和纤溶等多种生物功能，因此，血管内皮细胞功能失调是动脉粥样硬化的重要因素之一。研究发现，TMAO可促使炎性因子释放，激活信号转导途径，诱导血管内皮细胞发生相关的炎症反应，从而导致血管内皮细胞损伤[25-27]。总之，TMAO可能通过上调动脉粥样硬化相关巨噬细胞受体的表达，干扰胆固醇的逆向转运及抑制胆汁酸的合成，参与血管内皮细胞损伤及炎症反应，从而促进动脉粥样硬化的发生发展。TMAO还可促进血小板聚集，加快动脉血栓的形成，导致卒中的发生[28-30]，同时可通过影响卒中的其他危险因素（如胰岛素抵抗、糖尿病、肥胖等）而影响卒中的发生发展[31-33]。
目前对于TMAO与PSCI的关系研究较少。有研究证实，缺血性卒中患者简易智力状态检查量表（mini-mental state examination，MMSE）评分与TMAO浓度有关，TMAO浓度较高的急性缺血性卒中患者发生PSCI的可能性更高。PSCI与美国国立卫生研究院卒中量表（National Institutes of Health Stroke Scale，NIHSS）评分、卒中复发率、超敏C反应蛋白（hypersensitive C-reactive protein，hs-CRP）水平和TMAO浓度均呈正相关。但在控制上述混杂因素后，TMAO浓度仍为PSCI的独立预测因子[34]。TMAO浓度升高可加重动脉粥样硬化和颈动脉狭窄，同时与卒中患者的认知功能障碍有关[35-36]。
1.3 与帕金森病(Parkinson's disease，PD)认知功能障碍的关系
PD是一种常见的中老年神经退行性疾病。认知功能障碍是PD常见的非运动症状，包括帕金森病轻度认知功能障碍（mild cognitive impairment in Parkinson's disease，PD-MCI）和帕金森病痴呆（Parkinson's disease dementia，PDD），其中PD-MCI是PDD发生的独立危险因素。研究发现，PD-MCI在PD中的患病率约为40%[37]，每年进展为PDD的概率为6%~15%，且62%的PD-MCI患者在4年内进展为PDD[38]。PD认知功能障碍加之PD严重的运动障碍，给患者本身及照料者造成了沉重的生活和经济负担。因此，及早发现、及时干预、延缓病程进展是当前治疗PD认知功能障碍最有效的手段。
PD认知功能障碍的病理生理改变复杂，可能合并有皮质路易小体（Lewy body）形成、AD和脑微血管病变等多种病理改变，同时伴有多种神经递质的参与和广泛的神经元变性[39]。PD认知功能障碍患者存在明显的异质性，其发病既可在PD病程晚期，也可在PD诊断之初，累及的认知领域广泛，包括执行功能、记忆力、注意力、视空间能力、语言流畅性等。目前PD认知功能障碍发展的机制仍不清楚，脑脊液β淀粉样蛋白、血同型半胱氨酸、尿酸、抑郁症、快速眼动睡眠行为障碍、步态功能障碍、与白质病变相关的脑血管疾病、嗅觉功能障碍、APOE基因型、脑结构影像学改变等已被建议作为其发展的预测因子[40-41]。
研究发现，PD患者肠道菌群中乳杆菌科和肠杆菌科的丰度增高，而梭状芽胞杆菌、脆弱类杆菌和葡萄球菌的丰度较低，且肠杆菌科的丰度与姿势不稳和步态艰难的严重程度呈正相关，表明肠道菌群在PD临床表型中发挥了重要作用[42]。PD患者的肠道微生物群落发生改变[42-43]，而血浆中TMAO的浓度与肠道菌群有关[5]，故探讨PD患者血浆TMAO浓度与肠道菌群的关系已成为研究的重点。
目前关于TMAO与PD相关性的研究较少。Chung等[44]测定85例未接受药物治疗的早期PD患者和20名健康对照者的血浆TMAO浓度，依据TMAO浓度将PD患者分为高、中、低三组并进行随访（随访时间>2年），结果显示：（1）PD患者血浆TMAO浓度低于健康对照者；（2）PD各亚组间的纹状体多巴胺能去神经支配相似，但低TMAO浓度组较高TMAO浓度组需要更高剂量的多巴胺能药物才能有效控制症状；（3）血浆TMAO浓度预测PDD的最佳临界值为6.92 μmol/L，低TMAO浓度组的PDD转化风险高于高TMAO浓度组。该结果表明基线血浆TMAO浓度对早期PD患者具有一定预测作用，增高TMAO浓度可能有助于降低PD和PDD的发生风险。
一项关于唾液代谢的研究显示，与健康对照组相比，PD患者TMAO浓度升高；将PD患者分为早期PD组（疾病早期；H&Y分期≤2）与晚期PD组（疾病晚期；H&Y分期>2），早期PD组唾液中TMAO浓度明显高于健康对照组，但晚期PD组TMAO浓度并不高于健康对照组，推测TMAO浓度与PD分期有关[45]。另一项研究显示，晚期PD患者TMAO浓度升高，且有运动波动的患者脑脊液中TMAO浓度高于无运动波动的患者[11]。因此，PDD与TMAO浓度的关系目前尚未明确，推测与PD分期、标本来源等相关，今后仍需继续探索。
2 认知功能障碍疾病治疗前景：调控TMAO相关干预靶点
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TMAO与认知功能障碍疾病存在一定的关系，降低TMAO的浓度可能有利于认知功能障碍疾病的预防与治疗。目前研究表明，调整饮食结构、调整肠道菌群、抑制肝脏FMO活性均可影响TMAO的浓度，从而达到预防及治疗的目的。
由于磷脂酰胆碱、左旋肉碱是生成TMAO的主要原材料，而红肉及高脂肪乳制品是上述营养物质的主要来源，因此减少红肉摄入量、均衡膳食纤维可降低血浆中TMAO的浓度[46]。也有研究发现，一些天然小分子化合物如白藜芦醇、小檗碱等可降低外周血中TMAO的浓度[47-48]。
近年来，随着对肠道菌群认知的逐渐加深，学者们致力于通过重塑肠道菌群平衡而达到防治疾病的目的（表1），故肠道益生菌及粪菌移植成为研究热点。由于抗生素的广谱杀菌作用容易打破肠道固有生态菌群，目前不提倡应用抗生素治疗肠道菌群失调。目前研究显示，摄入益生菌对AD认知功能的改善有益，还有学者研究通过三联根除疗法消除幽门螺杆菌来改善AD患者的认知功能[49-50]。我国学者研制的甘露特钠胶囊（GV-971）可通过重塑肠道微生物群，抑制肠道细菌氨基酸型神经炎症，从而抑制AD的进展[51]。另有研究发现，我国传统针灸及补肾天水方也可调节肠道菌群，改善认知功能[52-53]。
3 总结与展望
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近年来研究表明，肠道菌群及其微生物代谢产物与认知功能障碍疾病存在一定的关系，肠道菌群失调可通过肠-脑轴对认知功能造成影响，在认知功能障碍疾病的发生发展中起着重要作用；通过饮食或非饮食方法调节肠道菌群能够对认知功能障碍疾病起到一定的干预作用，这为今后的临床诊治提供了新思路。TMAO作为肠道微生物的代谢产物，近年来已被公认为心血管疾病、糖尿病及肾脏疾病的独立危险因素，虽然部分研究表明血浆中的TMAO与认知功能障碍疾病相关，但二者之间的因果关系及具体影响机制尚未明确，甚至出现部分研究结果不一致的情况，这将是神经科学者们下一步探索的方向。
基金
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  • 云南省神经系统疾病诊疗中心应用基础研究(ZX2019-03-05)
文献
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2023年第48卷第3期
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doi: 10.11855/j.issn.0577-7402.2023.03.0339
  • 接收时间：2021-09-30
  • 首发时间：2025-12-03
  • 出版时间：2023-03-28
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  • 收稿日期：2021-09-30
  • 录用日期：2021-11-19
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Yunnan Neurological Disease Diagnosis and Treatment Center Applied Basic Research(ZX2019-03-05)
云南省神经系统疾病诊疗中心应用基础研究(ZX2019-03-05)
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    昆明医科大学第一附属医院老年神经内科，云南昆明 650000

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顾美娟，E-mail：
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2种不同金属材料的力学参数
科
Family		 属数
Number of
genus		 种数
Number of
species		 占总种数比例
Percentage of
total species (%)		 属
Genus		 种数
Number of
species		 占总种数比例
Percentage of total
species (%)
鹅膏菌科Amanitaceae 2 11 5.26 鹅膏菌属 Amanita 10 4.78
小菇科 Mycenaceae 2 12 5.74 丝盖伞属 Inocybe 5 2.39
多孔菌科 Polyporaceae 8 14 6.70 蜡蘑属 Laccaria 5 2.39
红菇科 Russulaceae 3 23 11.00 小皮伞属 Marasmius 6 2.87
小菇属 Mycena 11 5.26
光柄菇属 Pluteus 5 2.39
红菇属 Russula 17 8.13
栓菌属 Trametes 5 2.39
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