Article(id=1194684379910873487, tenantId=1146029695717560320, journalId=1192105938417971205, issueId=1194684377813717012, articleNumber=null, orderNo=null, doi=10.13343/j.cnki.wsxb.20250252, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1743091200000, receivedDateStr=2025-03-28, revisedDate=null, revisedDateStr=null, acceptedDate=1747756800000, acceptedDateStr=2025-05-21, onlineDate=1762764552333, onlineDateStr=2025-11-10, pubDate=1762185600000, pubDateStr=2025-11-04, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1762764552333, onlineIssueDateStr=2025-11-10, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1762764552333, creator=13701087609, updateTime=1762764552333, updator=13701087609, issue=Issue{id=1194684377813717012, tenantId=1146029695717560320, journalId=1192105938417971205, year='2025', volume='65', issue='11', pageStart='4721', pageEnd='5182', issueExtLink='null', onlineDate='null', pubDate='null', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1762764551833, creator=13701087609, updateTime=1762764551833, updator=13701087609, preIssue=null, nextIssue=null, ext=null, issueFiles=null}, startPage=4877, endPage=4888, ext={EN=ArticleExt(id=1194684381039141267, articleId=1194684379910873487, tenantId=1146029695717560320, journalId=1192105938417971205, language=EN, title=Effects of forest mycorrhizal types on responses of understory herbaceous plants to nitrogen deposition, columnId=1192149543992045670, journalTitle=Acta Microbiologica Sinica, columnName=Research Article, runingTitle=null, highlight=null, articleAbstract=

With the development of human industrial activities, nitrogen (N) emissions and atmospheric N deposition have increased significantly. When atmospheric N deposition exceeds the critical load that plants can bear, it may exert a negative effect on plants and lead to a decrease in species abundance. As an important biotic factor affecting species abundance, mycorrhizae can affect plant diversity and community structure through nutrient supply and hyphal network mechanism. Objective To compare the critical load of N deposition of understory herbaceous plants with different mycorrhizal types and explore how mycorrhizal types affect the responses of understory herbaceous plants to N deposition. Methods According to the database of long-term N deposition critical load of forest herbaceous plants, and the published literature, critical load database of herbaceous plants under different mycorrhizal types of forest forests in response to N deposition was established. We identified three mycorrhizal types of forest dominant species, which included arbuscular mycorrhiza (AM), ectomycorrhiza (ECM), and AM+ECM. The effects of mycorrhizal types on the critical load of N deposition of understory herbaceous plants were investigated. Results The critical load of N deposition of understory herbaceous plants varied in forests of different mycorrhizal types (P<0.05). The critical load of N deposition was the highest [9.28 kg N/(ha·a)] in the forest of the AM+ECM type, the second [8.41 kg N/(ha·a)] in the forest of the ECM type, and the lowest [7.19 kg N/(ha·a)] in the forest of the AM type. In forests of different mycorrhizal types, the critical loads of understory herbaceous plants of different functional groups (gramineous and non-gramineous) in response to N precipitation were consistent with the responses of all understory herbaceous plants of different mycorrhizal types. N deposition caused changes in the abundance of understory herbaceous plants. The species abundance of understory herbaceous plants in the forest of the AM type showed an increasing trend, while that in the forest of the ECM type mainly decreased. Conclusion Mycorrhizal types affect the critical load of N deposition of understory herbaceous plants (P<0.05), which is related to the niche differentiation, N concentration of litter, and N acquisition strategies of different mycorrhizal types of plants. In addition, the abundance of understory herbaceous plants will also vary due to different mycorrhizal types.

, correspAuthors=Zhaoyong SHI, authorNote=null, correspAuthorsNote=
*E-mail:
, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, authorCompany=null, fund=null, authors=null, authorsList=Zengmeihui XU, Zhaoyong SHI, Hao WANG, Yan LI, Luping MA, Mengge ZHANG, Shanwei WU, Jiakai GAO, Shuangshuang WANG), CN=ArticleExt(id=1194684882883420613, articleId=1194684379910873487, tenantId=1146029695717560320, journalId=1192105938417971205, language=CN, title=森林菌根类型对林下草本植物响应氮沉降的影响, columnId=1192149544164012138, journalTitle=微生物学报, columnName=研究报告, runingTitle=null, highlight=null, articleAbstract=

随着人类工业活动的开展,氮(N)排放和大气氮沉降显著增加。当大气氮沉降超过植物所能承受的临界负荷时可能使植物产生负面响应,导致物种丰度降低。菌根作为影响物种丰度的一个重要生物因素,可通过养分供给和菌丝网络机制影响植物的多样性和群落结构。 目的 分析不同菌根类型林下草本植物氮沉降临界负荷的差异,探讨菌根类型如何影响林下草本植物对氮沉降的响应。 方法 基于森林草本植物长期氮沉降临界负荷数据库,结合已发表的文献资料,建立了“不同菌根类型森林林下草本植物对氮沉降响应的临界负荷数据库”,补充了森林优势种的菌根类型数据。该数据库共涵盖3种菌根类型:丛枝菌根(arbuscular mycorrhiza, AM)、外生菌根(ectomycorrhiza, ECM),以及受2种菌根共同侵染的双菌根(AM+ECM),以探究菌根类型对林下草本植物氮沉降临界负荷的影响。 结果 不同菌根类型森林下草本植物的氮沉降临界负荷存在显著差异(P<0.05)。AM+ECM型森林林下草本植物的氮沉降临界负荷最高,为9.28 kg N/(ha·a);ECM型次之,为8.41 kg N/(ha·a);AM型最低,为7.19 kg N/(ha·a)。在不同菌根类型森林中,不同功能群的林下草本植物(禾本科和非禾本科)对氮沉降响应的临界负荷与不同菌根类型林下全部草本植物的反应一致。氮沉降会引起林下草本植物的丰度变化,AM型森林草本植物物种丰度呈增加趋势,而含ECM的森林草本植物物种丰度以降低为主。 结论 菌根类型对林下草本植物的氮沉降临界负荷具有显著影响(P<0.05),这与不同菌根类型植物的生态位分化、凋落物氮含量浓度以及氮获取策略有关。此外,林下草本植物的丰度也会因菌根类型的不同而存在差异。

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Ecological impacts of nitrogen deposition on terrestrial ecosystems: research progresses and prospects[J]. Chinese Journal of Plant Ecology, 2020, 44(5): 475-493 (in Chinese)., articleTitle=null, refAbstract=null), Reference(id=1194980534473437677, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, doi=null, pmid=null, pmcid=null, year=2019, volume=686, issue=null, pageStart=1129, pageEnd=1139, url=null, language=null, rfNumber=[48], rfOrder=61, authorNames=MEI LL, YANG X, ZHANG SQ, ZHANG T, GUO JX, journalName=Science of The Total Environment, refType=null, unstructuredReference=MEI LL, YANG X, ZHANG SQ, ZHANG T, GUO JX. Arbuscular mycorrhizal fungi alleviate phosphorus limitation by reducing plant N:P ratios under warming and nitrogen addition in a temperate meadow ecosystem[J]. 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caption=Frequency distribution of critical nitrogen deposition load of herbaceous plants in forest with different mycorrhizal types. A: AM; B: ECM; C: AM+ECM., figureFileSmall=w+SvqWWttnu+Ngm0EhaMdw==, figureFileBig=jECxNX2zr/jzhCSpw7a8Hg==, tableContent=null), ArticleFig(id=1194980522049909076, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=CN, label=图1, caption=不同菌根类型森林中草本植物氮沉降临界负荷的频率分布, figureFileSmall=w+SvqWWttnu+Ngm0EhaMdw==, figureFileBig=jECxNX2zr/jzhCSpw7a8Hg==, tableContent=null), ArticleFig(id=1194980523127845205, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=EN, label=Figure 2, caption=Critical load of nitrogen deposition of herbaceous plants in forest with different mycorrhizal types. The circle point represents the mean. Different lowercase letters mean significant difference at 0.05 level., figureFileSmall=sKCeK/r1SuAJS4txDe54Tw==, figureFileBig=Ls8BPT/3lbUB7T4pJil6HA==, tableContent=null), ArticleFig(id=1194980523190759766, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=CN, label=图2, caption=不同菌根类型森林中草本植物的氮沉降临界负荷, figureFileSmall=sKCeK/r1SuAJS4txDe54Tw==, figureFileBig=Ls8BPT/3lbUB7T4pJil6HA==, tableContent=null), ArticleFig(id=1194980523257868631, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=EN, label=Figure 3, caption=Frequency distribution of critical load of nitrogen deposition in different herbaceous plants with different mycorrhizal types. A: AM gramineous plants; B: ECM gramineous plants; C: AM+ECM gramineous plants; D: AM non-gramineous plants; E: ECM non-gramineous plants; F: AM+ECM non-gramineous plants., figureFileSmall=MEcRRHpIkvbkUKna4zS1yw==, figureFileBig=OQHQ6+gVt+zEU0zPYDYr7A==, tableContent=null), ArticleFig(id=1194980523438223704, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=CN, label=图3, caption=不同菌根类型不同草本植物氮沉降临界负荷的频率分布, figureFileSmall=MEcRRHpIkvbkUKna4zS1yw==, figureFileBig=OQHQ6+gVt+zEU0zPYDYr7A==, tableContent=null), ArticleFig(id=1194980523505332569, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=EN, label=Figure 4, caption=The critical load of different mycorrhizal types and herbaceous plants affected by nitrogen deposition. The circle point represents the mean. Different lowercase letters mean significant difference at 0.05 level., figureFileSmall=grbVbKJVrnWigjvZdzhx/A==, figureFileBig=fZ6aVjqUX7gWo3GvyvfqFQ==, tableContent=null), ArticleFig(id=1194980523622773082, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=CN, label=图4, caption=不同菌根类型不同草本植物受氮沉降影响的临界负荷, figureFileSmall=grbVbKJVrnWigjvZdzhx/A==, figureFileBig=fZ6aVjqUX7gWo3GvyvfqFQ==, tableContent=null), ArticleFig(id=1194980523706659163, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=EN, label=Figure 5, caption=Frequency distribution of species abundance of herbaceous plants with different mycorrhizal types in response to nitrogen deposition. A: AM decreased abundance; B: ECM decreased abundance; C: AM+ECM decreased abundance; D: AM increased abundance; E: ECM increased abundance; F: AM+ECM increased abundance., figureFileSmall=j+AiCD9xMIvcvsCJSzfMCQ==, figureFileBig=s+xB4ByILCl9rhJkYP8o7g==, tableContent=null), ArticleFig(id=1194980523782156636, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=CN, label=图5, caption=不同菌根类型草本植物物种丰度对氮沉降响应的频率分布, figureFileSmall=j+AiCD9xMIvcvsCJSzfMCQ==, figureFileBig=s+xB4ByILCl9rhJkYP8o7g==, tableContent=null), ArticleFig(id=1194980523853459805, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=EN, label=Figure 6, caption=Critical load of abundance change of herbaceous plants with different mycorrhizal types affected by nitrogen deposition. The circle point represents the mean. Different lowercase letters meant significant difference at 0.05 level., figureFileSmall=s6QNUkj1q3DX1N+mtdaAMg==, figureFileBig=yG2TT6PeDXs65/tq6Bh8Tw==, tableContent=null), ArticleFig(id=1194980523966706014, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=CN, label=图6, caption=不同菌根类型草本植物受氮沉降影响丰度变化的临界负荷, figureFileSmall=s6QNUkj1q3DX1N+mtdaAMg==, figureFileBig=yG2TT6PeDXs65/tq6Bh8Tw==, tableContent=null), ArticleFig(id=1194980524058980703, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=EN, label=Figure 7, caption=Critical load of decreasing abundance of different herbaceous plants under different mycorrhizal types affected by nitrogen deposition. A: Decreased abundance; B: Increased abundance. The line represents the mean. Different lowercase letters meant significant difference at 0.05 level., figureFileSmall=p6OyqlA0HRb4wfvKl0OhUg==, figureFileBig=uBfRvcJAbtb/9DwWQSbTpQ==, tableContent=null), ArticleFig(id=1194980524130283872, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=CN, label=图7, caption=不同菌根类型森林不同草本植物物种丰度对氮沉降的响应, figureFileSmall=p6OyqlA0HRb4wfvKl0OhUg==, figureFileBig=uBfRvcJAbtb/9DwWQSbTpQ==, tableContent=null), ArticleFig(id=1194980524209975649, tenantId=1146029695717560320, journalId=1192105938417971205, articleId=1194684379910873487, language=EN, label=Figure 8, caption=Percentage of abundance change of understory herbaceous plants affected by nitrogen deposition in different mycorrhizal types. 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森林菌根类型对林下草本植物响应氮沉降的影响
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许增美慧 1, 2, 3 , 石兆勇 1, 2, 3, * , 王浩 1 , 李琰 1, 2, 3 , 马路平 1, 2, 3 , 张梦歌 1, 2, 3 , 吴姗薇 1, 2, 3 , 高佳凯 1, 2, 3 , 王双双 1, 2, 3
微生物学报 | 研究报告 2025,65(11): 4877-4888
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微生物学报 | 研究报告 2025, 65(11): 4877-4888
森林菌根类型对林下草本植物响应氮沉降的影响
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许增美慧1, 2, 3, 石兆勇1, 2, 3, * , 王浩1, 李琰1, 2, 3, 马路平1, 2, 3, 张梦歌1, 2, 3, 吴姗薇1, 2, 3, 高佳凯1, 2, 3, 王双双1, 2, 3
作者信息
  • 1 河南科技大学 农学院,河南 洛阳
  • 2 河南省乡村人居环境工程中心,河南 洛阳
  • 3 洛阳市共生微生物与绿色发展重点实验室,河南 洛阳
Effects of forest mycorrhizal types on responses of understory herbaceous plants to nitrogen deposition
Zengmeihui XU1, 2, 3, Zhaoyong SHI1, 2, 3, * , Hao WANG1, Yan LI1, 2, 3, Luping MA1, 2, 3, Mengge ZHANG1, 2, 3, Shanwei WU1, 2, 3, Jiakai GAO1, 2, 3, Shuangshuang WANG1, 2, 3
Affiliations
  • 1 College of Agriculture, Henan University of Science and Technology, Luoyang, Henan, China
  • 2 Henan Rural Human Settlement Environment Engineering Center, Luoyang, Henan, China
  • 3 Luoyang Key Laboratory of Symbiosis Microorganism and Green Development, Luoyang, Henan, China
出版时间: 2025-11-04 doi: 10.13343/j.cnki.wsxb.20250252
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随着人类工业活动的开展,氮(N)排放和大气氮沉降显著增加。当大气氮沉降超过植物所能承受的临界负荷时可能使植物产生负面响应,导致物种丰度降低。菌根作为影响物种丰度的一个重要生物因素,可通过养分供给和菌丝网络机制影响植物的多样性和群落结构。 目的 分析不同菌根类型林下草本植物氮沉降临界负荷的差异,探讨菌根类型如何影响林下草本植物对氮沉降的响应。 方法 基于森林草本植物长期氮沉降临界负荷数据库,结合已发表的文献资料,建立了“不同菌根类型森林林下草本植物对氮沉降响应的临界负荷数据库”,补充了森林优势种的菌根类型数据。该数据库共涵盖3种菌根类型:丛枝菌根(arbuscular mycorrhiza, AM)、外生菌根(ectomycorrhiza, ECM),以及受2种菌根共同侵染的双菌根(AM+ECM),以探究菌根类型对林下草本植物氮沉降临界负荷的影响。 结果 不同菌根类型森林下草本植物的氮沉降临界负荷存在显著差异(P<0.05)。AM+ECM型森林林下草本植物的氮沉降临界负荷最高,为9.28 kg N/(ha·a);ECM型次之,为8.41 kg N/(ha·a);AM型最低,为7.19 kg N/(ha·a)。在不同菌根类型森林中,不同功能群的林下草本植物(禾本科和非禾本科)对氮沉降响应的临界负荷与不同菌根类型林下全部草本植物的反应一致。氮沉降会引起林下草本植物的丰度变化,AM型森林草本植物物种丰度呈增加趋势,而含ECM的森林草本植物物种丰度以降低为主。 结论 菌根类型对林下草本植物的氮沉降临界负荷具有显著影响(P<0.05),这与不同菌根类型植物的生态位分化、凋落物氮含量浓度以及氮获取策略有关。此外,林下草本植物的丰度也会因菌根类型的不同而存在差异。

菌根类型  /  草本植物  /  氮沉降  /  临界负荷

With the development of human industrial activities, nitrogen (N) emissions and atmospheric N deposition have increased significantly. When atmospheric N deposition exceeds the critical load that plants can bear, it may exert a negative effect on plants and lead to a decrease in species abundance. As an important biotic factor affecting species abundance, mycorrhizae can affect plant diversity and community structure through nutrient supply and hyphal network mechanism. Objective To compare the critical load of N deposition of understory herbaceous plants with different mycorrhizal types and explore how mycorrhizal types affect the responses of understory herbaceous plants to N deposition. Methods According to the database of long-term N deposition critical load of forest herbaceous plants, and the published literature, critical load database of herbaceous plants under different mycorrhizal types of forest forests in response to N deposition was established. We identified three mycorrhizal types of forest dominant species, which included arbuscular mycorrhiza (AM), ectomycorrhiza (ECM), and AM+ECM. The effects of mycorrhizal types on the critical load of N deposition of understory herbaceous plants were investigated. Results The critical load of N deposition of understory herbaceous plants varied in forests of different mycorrhizal types (P<0.05). The critical load of N deposition was the highest [9.28 kg N/(ha·a)] in the forest of the AM+ECM type, the second [8.41 kg N/(ha·a)] in the forest of the ECM type, and the lowest [7.19 kg N/(ha·a)] in the forest of the AM type. In forests of different mycorrhizal types, the critical loads of understory herbaceous plants of different functional groups (gramineous and non-gramineous) in response to N precipitation were consistent with the responses of all understory herbaceous plants of different mycorrhizal types. N deposition caused changes in the abundance of understory herbaceous plants. The species abundance of understory herbaceous plants in the forest of the AM type showed an increasing trend, while that in the forest of the ECM type mainly decreased. Conclusion Mycorrhizal types affect the critical load of N deposition of understory herbaceous plants (P<0.05), which is related to the niche differentiation, N concentration of litter, and N acquisition strategies of different mycorrhizal types of plants. In addition, the abundance of understory herbaceous plants will also vary due to different mycorrhizal types.

mycorrhizal type  /  herbaceous plants  /  nitrogen deposition  /  critical load
许增美慧, 石兆勇, 王浩, 李琰, 马路平, 张梦歌, 吴姗薇, 高佳凯, 王双双. 森林菌根类型对林下草本植物响应氮沉降的影响. 微生物学报, 2025 , 65 (11) : 4877 -4888 . DOI: 10.13343/j.cnki.wsxb.20250252
Zengmeihui XU, Zhaoyong SHI, Hao WANG, Yan LI, Luping MA, Mengge ZHANG, Shanwei WU, Jiakai GAO, Shuangshuang WANG. Effects of forest mycorrhizal types on responses of understory herbaceous plants to nitrogen deposition[J]. Acta Microbiologica Sinica, 2025 , 65 (11) : 4877 -4888 . DOI: 10.13343/j.cnki.wsxb.20250252
随着人类活动的激增以及工农业的迅速发展,大气氮沉降迅速增加,被认为是全球生物多样性的一个关键压力源[1]。尽管氮沉降为植物提供了一种新型的肥料来源,但过量的氮沉降会导致土壤富营养化、叶片营养富集、病虫害增加[2-3],进而改变生态系统的结构和功能。为了评估大气沉积对生态系统的影响,生态学家们提出了“临界负荷”这一概念[4]。临界负荷被定义为“基于定量法确定污染物的暴露阈值时,当污染物的暴露水平低于该阈值不会对环境中的敏感要素产生显著的有害影响”[5]。简单来说,临界负荷是特定有害物质开始产生影响的定量阈值。近年来,研究人员开始重点关注生物多样性的临界负荷[5],通常基于物种出现的频率来评价生境中植物物种多样性的变化。在本研究中,临界负荷主要指因大气氮沉降而引起林下草本植物物种丰度产生变化的临界点。
草本植物对氮沉降具有较高的敏感性,其物种丰度和多样性会受到显著影响[6]。例如,英国洛桑试验站自1856年起开展的长期定点试验数据表明,氮素的长期添加会降低草本植物的多样性[7]。Walter等[8]的研究结果同样指出,长期氮添加使草本植物丰度降低了13.6%。随着氮素有效性的提高,生态系统中部分本地植物物种通常会被具有更高生产力的外来物种所取代[1]。这进一步表明植物群落的组成及物种丰度的变化可能会导致植物多样性的丧失[9]。此外,生物因素在草本植物丰度变化中的作用也不容忽视。菌根真菌广泛分布于全球陆地生态系统[10-11],与约92%的陆地植物形成共生关系[12],并在植物微生物群中起着至关重要的作用[13-14]。其中,丛枝菌根真菌和外生菌根真菌是自然界中分布最广的2种菌根真菌,它们与植物互利共生,形成丛枝菌根(arbuscular mycorrhiza, AM)和外生菌根(ectomycorrhiza, ECM)[15-16]。菌根真菌通过提供养分和构建菌丝网络机制增强植物的存活率,并进一步影响植物群落的多样性与结构[17-18]。Guy等[17]的研究表明,菌根介导作用是林下草本植物物种丰度的重要驱动因素。Zhang等[19]同样指出,菌根真菌共生通过促进植物多样性来影响有机碳储量。作为与草本植物关系密切的上层木本植物,其菌根类型也会对林下草本植物的丰度产生影响[20]。巢林等[21]研究证明,不同菌根类型树种的叶凋落物养分差异会对土壤生态产生影响,这也会影响林下草本植物的物种丰度及多样性。高旭硕等[22]研究指出,不同菌根类型对林下草本植物的物种丰富度具有显著影响(P<0.05)。此外,该研究还证实了氮沉降作为环境因素对草本植物丰富度的影响。Yuan等[23]的研究也指出,菌根类型会影响植物对氮沉降的响应。值得注意的是,高旭硕等[22]的研究缺乏对菌根类型影响森林草本植物氮沉降临界负荷机制的研究。基于此,探讨森林草本植物受氮沉降影响的临界负荷,分析在长期氮沉降背景下菌根类型如何介导森林草本植物的氮沉降临界负荷,以及不同菌根类型对草本植物物种丰度的响应差异极为重要。
本研究对不同菌根类型下森林草本植物的氮沉降临界负荷进行研究,探究不同菌根类型森林之间林下草本植物的氮沉降临界负荷是否存在差异,以及菌根类型如何影响林下草本植物对氮沉降的响应,以期为后续研究不同菌根类型林下草本植物群落响应气候变化提供数据支撑,为全球气候变化下保护林下草本植物多样性提供理论参考。
本研究基于Wilkins等[5]发表的森林草本植物长期氮沉降临界负荷数据库,该数据库包含1 800种林下草本植物(共计3 592个个体)的丰度变化及氮沉降临界负荷数据。数据库中的草本植物氮沉降临界负荷是通过指示种阈值分析法(threshold indicator taxa analysis, TITAN)确定的。TITAN使用指示物种分数(IndVal分数)作为评估物种分布变化的指标,IndVal分数综合了物种的丰度和相对频率,能够反映物种在特定环境条件下的分布特征。随后,TITAN采用非参数变化点分析确定物种的分布变化点,识别在氮沉降梯度上物种分布发生显著变化的位置,即物种的氮沉降临界负荷,并确定每个物种的丰度对氮沉降的响应格局。
为研究不同菌根类型下林下草本植物氮沉降临界负荷的差异,针对原数据中森林的优势种,根据已发表的相关文献补充其菌根类型数据。同时,参考Vargas等[24]和石兆勇等[25]对不同菌根类型森林的划分方法,以优势种的菌根类型代表森林群落的菌根类型。当森林中的优势种不止一种菌根类型,或者该群落中存在共优种时,则将几种菌根结合的类型代表森林群落的菌根类型。本研究的森林植被类型涉及2种菌根,即AM和ECM。根据上述方法,最终确定原数据中所有森林群落的菌根类型,将森林群落的菌根类型分为AM型、ECM型以及AM+ECM型,并建立新的数据库“不同菌根类型森林林下草本植物对氮沉降响应的临界负荷数据库”。该数据库包含619个AM型森林林下草本植物、2 445个ECM型森林的林下草本植物和528个AM+ECM型森林的林下草本植物。
本研究使用SPSS 27.0软件分析森林草本植物物种的氮沉降临界负荷在不同菌根类型下的差异。对于呈正态分布的数据采用单因素方差分析(one-way ANOVA);对于不呈正态分布的数据采用非参数检验(Kruskal-Wallis)[26]。在显著水平P<0.05下,分析不同菌根类型间林下草本植物氮沉降临界负荷的差异性。基于SPSS的分析结果,使用Origin 2024进行作图,并用直方图表述数据的分布状况。
根据直方图,3种菌根类型森林草本植物的氮沉降临界负荷分布存在差异(图1)。AM森林呈三段式分布,主要分布在3、7、12 kg N/(ha·a);ECM森林呈两段式分布,氮沉降阈值分布在 3、10 kg N/(ha·a);AM+ECM森林则呈一段式,分布在9 kg N/(ha·a)。从直方图可看出数据分布不符合正态分布,因此采用非参数检验对3种菌根类型森林草本植物的氮沉降临界负荷进行分析,结果表明它们之间存在显著差异(P<0.05) (图2)。AM+ECM森林林下草本植物的氮沉降阈值最高,为9.28 kg N/(ha·a),显著高于AM森林[7.19 kg N/(ha·a)]和ECM森林[8.41 kg N/(ha·a)]。
将草本植物分为禾本科和非禾本科植物,分析其不同菌根类型的氮沉降临界负荷。结果表明,无论是禾本科还是非禾本科植物,在不同菌根类型影响下的氮沉降临界负荷分布趋势一致。AM森林均为三段式分布,其阈值分布在3、7、11 kg N/(ha·a) (禾本科)和3、7、12 kg N/(ha·a) (非禾本科);ECM森林为两段式分布,分别 在3、9 kg N/(ha·a) (禾本科)和3、12 kg N/(ha·a) (非禾本科);AM+ECM森林则为一段式分布,禾本科主要分布在11 kg N/(ha·a),非禾本科分布在9 kg N/(ha·a) (图3)。非参数检验结果显示,不同菌根类型森林下,禾本科和非禾本科植物的氮沉降临界负荷呈现显著差异(P<0.05) (图4),均为AM+ECM型森林的氮沉降临界负荷[禾本科植物为9.30 kg N/(ha·a),非禾本科植物为9.37 kg N/(ha·a)]显著高于AM和ECM,AM型森林林下草本植物的临界负荷最低,分别为6.82 kg N/(ha·a) (禾本科)、7.24 kg N/(ha·a) (非禾本科)。
三种菌根类型林下草本植物受氮沉降影响的丰度变化,其阈值分布情况一致(图5)。无论丰度降低还是增加,AM森林的分布均呈现三段式,临界负荷主要分布在2、7、11 kg N/(ha·a) (丰度降低)和3、6.5、12 kg N/(ha·a) (丰度增加);ECM型呈两段式,丰度降低的氮沉降临界负荷主要分布在2.5 kg N/(ha·a)和9 kg N/(ha·a),丰度增加则主要分布在3 kg N/(ha·a)和 12 kg N/(ha·a);AM+ECM森林林下草本植物丰度变化的氮沉降临界负荷阈值分别主要集中在9 kg N/(ha·a) (丰度降低)和11 kg N/(ha·a) (丰度增加)。针对草本植物不同的丰度变化,3种菌根类型之间具有差异(图6)。对于草本植物丰度降低的阈值,Kruskal-Wallis检验结果表明3种菌根类型之间无显著差异。对于草本植物丰度增加的氮沉降临界负荷,AM+ECM型森林的林下草本植物为10.13 kg N/(ha·a),显著高于其他2种菌根类型(P<0.05)。将草本植物分为禾本科和非禾本科,受氮沉降影响物种丰度降低的禾本科植物,在不同菌根类型森林之间无显著差异,而对于非禾本科植物,ECM森林植物的氮沉降临界负荷显著高于AM森林植物(P<0.05) (图7A)。此外,不同菌根类型森林下无论是禾本科还是非禾本科植物,受氮沉降影响物种丰度增加的氮沉降临界负荷均表现为AM+ECM森林>ECM森林>AM森林(图7B)。
不同菌根类型森林的林下草本植物的丰度变化占比情况不同,其中AM森林的林下草本植物受氮沉降影响的丰度变化主要以增加为主;ECM和AM+ECM森林的草本植物则以丰度降低为主(图8)。
本研究基于长期氮沉降研究试验所建立的数据库,从森林树木不同菌根类型的视角探讨了不同菌根类型优势种森林下草本植物响应氮沉降的临界负荷及其影响因素。研究表明氮沉降能够引起不同菌根类型森林林下草本植物物种多样性的不同响应[22]。本研究进一步探究了不同菌根类型森林中草本植物对氮沉降的承受能力,并评估了其因氮沉降而导致物种丰度增加或降低的氮沉降阈值,为准确评估林下草本植物对氮沉降的响应提供了数据支撑,也为全球气候变化生态学提供了理论参考。
不同菌根类型,特别是AM与ECM植物群落生态功能的差异研究一直是学者聚焦的热点科学问题[27-28]。本研究发现,AM和ECM森林林下草本植物对氮沉降的响应具有显著差异(P<0.05),这与以往针对2种菌根类型植物在生存环境[29]、凋落物[30]以及氮获取策略[31]等方面的研究结果一致。本研究结果的具体差异表现为:AM森林林下草本植物的氮沉降临界负荷显著低于ECM森林林下草本植物(P<0.05),造成这一结果可能有以下3个原因。(1) 不同菌根类型植物的生态位分化差异所致。因为AM植物更倾向于生存在氮丰富的生境中,而ECM植物则倾向于生存在氮缺乏的地区[29],这表明AM林下土壤的氮含量本身就高于ECM林下土壤。在氮沉降影响下,AM森林林下草本植物比ECM森林林下草本植物更容易达到氮饱和状态。(2) AM和ECM菌根类型不同引起的凋落物氮浓度不同所致。Yuan等[30]研究表明,AM树种凋落叶的氮浓度显著高于ECM树种的凋落叶氮浓度(P<0.001),这使得AM森林林下土壤的氮含量高于ECM森林。陈亮等[32]研究证明,AM森林的硝态氮含量是ECM森林的2倍以上。Lin等[33]的研究也指出,AM森林中表征无机氮有效性的指标均高于ECM森林。因此,AM森林林下草本植物的氮沉降临界负荷低于ECM森林林下草本植物的氮沉降临界负荷。(3) AM和ECM帮助植物获取氮的策略不同所致。研究证明AM植物相对于ECM植物更容易从土壤中获取无机氮[34],而ECM真菌不仅能利用无机氮,还可以通过分泌氧化酶分解土壤中的有机氮,将部分氮固定在土壤有机质中[31]。因此,AM林下草本植物在氮沉降背景下更容易达到氮饱和,而ECM林下草本植物则由于养分缓冲机制从而延缓氮饱和。不同菌根类型森林林下草本植物对氮沉降响应的状态不同,也会导致不同菌根类型林下草本植物氮沉降临界负荷的差异。在本研究中,AM+ECM森林林下草本植物的氮沉降临界负荷在3种菌根类型森林中最高。这是不同菌根类型林下草本植物在氮沉降影响下不同频率分布导致的。具体而言,AM和ECM林下草本植物表现出多峰分布特征,其中AM林下草本植物表现为3个分布特征,ECM林下草本植物表现为2个分布特征,这表明2种菌根类型森林林下草本植物群落在氮沉降的影响下可能具有多个稳定状态[35]。相比之下,AM+ECM林下草本植物受氮沉降影响只表现出一个分布特征,表明AM+ECM森林林下草本植物群落对氮沉降的响应更稳定。此外,AM和ECM的协同作用也会使AM+ECM林下草本植物对氮沉降的忍耐力更强。菌根真菌可以协同调控根际土壤生态化学计量比[36],增强AM+ECM森林林下草本植物对氮沉降的适应性。AM真菌通过菌丝网络扩大宿主植物对磷的吸收范围[37],而ECM真菌则通过分泌胞外酶促进有机氮矿化能力[38]并活化难溶性磷库[39],二者共同维持土壤的N:P平衡。同时,ECM真菌能够从宿主植物中获得更多的碳,将更多碳储存在土壤中[40],并通过自身的代谢功能调节土壤中的C:N比。因此,在长期大气氮沉降背景下,AM和ECM的协同作用可以调节C:N:P的土壤化学计量平衡,进而增强根际微生态环境对氮沉降的适应能力[36],最终使得AM+ECM森林草本植物的氮沉降临界负荷高于单一菌根类型森林的草本植物。除此之外,不同菌根类型森林中不同功能群草本植物(禾本科和非禾本科)对氮沉降响应的临界负荷状况与不同菌根类型林下全部草本植物的反应完全一致。可见,植物功能群对林下草本植物响应氮沉降无显著影响。这一结果有力地支持了前人关于菌根类型对不同植物功能群研究方面的结果,Shi等[41]研究表明,在不同菌根类型下不同功能群植物的单位面积叶片质量不存在显著差异。当然,也有大量研究表明菌根类型在不同植物功能群中发挥着显著不同的作用,例如Wu等[42]研究发现在不同菌根类型中落叶植物、常绿植物以及阔叶植物的衰老叶片C含量存在显著差异(P<0.01)。之所以在本研究中不同菌根类型林下草本植物功能群之间无差异,可能是由于未能根据植物功能群对林下草本植物进行更精准划分导致的。鉴于此,在今后的研究中可以对林下草本植物进行更为细致的功能群划分,以更深入地探究不同菌根类型下不同功能群林下草本植物对氮沉降的响应。
大气氮沉降会导致林下草本植物的物种丰度发生变化,这一过程会受到菌根类型的影响。本研究发现,AM森林草本植物受氮沉降影响物种的丰度主要呈增加趋势,含ECM优势种森林的草本植物物种丰度则主要呈降低趋势。氮沉降会增强物种间的竞争[43],导致森林中氮敏感物种逐渐减少,物种丰度降低。相比之下,AM主要帮助植物吸收磷,当土壤中的氮增加时AM有助于植物更好地维持氮磷平衡[44],帮助植物在氮丰富的环境中更好地生长。基于大尺度的生态学研究指出,大气氮沉降会使陆地上植物群落的菌根类型从ECM向AM转变[13,45-46]。在大气氮沉降的条件下,随着土壤中氮的增加,ECM真菌帮助宿主植物吸收氮的生态功能被削弱,一些草本植物不需要通过ECM真菌来获取氮元素。植物生长的磷限制被放大导致植物更倾向于与AM真菌共生从而缓解自身磷限制[47-48],进而导致AM森林的林下草本植物物种丰度增加,而含ECM森林的林下草本植物物种丰度降低。
本研究探讨了不同菌根类型森林草本植物氮沉降临界负荷之间的差异,以及在菌根类型介导下草本植物物种丰度对氮沉降的响应。本研究所采用的数据涵盖了美国24个森林群落,并且试验时间从1985年持续到2012年,数据较为综合且时期较长,并以菌根类型为关键因素分析其对林下草本植物响应氮沉降的直接影响,为今后探究林下草本植物如何应对气候变化提供了科学依据和理论支撑。未来研究可通过补充实验和多变量分析,加入相关生物和非生物等驱动因素,进一步完善不同菌根类型森林草本植物对氮沉降响应机制的研究。
本研究基于大尺度分析了不同菌根类型森林草本植物氮沉降临界负荷之间的差异,以及不同菌根类型森林草本植物物种丰度对氮沉降的响应。研究发现菌根类型对林下草本植物氮沉降临界负荷有显著影响,具体表现为ECM森林草本植物的氮沉降临界负荷显著高于AM森林草本植物(P<0.05)。然而,AM+ECM森林草本植物的氮沉降临界负荷最高,说明二者协作可以加强宿主植物对长期氮沉降的适应能力。此外,氮沉降对林下草本植物物种丰度的影响因菌根类型而异,AM森林林下草本植物的物种丰度增加,ECM和AM+ECM森林林下草本植物的物种丰度减少。
许增美慧:数据处理、数据分析、制作图表和稿件撰写;石兆勇:稿件选题、研究思路和稿件修改;王浩:数据处理方法、图表制作方法、稿件修改和校对;李琰:图表制作方法、稿件修改和校对;马路平:图表制作方法和稿件修改;张梦歌:数据处理方法和稿件修改;吴姗薇:图表制作方法、稿件修改和参与论文讨论;高佳凯:稿件修改和参与论文讨论;王双双:稿件修改。
作者声明不存在任何可能会影响本文所报告工作的已知经济利益或个人关系。
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2025年第65卷第11期
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doi: 10.13343/j.cnki.wsxb.20250252
  • 接收时间:2025-03-28
  • 首发时间:2025-11-10
  • 出版时间:2025-11-04
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  • 收稿日期:2025-03-28
  • 录用日期:2025-05-21
基金
Central Plains Science and Technology Innovation Leading Talents Program(254200510006)
中原科技创新领军人才项目(254200510006)
Key Research and Development Program of Hainan Province(ZDYF2024XDNY172)
海南省重点研发项目(ZDYF2024XDNY172)
Scientific and Technological Research Projects in Henan Province(252102320074)
河南省科技攻关项目(252102320074)
Scientific and Technological Research Projects in Henan Province(252102110191)
河南省科技攻关项目(252102110191)
作者信息
    1 河南科技大学 农学院,河南 洛阳
    2 河南省乡村人居环境工程中心,河南 洛阳
    3 洛阳市共生微生物与绿色发展重点实验室,河南 洛阳

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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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