Article(id=1274057400870003617, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1274057338156769818, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250833, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1762358400000, receivedDateStr=2025-11-06, revisedDate=null, revisedDateStr=null, acceptedDate=1772640000000, acceptedDateStr=2026-03-05, onlineDate=1781688555209, onlineDateStr=2026-06-17, pubDate=1780502400000, pubDateStr=2026-06-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1781688555209, onlineIssueDateStr=2026-06-17, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1781688555209, creator=13701087609, updateTime=1781688555209, updator=13701087609, issue=Issue{id=1274057338156769818, tenantId=1146029695717560320, journalId=1192105938417971205, year='2026', volume='66', issue='6', pageStart='2561', pageEnd='3114', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=1, specialIssue=null, createTime=1781688540257, creator=13701087609, updateTime=1781688602467, updator=13701087609, preIssue=null, nextIssue=null, ext={EN=IssueExt(id=1274057599193486082, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1274057338156769818, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1274057599193486083, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1274057338156769818, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null}, startPage=2617, endPage=2633, ext={EN=ArticleExt(id=1274057401268462499, articleId=1274057400870003617, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Research advances in mechanisms of microbial carbon fixation and sequestration in mangrove sediments, columnId=1192149543727808575, journalTitle=Acta Microbiologica Sinica, columnName=Review, runingTitle=null, highlight=null, articleAbstract=

Mangrove ecosystems, situated at the land-sea interface, serve as vital blue carbon sinks, playing a key role in the global carbon cycle and climate regulation with their efficient carbon sequestration capacity. Microorganisms are central drivers of carbon sequestration in mangrove sediments, capable of fixing carbon through diverse metabolic pathways. This review first summarizes the currently identified microbial carbon fixation pathways and carbon sequestration mechanisms in mangrove sediments, with a focus on three primary processes: the Calvin-Benson-Bassham cycle, the reductive tricarboxylic acid cycle, and the reductive acetyl-CoA (Wood-Ljungdahl) pathway. Furthermore, we discuss the influences of key environmental factors, such as vegetation type, sediment physicochemical properties, and nutrient inputs, on microbial carbon fixation and sequestration. Finally, we propose the future directions for studies on microbial carbon fixation and sequestration in mangrove sediments, including the couplings of nutrient cycling processes, microbiome engineering, and microorganism-plant interactions. This review proposes potential novel strategies for enhancing blue carbon capacity in mangrove ecosystems.

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红树林生态系统位于陆海交汇带,是重要的蓝碳汇,凭借其高效的碳汇能力,在全球碳循环和气候调节中发挥着关键作用。微生物是红树林沉积物中碳储存的核心驱动因素,可通过多种途径实现固碳。本文系统综述了目前红树林沉积物中已发现的微生物固碳途径及其储碳机制,重点关注卡尔文循环、还原型三羧酸循环和还原性乙酰辅酶A途径这3种主要固碳途径及其储碳机制。进一步地,本文阐述了植被类型、沉积物理化性质、营养输入等关键环境因子对微生物碳固定和碳储存的影响。最后,展望了红树林沉积物微生物固碳和储碳的研究前景,包括多元素循环与耦合储碳、微生物组工程以及微生物-植物互作与协同增汇。本文为提高红树林生态系统的蓝碳能力提出了潜在的新策略。

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作者贡献声明

何煦妍:文章撰写;黄华夏:协助文章撰写,表格绘制;闫博治:图片绘制;刘焕平:格式校对;于晓莉:文章修改;颜庆云:文章审阅与修改;贺志理:整体构思,文章审阅与修改。

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Summary of microbial carbon fixation pathways in mangrove sediments: taxa, functional genes, and niches

, figureFileSmall=null, figureFileBig=null, tableContent=
Carbon fixation pathwaysMicrobial taxaFunctional genesNiches
CBBPseudomonadota, Halobacteriota, Candidatus Asgardarchaeota, Actinomycetota, ThermoplasmatotacbbL/cbbM, cbbS, prkB, rbcL/SSurface layer
rTCACampylobacterota, Thermoplasmatota

aclA/B, korABCD,

frdABCDE, ccsA/B, ccl

Middle layer, deep layer
WLPseudomonadota, Chloroflexota, Acidobacteriota, Spirochaetota, Nitrospirota, Halobacteriota, Thermoproteota, ThermoplasmatotaacsA/B, cdhA/C, fhs, fdhADeep layer
3-HPHaven’t been explicitly reported in mangrove sedimentsmcr, mcl, meh, mct-K14470There is no evidence that these pathways constitute a complete function in mangrove sediments
DC/4HBporABCD, porG, mcmA1/A2, hed, K15039
3HP/4HBfumABCDE, sucD/C, hcd, K15016
rGlygcvP, gcvT, gcvH, fhs, fchA, folD, glyA
), ArticleFig(id=1274088179759092520, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1274057400870003617, language=CN, label=表1, caption=

红树林沉积物中不同固碳途径的类群、功能基因与生态位特征

, figureFileSmall=null, figureFileBig=null, tableContent=
Carbon fixation pathwaysMicrobial taxaFunctional genesNiches
CBBPseudomonadota, Halobacteriota, Candidatus Asgardarchaeota, Actinomycetota, ThermoplasmatotacbbL/cbbM, cbbS, prkB, rbcL/SSurface layer
rTCACampylobacterota, Thermoplasmatota

aclA/B, korABCD,

frdABCDE, ccsA/B, ccl

Middle layer, deep layer
WLPseudomonadota, Chloroflexota, Acidobacteriota, Spirochaetota, Nitrospirota, Halobacteriota, Thermoproteota, ThermoplasmatotaacsA/B, cdhA/C, fhs, fdhADeep layer
3-HPHaven’t been explicitly reported in mangrove sedimentsmcr, mcl, meh, mct-K14470There is no evidence that these pathways constitute a complete function in mangrove sediments
DC/4HBporABCD, porG, mcmA1/A2, hed, K15039
3HP/4HBfumABCDE, sucD/C, hcd, K15016
rGlygcvP, gcvT, gcvH, fhs, fchA, folD, glyA
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红树林沉积物微生物驱动的固碳与储碳机制研究进展
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何煦妍 , 黄华夏 , 闫博治 , 刘焕平 , 于晓莉 , 颜庆云 , 贺志理
微生物学报 | 综述 2026,66(6): 2617-2633
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微生物学报 | 综述 2026, 66(6): 2617-2633
红树林沉积物微生物驱动的固碳与储碳机制研究进展
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何煦妍, 黄华夏, 闫博治, 刘焕平, 于晓莉, 颜庆云, 贺志理
作者信息
  • 南方海洋科学与工程广东省实验室(珠海),海洋合成生态学研究中心,广东 珠海
Research advances in mechanisms of microbial carbon fixation and sequestration in mangrove sediments
Xuyan HE, Huaxia HUANG, Bozhi YAN, Huanping LIU, Xiaoli YU, Qingyun YAN, Zhili HE
Affiliations
  • Marine Synthetic Ecology Research Center, Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai), Zhuhai, Guangdong, China
出版时间: 2026-06-04 doi: 10.13343/j.cnki.wsxb.20250833
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红树林生态系统位于陆海交汇带,是重要的蓝碳汇,凭借其高效的碳汇能力,在全球碳循环和气候调节中发挥着关键作用。微生物是红树林沉积物中碳储存的核心驱动因素,可通过多种途径实现固碳。本文系统综述了目前红树林沉积物中已发现的微生物固碳途径及其储碳机制,重点关注卡尔文循环、还原型三羧酸循环和还原性乙酰辅酶A途径这3种主要固碳途径及其储碳机制。进一步地,本文阐述了植被类型、沉积物理化性质、营养输入等关键环境因子对微生物碳固定和碳储存的影响。最后,展望了红树林沉积物微生物固碳和储碳的研究前景,包括多元素循环与耦合储碳、微生物组工程以及微生物-植物互作与协同增汇。本文为提高红树林生态系统的蓝碳能力提出了潜在的新策略。

红树林  /  微生物固碳  /  沉积物  /  碳汇

Mangrove ecosystems, situated at the land-sea interface, serve as vital blue carbon sinks, playing a key role in the global carbon cycle and climate regulation with their efficient carbon sequestration capacity. Microorganisms are central drivers of carbon sequestration in mangrove sediments, capable of fixing carbon through diverse metabolic pathways. This review first summarizes the currently identified microbial carbon fixation pathways and carbon sequestration mechanisms in mangrove sediments, with a focus on three primary processes: the Calvin-Benson-Bassham cycle, the reductive tricarboxylic acid cycle, and the reductive acetyl-CoA (Wood-Ljungdahl) pathway. Furthermore, we discuss the influences of key environmental factors, such as vegetation type, sediment physicochemical properties, and nutrient inputs, on microbial carbon fixation and sequestration. Finally, we propose the future directions for studies on microbial carbon fixation and sequestration in mangrove sediments, including the couplings of nutrient cycling processes, microbiome engineering, and microorganism-plant interactions. This review proposes potential novel strategies for enhancing blue carbon capacity in mangrove ecosystems.

mangrove  /  microbial carbon fixation  /  sediment  /  carbon sink
何煦妍, 黄华夏, 闫博治, 刘焕平, 于晓莉, 颜庆云, 贺志理. 红树林沉积物微生物驱动的固碳与储碳机制研究进展. 微生物学报, 2026 , 66 (6) : 2617 -2633 . DOI: 10.13343/j.cnki.wsxb.20250833
Xuyan HE, Huaxia HUANG, Bozhi YAN, Huanping LIU, Xiaoli YU, Qingyun YAN, Zhili HE. Research advances in mechanisms of microbial carbon fixation and sequestration in mangrove sediments[J]. Acta Microbiologica Sinica, 2026 , 66 (6) : 2617 -2633 . DOI: 10.13343/j.cnki.wsxb.20250833
红树林是由盐生乔木和灌木物种组成的沿海潮间带湿地生态系统,主要分布于亚热带和热带地区,具有强大的碳汇能力,是世界上生态系统服务价值最高的生态系统之一[1]。尽管红树林生态系统仅占全球滨海生态系统总面积的0.5%,但其单位面积生产力是海草床的5倍、盐沼的2倍,贡献了全球滨海10%-15%的碳储量[2]。研究表明,红树林生态系统最主要的碳封存发生在沉积物中,其碳储量占总碳库的50%以上[3]。随着全球城市化进程加快以及气候变化加剧,红树林沉积物碳循环过程及其碳汇机制正受到越来越多的关注[4]。有研究关注不同水文地貌类型红树林沉积物碳储量的差异:河口型红树林沉积物的碳储量占比为71%-98%,而海洋型红树林为49%-90%[5]。在全球尺度上,Atwood等[6]基于多源数据整合,估算全球红树林沉积物碳储量约为2.6 Pg C。Kauffman等[3]基于对全球不同地区红树林碳储量的系统评估,认为红树林地下碳储量可达10.2 Pg C。在此背景下,基于全球红树林20年土地变化的研究显示,红树林沉积物在此期间可累计固存约19.4 Tg C,并通过蓝碳交易产生约0.69-2.36亿美元的潜在收益,凸显了以沉积物碳封存为核心的红树林蓝碳恢复项目的巨大潜力[7]
红树林沉积物微生物是推动碳汇形成、维持碳封存的主要驱动因素。红树林一方面可通过红树植物光合作用吸收二氧化碳(carbon dioxide, CO2),并以凋落物和根际分泌物形式将有机碳输送到沉积物中[8-9];另一方面,潮汐交换过程会将近岸水体中的有机碳和无机碳输入红树林沉积物,从而形成重要的外源碳补给途径[2]。基于此,沉积物微生物通过多种代谢途径将无机碳转化为有机碳,进而增强红树林生态系统的碳汇功能。红树林这一特殊生境通过塑造独特的沉积物微环境,驱动生态位的分化与多样性,孕育了多样化的微生物群落和功能[10]。红树林沉积物微生物不仅能以光能自养、化能自养及异养代谢直接或间接参与沉积物的碳固定,还可通过与氮、硫、磷等元素循环耦合,共同调控碳转化与封存效率[11]。在氧气长期匮乏的深层沉积物中,厌氧微生物介导的碳固定过程构成红树林蓝碳系统中的关键环节,大量有机碳通过微生物转化储存于沉积物中,形成稳定碳库[12-14]。尽管目前已从自然界中发现了7种微生物自养固碳途径[15-16],但红树林生态系统中仅发现了卡尔文循环(Calvin-Benson-Bassham cycle, CBB)、还原型三羧酸循环(reductive tricarboxylic acid cycle, rTCA)、还原性乙酰辅酶A途径(reductive acetyl-CoA pathway, Wood-Ljungdahl pathway, WL)、二羧酸/4-羟基丁酸循环(dicarboxylate/4-hydroxybutyrate cycle, DC/4HB)、3-羟基丙酸/4-羟基丁酸循环(3-hydroxypropionate/4-hydroxybutyrate cycle, 3HP/4HB)和3-羟基丙酸循环(3-hydroxypropionate bicycle, 3HP)这6种固碳途径的关键功能基因,尚未发现还原性甘氨酸途径(reductive glycine pathway, rGly)的相关证据[17],而且3HP/4HB的关键功能基因丰度较低,DC/4HB和3HP的关键功能基因缺失[18]。因此,目前普遍认为红树林沉积物微生物的固碳过程主要依赖CBB、rTCA和WL这3种途径。
近年来,红树林沉积物中微生物驱动的固碳与储碳机制研究取得了突破性进展,进一步明确了CBB、rTCA和WL这3种核心固碳途径的生态分布特征、环境因子响应规律及与其他生物地球化学循环过程的耦合机制(图1)。本文重点综述了红树林沉积物中微生物驱动的CBB、rTCA和WL这3种核心固碳途径、机制及其关键影响因素,并探讨了相关研究的发展趋势及其主要挑战。
在红树林沉积物体系中,目前重点关注的CBB、rTCA和WL途径在能量需求、主导微生物类群及环境适应性上存在明显差异(表1)[16,18]
卡尔文循环(CBB)是自然界中最常见的固碳途径,广泛存在于藻类和多种化能自养微生物中,被认为是海洋与陆地生态系统自养微生物的主要固碳途径[19],通常在好氧或微氧、能量供应充足的环境中发挥作用。红树林沉积物中广泛存在CBB循环固碳过程,近期研究还发现超出传统认知的好氧光合类群也参与其中。例如,在深圳福田红树林自然保护区首次通过宏基因组鉴定出118个含CBB循环关键基因的宏基因组装基因组(metagenome-assembled genomes, MAGs),主要归属于假单胞菌门(Pseudomonadota)、MethanobacteriotaHalobacteriotaCandidatus Bathyarchaeota和Asgard archaea等[20]。此外,针对中国与南美洲21个红树林近400份沉积物样本的研究显示,不同区域样品中均有超10%的微生物携带CBB循环相关基因,且主要归属于假单胞菌门,这一结果表明CBB循环是红树林沉积物微生物的重要固碳途径[21]。本课题组对广东淇澳岛红树林沉积物的研究发现,CBB循环是该区域沉积物微生物最主要的化能自养固碳途径;其核心类群为化能自养硫氧化菌伯克霍尔德氏菌目(Burkholderiales)新谱系,该类群贡献了超75%的CBB循环功能,其基因组中不仅携带完整的CBB循环关键基因,还具有硫氧化、硝酸盐还原等功能基因,进一步支撑其在红树林沉积物化学自养固碳中的核心地位[22]
早期研究普遍认为CBB循环主要由细菌主导,随着研究的深入,近年来还在红树林沉积物中发现了古菌介导的CBB循环新证据。对中国香港米埔红树林沉积物中Asgard超门中Candidatus Thorarchaeota的分析发现,该类古菌门携带的CBB循环相关基因近乎完整,但其基因组均缺失CBB循环的关键基因prk,且其通过CBB循环固碳的机制尚未得到验证[23]。随后,进一步分析了红树林沉积物中Candidatus Bathyarchaeota的基因组,首次在古菌中识别出III-a型核酮糖-1,5-二磷酸羧化酶/加氧酶(ribulose-1,5-bisphosphate carboxylase/oxygenase, RuBisCO),并检测到部分亚群中prk基因的活性,表明这些古菌类群具备完整的CBB循环代谢潜力,并可能在微氧甚至厌氧环境中完成碳固定[24]。此外,一些古菌MAGs还同时携带CBB与WL循环关键基因,推测这些微生物具有灵活的代谢策略,可在不同能量与氧化还原条件下切换碳固定机制以适应红树林沉积物复杂多变的环境[20]
随着对CBB固碳微生物功能研究的不断深入,越来越多的研究表明,CBB循环常与硫循环、氮循环及有机碳分解等过程协同互作,形成复杂耦合网络。例如,在本地种红树林沉积物中发现硫氧化、碳降解、反硝化等功能基因与CBB循环基因通常协同表达,推测其可能通过硫氧化产生的能量驱动CO₂固定,同时利用硝酸盐/亚硝酸盐作为电子受体增强固碳效率[22]。通过分析巴西热带红树林沉积物的宏基因组发现,Thiohalomonadales代表种TH94含有CBB循环固碳的关键基因rbcLSprk,且能够利用硫氧化产生的能量驱动反硝化,从而实现硫循环、氮循环与碳固定的耦合[25]。此外,针对印度孙德尔本斯红树林碳调节基因的研究发现,红树林沉积物中富集的三酰甘油脂肪酶、硝酸盐还原酶(NarG基因编码)等功能基因通常与CBB循环关键基因协同表达,形成“碳分解-氮还原-硫代谢-碳固定”耦合网络[26]。这种协同作用为微生物在沉积物中实现碳汇调控与固碳效率提升提供了代谢基础。
与邻近的盐沼沉积物相比,红树林沉积物中CBB循环关键功能基因的丰度普遍较高,且相关基因主要来自硫氧化细菌的硫碱弧菌属(Thioalkalivibrio)和硫杆菌属(Thiobacillus)等[27]。这些细菌通过CBB循环促进红树林沉积物固碳,减少红树林区域CO2排放,在红树林生态系统整体功能运转中发挥作用。
还原型三羧酸(rTCA)循环是一种在厌氧或微氧条件下的自养固碳途径,其本质是三羧酸循环(tricarboxylic acid cycle, TCA)的逆过程,通过一系列还原反应将3个CO2分子整合为1个乙酰辅酶A分子,为细胞提供碳骨架。rTCA具有较高的能量效率,每固定1 mol CO2仅需约1 mol ATP[28]。这种途径在红树林沉积物的中深层厌氧环境中广泛存在,尤其适配于氧气匮乏、有机质丰富的还原性生境,是沉积物中自养固碳的重要组成途径之一[19,29]
在红树林沉积物中,rTCA循环主要发生在中层与深层厌氧区域,有机质积累和还原性增强为rTCA循环提供了适宜的能量与电子供体。已有研究表明,尽管rTCA循环在红树林沉积物中的分布不如CBB和WL途径广泛,但多种厌氧微生物固碳均依赖rTCA途径。Laux等[25]从炼油厂附近的热带红树林沉积物宏基因组中重建的港口球菌科(Porticoccaceae)、脱硫杆状菌目(Desulfobacterales)和Candidatus Zixiibacteriota等微生物中发现了rTCA循环的关键基因,包括编码ATP依赖的柠檬酸裂解酶基因(aclAB)和2-氧戊二酸:铁氧还蛋白氧化还原酶基因(korAB)等,这些基因在自养条件下具有活性,能够通过rTCA循环固定CO2。在广东淇澳岛红树林沉积物中,rTCA途径的主要贡献者为弯曲杆菌目(Campylobacterales)和产甲烷线菌目(Methanotrichales),其相对丰度分别为60.5%和39.5%[22]。此外,福建漳州红树林湿地的分析结果也显示,来自Campylobacteria的微生物携带有完整的rTCA基因簇,是rTCA途径的主要贡献者[30]
随着研究的深入,多种具有rTCA途径的新型微生物不断从红树林沉积物中发现。例如,Zhang等[20]利用基于二代三代混合拼装的宏基因组技术从深圳福田红树林沉积物中鉴定了一个具有rTCA途径的细菌新门,命名为Candidatus Cosmopoliota。Zhong等[31]从福建九龙江口红树林沉积物中分离鉴定了2个硫单胞菌属的新种Sulfurimonas microaerophilic HSL1-7ᵀ和Sulfurimonas diazotrophicus HSL3-1ᵀ,发现这2个新种的基因组均编码rTCA循环的aclABoorABCDporABCD等关键基因。随后,Cui等[32]也从该区域分离到7株隶属于弯曲杆菌纲的潜在新种,这些菌株均携带了完整的rTCA循环关键基因,包括ATP柠檬酸裂合酶(ATP-citrate lyase, Acl)、丙酮酸:铁氧还蛋白氧化还原酶(pyruvate:ferredoxin oxidoreductase, Por)、2-氧戊二酸:铁氧还蛋白氧化还原酶(2-oxoglutarate:ferredoxin oxidoreductase, Oor)这3个关键酶的基因。
研究表明,rTCA循环的关键酶Por和Oor对氧气高度敏感。四亚基型Por蛋白能够在低氧条件下高效运行,而三亚基和四亚基型Oor蛋白分别适用于微氧与厌氧环境[19]。福建九龙江口红树林沉积物中发现的7株菌均含有三亚基和四亚基型Oor,以及四亚基Por,而硫单胞菌属中的Oor和Por几乎均为四亚基型,推测相关酶结构组成有效提高了菌株在动态潮汐梯度下固碳活性的稳定性[32]。值得注意的是,这些微生物通常同时携带硫代谢相关Sox基因簇(如soxABXYZsoxCDYZ),并与rTCA关键基因协同表达,形成以还原性硫化物氧化为能量来源、以rTCA循环为碳固定途径的耦合代谢网络。正是由于rTCA循环可与硫氧化等高效能量获取过程紧密耦合,使相关微生物在红树林沉积物这种高硫、低氧且氧化还原条件剧烈波动的环境中,能够以较低能量成本持续固碳,从而在生态位竞争中占据优势。
还原性乙酰辅酶A (WL)途径是一种高能效的厌氧固碳途径,能够在不依赖光的情况下利用1分子ATP固定1分子CO2,是目前已知能耗最低的碳固定途径[33-34]。该途径与氢代谢、甲烷代谢及硫还原过程紧密耦合,利用铁氧还蛋白作为电子供体,在对氧高度敏感的一氧化碳脱氢酶/乙酰辅酶A合成酶复合物(carbon monoxide dehydrogenase/acetyl‑CoA synthase, CODH/ACS)的驱动下进行碳还原反应。由于红树林深层沉积物中电子供体丰富且缺乏氧气,WL途径成为其中关键的固碳途径,可提升红树林的碳储存功能与效率[35-37]。已有研究表明,WL途径在红树林沉积物碳循环中发挥的作用远不只是单纯的碳固定功能,而是作为一个代谢枢纽,连接了碳、硫、氮等多种元素的生物地球化学循环。
在碳循环方面,WL途径与产甲烷过程密切关联,产甲烷菌通过WL途径在碳降解和固碳底物供应中起双重作用。红树林沉积物中的甲烷微菌目(Methanomicrobiales)含有完整的WL途径甲基分支基因簇,包括fwdftrmchmtd等关键基因,这些基因可将CO2逐步转化为甲基四氢叶酸(methyl-tetrahydrofolate, methyl-THF),再通过甲基还原酶生成甲烷,而代谢中间产物乙酰辅酶A则可作为WL途径的固碳底物[38]。其次,甲烷代谢与碳固定过程紧密耦合,甲烷氧化过程中产生的甲酸可作为代谢桥梁,经甲酸脱氢酶催化后转化为WL途径甲基分支的起始底物。
硫循环与WL途径的耦合是红树林深层沉积物能量利用的另一特征。硫氧化菌通过降解有机碳生成甲酸或CO2等固碳底物,为硫酸盐还原菌通过WL途径固碳提供必要的底物[39]。在这一过程中,微生物利用这些底物进行碳固定,而硫酸盐还原过程产生的能量则可反向驱动WL途径的持续运行。这种群落水平的代谢协同模式不同于其他生态系统中由单一类群驱动WL途径的情况,凸显了红树林生境下微生物群落的协同特性[40-41]。近年来的研究进一步阐明了这种联系的普遍性和重要性。针对微生物驱动的化能自养固碳过程分析发现,WL途径相关类群在深层沉积物(尤其是低氧或厌氧区)丰度更高,主要包括厌氧绳菌目(Anaerolineales)和脱硫杆状菌目(Desulfobacterales)[22]。这些类群的MAGs携带WL途径关键基因,包括编码一氧化碳脱氢酶和乙酰辅酶A合成酶的基因,且通常与硫酸盐还原基因(dsrAB)同时被检测到。同样地,脱硫杆状菌目(Desulfobacterales)和隶属硝化螺菌门(Nitrospirota)的Candidatus Sulfobium属携带WL途径的全套关键基因,并与dsrAB存在明显的协同作用[25]。在有机质相对匮乏的深层沉积物中,硫还原供能与WL途径固碳结合模式提升了厌氧环境的碳利用效率。
在氮限制环境下,WL途径通过与固氮过程的耦合维持了红树林的生产力。研究发现,来自红树林深层沉积物的3个固氮群落[分别隶属于厌氧绳菌纲(Anaerolineae)、地杆菌科(Geobacteraceae)、脱硫单胞菌科(Desulfuromonadaceae)]的MAGs携带了完整的WL途径相关基因;这些固氮微生物多为兼性厌氧菌,含有低氧代谢基因,能够通过WL途径合成乙酰辅酶A,从而进入TCA循环进行氧化供能,支持高能耗的固氮过程[42]。这种机制表明,部分固氮微生物可通过混合营养的策略适应低氧的极端环境,在红树林深层沉积物的氮限制环境中生存和繁衍,为红树林生产力维持及沉积物碳汇形成提供了微生物层面的支撑机制。
综上所述,CBB、rTCA和WL途径共同构成了红树林沉积物中微生物固碳的主要代谢途径,但现有研究多基于功能基因分布或代谢潜力推断其固碳作用,尚缺乏对不同固碳途径在原位固碳贡献的定量评估。现有研究大多基于系统发育和宏基因组进行推断,也有研究通过原位过程测量方法直接量化红树林沉积物中微生物驱动的暗碳固定(dark carbon fixation, DCF)。例如,基于14C标记碳酸氢盐法的研究结果显示,红树林沉积物中DCF速率为0.02-3.27 mmol C/(m2·d),结合宏基因组和宏转录组分析确认了CBB途径在表层沉积物活性碳固定中的贡献,以及rTCA和WL途径在深层的实际运行[18]。然而,在微生物固碳与氮、硫循环已被证实存在耦合的背景下,不同固碳途径在该耦合体系中的实际固碳效率、对净碳汇的相对贡献仍有待通过通量测量进一步厘清。尤其是固碳产物在后续有机质转化过程中如何参与微生物残体碳的形成,目前相关的系统研究仍然有限。
红树林沉积物中微生物固碳过程受多重环境与生物因子的共同调控,其强度与稳定性在不同空间尺度上存在异质性。其中,红树植物类型及其与根际微生物的互作决定了沉积物中碳源输入方式和微环境特征,沉积物理化性质直接影响固碳微生物的代谢模式,而外源污染输入则可能通过干扰微生物功能过程改变固碳效率。本部分围绕植物-微生物互作、沉积物理化条件及人为扰动3类关键调控因素,综述其作用机制对微生物固碳过程的影响。
红树植物类型是影响红树林沉积物微生物固碳的关键生物因子。不同红树植物凭借其独特的生物学特性和根际过程,通过根际互作富集特定固碳功能菌群,进而调控沉积物中微生物的固碳效率。Mai等[43]对3种类型红树植物沉积物微生物群落的比较显示,正红树(Rhizophora apiculata)树种的根际环境对自养固碳菌的富集最为明显,其根际沉积物中与碳固定直接相关的功能菌群[如Rhizobiales和脱硫杆状菌目(Desulfobacterales)]的丰度明显高于杯萼海桑(Sonneratia alba)和小花木榄(Bruguiera parviflora)树种。红树主要通过根际分泌物与次生代谢物为这些功能菌群提供特异性碳源,改善微环境与功能菌群结构,从而促进微生物固碳作用。
进一步从宏观进化视角来看,红树物种的种类也对微生物群落结构和功能产生深刻影响。本地种红树林与引进种红树林根际微生物的对比研究表明,本地种秋茄(Kandelia obovata)在演化过程中筛选出基因组精简、固碳功能特化的微生物群落;这些微生物群落通过基因丢失过程剔除冗余基因,使硫酸盐还原基因与硫氧化基因在群落中具有较高的相对丰度,从而实现硫代谢与固碳过程的耦合;加之秋茄根际的高铵浓度环境也缓解了氮限制,强化了固氮与固碳的协同作用,最终形成一套碳-氮-硫协同的高效固碳体系[44]。相比之下,引进种无瓣海桑(Sonneratia apetala)的根际环境波动较大,微生物通常需要通过水平基因转移富集可移动遗传元件以适应环境,导致基因组扩张而稀释了固碳功能基因,从而造成碳、氮、硫循环功能潜力降低,最终限制了固碳效率[44]
外来植被入侵通过改变根际环境重塑红树林沉积物中不同微生物固碳途径的相对贡献。基于宏基因组与宏转录组的研究显示,在未受入侵的红树林中CBB循环相关基因在表层沉积物中表达丰度最高,约为59.48 TPM (transcripts per million),而WL与rTCA途径关键基因则在中深层沉积物中相对富集,分别达到44.53 TPM和37.82 TPM,反映了氧化还原梯度对固碳途径分异的调控[18]。互花米草入侵进一步放大了微生物固碳的重组效应。基因表达分析结果表明,在互花米草入侵背景下,红树林沉积物中CBB循环相关基因cbbL以及rTCA循环关键基因aclA/BkorA的丰度升高,同时RuBisCO酶活性显著增强,表明入侵植被可能通过提高底物有效性促进多种自养固碳途径的调整[45]
植物根际效应是影响不同红树植物招募特定固碳菌群的核心驱动力。其表现之一是,红树林根系通过构建距离依赖的微环境梯度,在根际不同区域形成差异化的固碳微环境。研究表明,根际不同区域(0-2、2-4、4-6 mm)的微生物多样性均高于非根际区域,且近根区域微生物生物量碳(microbial biomass carbon, MBC)与土壤微生物呼吸(soil microbial respiration, SMR)强度也明显升高[46]。在此基础上,红树林根系分泌物主要由糖类和低分子有机酸等小分子化合物组成,既作为碳源和能量来源,又作为化学信号直接调控根系微生物种群结构和代谢功能。相关研究表明,糖类和有机酸的持续输入可通过启动效应加速土壤有机碳(soil organic carbon, SOC)的分解,并促进有机碳向无机碳转化[47]。同时,在潮间带,有机酸还可改变红树根际溶解有机碳、氮和磷含量,提高根际微生物多样性及特定功能类群的丰度[48]。实证结果表明,放线菌在红树林根表圈中明显富集,固氮菌在根际圈和根表圈呈现特异性分布,且生丝微菌属(Hyphomicrobium)与甜菜碱、水杨酸、肉豆蔻酸、油酸、硬脂酸和棕榈酸等多种高丰度根系分泌物组分呈正相关[49]。可见,红树林根系分泌物通过调控微生物群落组成和代谢活性间接影响红树林沉积物中微生物固碳途径的功能发挥。
红树林沉积物的理化性质,特别是沉积物氧化还原条件、盐度、pH、有机质含量、粒径结构和沉积速率等因素直接影响微生物群落结构、代谢途径及其固碳效率。红树林沉积物普遍呈缺氧或厌氧状态,氧化还原电位随沉积物深度而发生梯度变化,进而驱动功能性微生物群落在垂直方向上的分异格局[11]。表层有氧环境富集了采用高氧适应策略的微生物,固碳相关功能基因丰度高;深层低氧或厌氧环境则富集了专性厌氧菌群,依赖厌氧呼吸耦合木质素单体衍生物(lignin monomer derivatives, LMDs)矿化,固碳基因丰度相对较低[50]。此外,红树林的生物扰动也可改变沉积物的氧化还原梯度,从而间接调控微生物固碳效率。基于全球典型红树林的研究表明,底栖生物扰动可通过改变沉积物中有机物积累和降解作用的空间分布从而影响微生物固碳效率;例如,蟹的掘穴活动可构建三维孔隙网络,改变沉积物中氧气的渗透,使洞穴壁的氧化还原电位高于非扰动沉积物,从而影响微生物群落的组成与分布[51]
盐度和pH也是影响红树林微生物结构和功能的关键环境因子。高盐度不仅直接筛选耐盐固碳菌群,还会抑制微生物残体碳(microbial necromass carbon, MNC)积累,加速有机碳的分解;pH则主要通过调控溶解无机碳形态和酶活性影响自养固碳过程[52]。沉积物中有机质和养分[总氮(total nitrogen, TN)、总磷(total phosphorus, TP)]协同影响微生物代谢模式。高有机质环境既可为异养和混合营养微生物提供丰富的碳源,还能影响微生物胞外聚合物(extracellular polymeric substances, EPS)的分泌,间接调控沉积物中微生物的固碳能力[53]。从长期尺度看,红树林沉积物理化性质对固碳效率的调控具有持续效应:沉积物中的有机碳密度、碳氮比、放射性碳年龄等环境理化性质分异能够长期调控微生物固碳效率;高C:N环境通过筛选高效氮利用型固碳菌提升单位氮固碳效率;泥炭的高碳吸附能力与黏土覆盖层的保护使碳留存率在千年尺度下仍保持较高水平[54]
沉积物的粒径结构与沉积速率则从物理层面调控碳的保存环境。较细的沉积物颗粒有助于有机碳的埋藏和稳定,同时创造稳定的厌氧环境以维持特定自养固碳途径;高沉积速率则能快速掩埋有机质,减少其分解潜力,从而促进长期碳固定。对巴西帕科蒂河红树林的研究表明,淤泥-黏土比例是影响细菌和古菌群落分布的关键因子,其中细颗粒条件下有机质保留率更高,且微生物多样性更高[55]。对广西6个代表性红树林湿地的研究结果进一步表明,沉积物粒径对不同微生物类群具有差异化筛选效应,黏土对绿屈挠菌门(Chloroflexota)呈正效应,而对蓝细菌门(Cyanobacteriota)呈负效应;砂粒则对蓝细菌门(Cyanobacteriota)有正效应[56]。此外,沉积物大团聚体(>0.25 mm)占比增加可以提升团聚体结构的稳定性,并增强碳汇能力[57]。从地貌类型及其演替背景看,河口型红树林因持续接受陆源矿物颗粒和颗粒有机碳输入,沉积速率和细颗粒比例较高,但增强的矿物输入也可能稀释沉积物中的SOC含量[8]。全球尺度的大样本分析表明,河口型红树林的单位面积SOC储量整体仍高于海洋型红树林,这主要受沉积环境、林龄结构及矿物稀释效应的共同影响[8,58]。总体而言,沉积物粒径结构、沉积速率及地貌背景通过共同塑造微生物生态位分化和有机质埋藏条件,调控红树林沉积物中微生物固碳途径的活性与长期碳汇潜力。
外源污染输入是影响红树林沉积物微生物固碳功能的主要人为干扰因素。随着沿海地区工农业生产经营活动的快速发展,尤其是养殖排放量的持续增加,营养盐、微塑料、重金属等多种污染物大量进入红树林生态系统,从菌群、基因到环境等多个维度破坏微生物固碳基础,对红树林碳汇功能构成严重威胁。
养殖排放作为人为外源污染,主要通过改变红树林沉积物环境来影响微生物群落结构与固碳功能。研究表明,虾塘养殖废水的排放通过引入高硫化物和低盐度等因子改变沉积物环境,降低伸长杆菌科(Prolixibacteraceae)等顽固性碳降解菌群的丰度,同时导致微生物胞外有机碳降解基因丰度下降、有氧呼吸增强,从而降低微生物碳利用效率(microbial carbon use efficiency, CUE),减少SOC储存[59]。此外,养殖排放还抑制CBB循环关键固碳微生物伯克霍尔德氏菌科(Burkholderiaceae)的活性,导致CBB和WL途径相关基因丰度下降[59]。这种由环境因素驱动,微生物群落及其基因组成共同作用的抑制效应削弱了红树林沉积物中微生物的固碳潜力。然而,外源营养输入对红树林沉积物微生物固碳的影响因环境而异:在富营养化区域,自然潮汐营养输入与人为污染叠加可能增强固碳作用,提升总有机碳(total organic carbon, TOC)与颗粒有机碳(particulate organic carbon, POC)含量,优化固碳菌所需的碳源供给与营养平衡;而在贫营养区域,同类输入则可能打破原有碳平衡[60]
微塑料主要通过群落结构重塑和代谢干扰影响固碳过程。首先,微塑料表面可形成特殊的生物膜,选择性富集具有塑料降解能力的微生物改变原有微生物群落结构,导致固碳功能菌群被竞争性排斥。其次,微塑料在沉积物中的积累改变沉积物的孔隙结构和氧气扩散,影响微生物代谢途径。He等[61]基于全国尺度的红树林研究表明,微塑料会定向富集产甲烷古菌[如甲烷八叠球菌属(Methanosarcina)]促进有机碳向甲烷转化,同时抑制甲烷氧化细菌[如甲基胞囊菌属(Methylocystis)]导致碳源流失与再固碳受阻。最近的研究还发现,毫米级聚乳酸(millimeter-scale polylactic acid, mm-PLA)微塑料的输入使红树林沉积物CO2排放量增加194%,表层土甲烷释放量提升17.95%,使沉积物源溶解性有机碳(dissolved organic carbon, DOC)以类蛋白质为主,生物指数(biological index, BIX)升高,从而调控红树林沉积物固碳功能[62]
汞、铅、镉等重金属主要通过毒性抑制与群落失衡破坏微生物固碳功能。一项涵盖38项研究的国际综合分析显示,重金属污染可导致红树林沉积物有机碳减少0.57%-8.06%;其作用机制包括:直接破坏固碳微生物细胞膜结构与酶活性,干扰有机碳固定与转化;筛选耐重金属优势类群,取代高效固碳微生物;打破碳收支平衡,使甲烷排放量增加90-150 μg/(g·d)导致有机碳以温室气体形式流失[63]。重金属的毒性累积效应显著,即使低浓度暴露也会长期抑制微生物固碳效率,且影响具有不可逆性。
此外,多种外源污染物共存时易产生协同效应,加剧对微生物固碳的破坏。微塑料-重金属复合污染可通过多种途径抑制固碳:(1) 选择性富集芽胞杆菌等耐污染微生物菌群,挤压固碳功能菌群的生态位;(2) 通过加剧毒性胁迫,下调固碳核心途径关键基因的表达;(3) 通过干扰硫、氮等元素循环切断固碳能量供给,或通过氮限制抑制固碳酶合成,破坏固碳微环境[64]。值得注意的是,重金属具有明显的毒性累积效应,且影响具有不可逆性,即使低浓度暴露也会长期抑制微生物固碳效率。有机污染物与重金属共存时可能通过影响沉积物理化性质(如pH、氧化还原电位)间接叠加干扰,导致微生物群落结构更趋简化,固碳效率大幅降低。目前,关于红树林沉积物中复合污染的具体作用机制、浓度阈值及长期生态效应仍需更多针对性原位观测与控制实验验证。
现有研究表明,红树植物类型、沉积物理化性质及外源污染输入均会影响红树林沉积物中微生物固碳潜力,但多数研究仍停留在环境因子与群落结构或功能基因丰度的相关性层面,对固碳过程本身的调控机制缺乏直接验证。特别是在多种环境因子共同作用的情景下,不同理化条件和生物因素如何通过作用于特定关键固碳功能微生物,进而影响沉积物净碳汇形成,尚未形成清晰的机制框架。与此同时,植物根系活动及其与微生物的协同互作虽被认为是调控固碳的重要环节,但其在微尺度上改变根际环境、引导微生物固碳的过程与机制尚不明确。
尽管有关红树林沉积物微生物驱动的固碳与储碳研究已取得重要进展,但对其在复杂环境背景下形成稳定碳汇的关键机制仍缺乏系统性和定量化认知[65]。这一认知局限主要源于两方面因素:(1) 受气候、潮汐节律、地形地貌等环境条件影响,红树林沉积物环境异质性高,微生物群落组成与功能时空变化大;(2) 气候变化及人类活动也不断改变红树林沉积物生态环境,进而影响微生物固碳过程[4,66]。因此,从多元素生物地球化学循环耦合、功能微生物重构到植物-微生物协同调控的多层级视角,深入解析红树林沉积物中微生物固碳储碳过程及其调控机制(图2),不仅对阐明红树林蓝碳的形成机制具有关键科学意义,同时也为推动微生物组工程在红树林生态系统中的应用提供了理论依据。
红树林沉积物中微生物驱动的碳循环与氮、硫、磷等元素循环深度耦合,这种多元素协同循环有助于调控碳汇效率与稳定性。尽管当前研究对红树林沉积物微生物的固碳潜力已有初步认识,但未来研究的核心科学问题在于阐明微生物固碳过程如何与氮、硫循环及有机质分解过程协同作用,从而促进微生物残体碳的形成与积累,以及在不同环境扰动背景下,这种元素循环耦合关系是否发生解耦并导致沉积物碳封存能力下降。未来研究需借助多组学技术(宏基因组、宏转录组、代谢组学)与高分辨率原位监测相结合的策略,系统解析红树林沉积物中固碳、氮硫转化与有机质分解等关键过程之间的功能联系。在此基础上,引入稳定同位素示踪[如13C标记的溶解无机碳(13C-labeled dissolved inorganic carbon, 13C-DIC)]与单细胞尺度分析技术[如荧光原位杂交联用纳米级次级离子质谱(fluorescence in situ hybridization coupled with nanoscale secondary ion mass spectrometry, FISH-nanoSIMS)]可定量评估微生物自养固碳通量及其向残体碳的转化效率,并在单细胞水平识别在不同环境条件下实际参与无机碳同化的关键类群。进一步地,将同位素通量数据与功能基因表达信息相整合,有助于理解微生物残体碳形成等长期碳封存过程,以及环境因子变化对元素循环耦合固碳过程的干扰效应(图3)。基于上述多尺度数据整合,未来研究还应发展融合关键微生物功能基因与多元素耦合过程的生态模型,用以模拟在气候变化背景下红树林沉积物碳汇功能的动态响应与反馈机制,从而为红树林蓝碳潜力的评估提供更具预测性的理论支撑。
微生物组工程通过精准调控核心功能微生物丰度、激活功能基因表达和优化代谢路径,为定向提升红树林沉积物碳汇潜力提供了新的技术途径。构建靶向固碳功能微生物组筛选与组装技术体系是微生物组工程技术应用的一个研究方向。微生物作为红树林沉积物碳循环的核心驱动者,其群落结构与功能直接决定碳转化效率与碳库稳定性。其中固碳菌、甲烷氧化菌、顽固性碳降解菌等功能微生物是影响红树林沉积物碳汇的关键微生物。目前,对红树林沉积物中碳固定与长期碳封存相关功能微生物的系统性认知仍然不足,并且对于不同微生物固碳途径在特定沉积环境中的生态位适配性及其对净碳汇的实际贡献尚缺乏直接证据。虽然已在海草床沉积物、深海热液、冷渗透区等缺氧生态系统检测到DC/4HB、3HP/4HB、3HP和rGly等固碳途径,但这些途径在特定红树林沉积物中的分布特征、环境适应条件及固碳通量还有待进一步研究[67]。未来研究可结合宏基因组功能注释与单细胞分选技术,从不同红树植被类型和特定沉积物生境筛选依赖不同固碳途径的核心功能菌,明确不同固碳途径的环境适配条件。进一步地,基于合成微生物生态学理论和方法,依据功能互补与环境适配原则,构建以高效固碳类群为核心,并与碳转化和稳定过程协同的人工微生物组,实现红树林沉积物微生物组工程的定向设计(图3)。需要强调的是,在引入外源功能菌群或构建人工微生物组的过程中应同步开展严格的生态环境风险评估,重点分析潜在的生物入侵风险,减少对本地微生物群落结构和生态功能稳定性的影响。此外,还需探索基于代谢调控的微生物功能强化路径,通过代谢工程手段减少甲烷生成相关基因表达,引导更多碳流向微生物残体碳或有机碳库。例如:基于“酶锁-微生物碳利用效率”机制的研究启示,可通过引入聚羟基脂肪酸酯(polyhydroxyalkanoate, PHA)合成基因调控工程菌群的碳代谢流向,引导更多LMDs转化为稳定的微生物残体碳,实现木质素高效降解和碳源精准固存的协同增汇[50]
红树植物根系与微生物群落协同互作是调控沉积物碳输入、转化与固存的关键环节,也是未来提升红树林碳汇功能的核心突破口。已有研究表明,红树植物根系分泌物的组成与沉积物菌群密切相关[68],但特定根系分泌物如何激活微生物固碳关键基因表达,共生微生物与根系的互作是否通过改善根际微环境优化固碳途径,仍缺乏分子水平的直接证据。在机制上,结合多组学手段筛选参与根际微生物-植物互作调控固碳的关键信号分子与功能模块,揭示植物通过化学信号和营养供给调控根际微生物固碳潜力的分子路径,并解析微生物代谢产物对植物根系碳分配策略的反馈作用。在功能验证层面,通过引入高分辨率原位监测技术(如微电极系统)和可控尺度的根际模拟实验体系可在微尺度上解析植物根系活动改变根际微环境的动态过程,从而评估根际氧化还原状态、pH及有机底物供给变化对微生物固碳功能的调控效应(图3)。在应用方面,发展基于根际微生态调控的固碳功能提升策略。当前红树林修复多聚焦植物补种,忽视了根际微生态的优化。针对退化红树林根际有害代谢产物积累、固碳功能微生物活性抑制等问题,可通过接种与红树植物共生的功能固碳菌剂重构高效固碳根际微生态体系。
综上所述,在未来的红树林湿地碳汇研究中,可从系统解析微生物驱动的元素循环与耦合机制、创新微生物组工程理论和技术、强化微生物-植物协同互作三方面入手,揭示红树林湿地生态系统中的固碳与储碳协同机制。
  • 南方海洋科学与工程广东省实验室(珠海)项目(SML2020SP004)
  • 南方海洋科学与工程广东省实验室(珠海)项目(SML2024SP002)
  • 南方海洋科学与工程广东省实验室(珠海)项目(SML2024SP022)
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2026年第66卷第6期
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doi: 10.13343/j.cnki.wsxb.20250833
  • 接收时间:2025-11-06
  • 首发时间:2026-06-17
  • 出版时间:2026-06-04
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  • 收稿日期:2025-11-06
  • 录用日期:2026-03-05
基金
the Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai) Projects(SML2020SP004)
南方海洋科学与工程广东省实验室(珠海)项目(SML2020SP004)
the Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai) Projects(SML2024SP002)
南方海洋科学与工程广东省实验室(珠海)项目(SML2024SP002)
the Southern Marine Science and Engineering Guangdong Laboratory (Zhuhai) Projects(SML2024SP022)
南方海洋科学与工程广东省实验室(珠海)项目(SML2024SP022)
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    南方海洋科学与工程广东省实验室(珠海),海洋合成生态学研究中心,广东 珠海
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2种不同金属材料的力学参数

Family
属数
Number of
genus
种数
Number of
species
占总种数比例
Percentage of
total species (%)

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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