叶片凋落物分解的主场优势研究进展
Research progress on home-field advantage of leaf litter decomposition
通讯作者: *刘峰: ORCID:0000-0003-3383-7598(liufeng@wbgcas.cn)
编委: 孙建新
责任编辑: 赵航
收稿日期: 2022-04-14 接受日期: 2022-09-8
基金资助: |
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Received: 2022-04-14 Accepted: 2022-09-8
Fund supported: |
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凋落物在原生生境(“主场”)中比在非原生生境(“客场”)中分解得更快的现象被称为凋落物分解的“主场优势”。探究凋落物分解的主场优势的主要影响因素及驱动机制对预测植物养分的归还过程和生态系统碳收支有重要意义。该文主要从主场优势的计算方法、影响因素及驱动机制出发, 综述了近年来凋落物分解的主场优势的研究进展, 并对未来的研究方向进行了展望。度量凋落物分解的主场优势有4种常见的计算方法, 其中采用线性模型计算主场优势在当前最为合适。凋落物质量(化学成分等)、土壤微生物群落结构是影响凋落物分解的主场优势的主要因素, 土壤动物、气候条件、分解时间、植物生活型及生长型也能改变主场优势的强度。凋落物之间质量差异越大, 产生的主场优势越大。土壤微生物群落驱动着凋落物分解的主场优势, 但其作用时常受到动物的干扰及气候的制约。此外, 带有叶际微生物的凋落物比去除了叶际微生物的凋落物有更强的主场优势。凋落物化学性质趋同假说、分解者控制假说及凋落物质量与环境相互作用假说是解释主场优势产生的主要假说, 但它们均有不足之处。该文认为凋落物和土壤微生物的协同作用可能是产生和驱动主场优势的主要机制。当前的研究存在着各因素对主场优势的影响探究不够深入、关注的生态系统类型较为单一等问题, 在未来的研究中需要进一步深入探究各因素对主场优势效应的影响及其相对贡献, 关注更多不同的生态系统类型, 从而增强对主场优势相关机制的理解。
关键词:
Litter often decomposes more rapidly in its native habitat (“home”) than in non-native habitats (“away”), a phenomenon called the “home-field advantage”. To explore the driving mechanism of home-field advantage of litter decomposition is important to predict the process of plant nutrient return and ecosystem carbon budget. This study reviewed the research progress on the home-field advantage of litter decomposition in recent years by discussing the quantification of home-field advantage, the controlling factors, and related driving mechanisms. There are four common metrics to describe home-field advantage in litter decomposition, and the use of linear model analysis to calculate home-field advantage is more appropriate. Litter quality (chemical composition, etc.) and soil microbial community structure are the main factors influencing the home-field advantage of litter decomposition, and soil fauna, climatic conditions, decomposition time, plant life form and growth form can also influence the intensity of the home-field advantage. Greater differences in litter quality usually generate stronger home-field advantage. Microbial taxa in the soil drive the home-field advantage of litter decomposition, but the role of soil microbes is often mediated by animal and climatic disturbances. In addition, the existence of phyllosphere microbes makes the home-field advantage of litter decomposition stronger. The litter chemical convergence hypothesis, decomposer control hypothesis and substrate quality-matrix quality interaction hypothesis are major hypotheses explaining the home-field advantage in litter decomposition, but they are not impeccable. We believe that the association between litter and soil microbial community is the driving force behind home-field advantage. The current researches on the factors and relative contribution of home-field advantage are not deep enough and usually focusing on a single ecosystem. Future investigations should explore deeper on the factors and their relative contributions of home-field advantage, and focus on more ecosystem types to improve the understandings of the mechanism of home-field advantage.
Keywords:
引用本文
赵小祥, 朱彬彬, 田秋香, 林巧玲, 陈龙, 刘峰.
ZHAO Xiao-Xiang, ZHU Bin-Bin, TIAN Qiu-Xiang, LIN Qiao-Ling, CHEN Long, LIU Feng.
凋落物分解是调节生态系统碳平衡和养分循环的一个基本生态过程, 也是土壤有机质的重要来源。已有的研究发现在大部分生态系统中凋落物在其原生生境(“主场”)比在非原生生境(“客场”)分解得更快, 这一现象被称为凋落物分解的“主场优势(home-field advantage)” (Gholz et al., 2000; Prescott et al., 2000; Vivanco & Austin, 2008; Ayres et al., 2009b)。这一现象的产生主要归因于不同生境中凋落物质量(化学成分等)的差异和土壤生物对同生境植物凋落物的分解特化作用(Gholz et al., 2000; 查同刚等, 2012)。因此, 在对凋落物分解模型进行参数化的过程中, 需要弄清凋落物分解数据来源于“主场”还是“客场”。探究凋落物分解的主场优势的驱动机制和影响因素, 能为优化凋落物分解模型及预测陆地生态系统碳收支和植物养分的归还过程提供数据支撑和理论基础(St. John et al., 2011)。
在过去20多年中, 凋落物分解的主场优势备受关注, 其研究对象从单一的生态系统延伸到多类型的生态系统(Fanin et al., 2016), 关注的物种也从单一走向多元化(Ayres et al., 2009a; Milcu & Manning, 2011; Sterkenburg et al., 2018; Lin et al., 2019)。利用凋落物分解互置实验, 大多数研究发现凋落物在“主场”比在“客场”生境下分解得更快(Perez et al., 2013; Jewell et al., 2015; Keiser & Bradford, 2017; Yuan et al., 2019) (图1), 然而还有部分研究发现凋落物分解速率在“主场”和“客场”间无显著差异(St. John et al., 2011; Bachega et al., 2016), 也有少量研究结果为凋落物在“主场”生境下分解得更慢(Wang et al., 2013; Yu et al., 2015)。最新的综述研究表明在全球大多数生态系统中约70%的凋落物分解研究表现出显著的主场优势(Fanin et al., 2021)。
图1
图1
落叶阔叶林和常绿针叶林叶片凋落物分解的主场优势示意图。凋落物在原生生境(“主场”)比在非原生生境(“客场”)中分解得更快, 这种现象称为凋落物分解的“主场优势”。
Fig. 1
Schematic diagram of the home-field advantage of litter decomposition of broadleaf deciduous forests and needleleaf evergreen forests. Litter decomposes more rapidly in its native habitat (“home”) than in non-native habitats (“away”), a phenomenon called the “home-field advantage”.
1 主场优势效应的计算方法
凋落物分解的主场优势效应的度量需通过凋落物分解互置实验, 即各凋落物需同时在“主场”和“客场”进行分解实验。目前, 计算凋落物分解的主场优势的方法主要有4种。第一种方法是直接通过对比凋落物在“主场”和“客场”的分解速率差异来计算主场优势(Austin et al., 2014)。这种方法简单、快捷, 但只给出单个物种在特定“主场”和“客场”组合时的优势效应, 且忽略了“主客场”间气候条件和土壤性质的差异对分解的影响。因此, 这种方法在实例研究中较少, 计算公式为:
式中, HFA为主场优势, kin situ为凋落物在“主场”环境中的分解速率常数, kex situ为凋落物在“客场”环境中的分解速率常数。当HFA = 0时, 表示不存在主场优势; HFA > 0时, 表示正主场优势; 当HFA < 0时, 表示负主场优势。
式中, HFAi为主场优势效应指数, A、B分别表示物种, a、b为对应的生境, Aa表示物种A在地点a的质量损失, Ba表示物种B在地点a的质量损失, ARMLa表示为物种A在地点a的相对质量损失, BRMLb表示为物种B在地点b的相对质量损失, ARMLb表示为物种A在地点b的相对质量损失, BRMLa表示为物种B在地点a的相对质量损失。当HFAi = 0时, 表示不存在主场优势; HFAi > 0时, 表示正主场优势; HFAi < 0时, 表示负主场优势。利用这一计算方法, 文献中报道的凋落物分解的主场优势范围为-24.65%-56.28% (Fanin et al., 2021)。
式中, ADHi表示I物种的主场优势附加分解指数; I, J, K分别表示不同物种; i, j, k为对应的生境; D为凋落物质量损失的百分比; HDDi表示在生境i中“主场”物种与其他2种“客场”物种的分解差异之和(当分解的物种超过3种时, HDDi为在生境i中“主场”物种与其他所有“客场”物种的分解差异之和); ADDi表示物种I凋落物在“客场”与该生境下“主场”物种分解差异之和(当分解的物种超过3种时, ADDi表示物种I凋落物在“客场”与该生境下其他所有“主场”物种分解差异之和); H表示所有物种的平均主场优势效应; N表示物种数量。将ADHi与0进行t检验, 如果p < 0.05, 表示产生主场优势, 相反则未产生主场优势。在p < 0.05的条件下, ADHi = 0, 表示物种I不存在主场优势; ADHi > 0, 表示物种I存在正主场优势; ADHi < 0, 表示物种I存在负主场优势。根据当前已有的研究结果, 文献中报道的ADHi范围为-10%-10%。
式中, Yi为凋落物分解质量剩余百分比; α为在控制凋落物、生境和主场优势配对之后, 所有观察到的分解速率的平均值; βl表示凋落物l的影响;
2 主场优势效应的影响因素
2.1 凋落物质量的影响
凋落物质量(化学成分等)控制着凋落物分解速率, 同时也是影响主场优势效应最重要的因素之一 (Ayres et al., 2009b; Vauramo & Setälä, 2011)。多项研究发现凋落物易分解的成分含量越高, 凋落物分解的主场优势越强(Veen et al., 2018; Lin et al., 2020)。例如, 在夏威夷热带雨林中, 拥有高氮、磷含量和低木质素、可溶性多酚含量的白蜡树(Fraxinus uhdei)凋落物的分解速率和主场优势效应显著高于Metrosideros polymorpha (Rothstein et al., 2004)。然而, 有研究却认为凋落物易分解的成分含量越低, 其分解的主场优势越强(Milcu & Manning, 2011; Austin et al., 2014)。这可能归因于凋落物中难降解的化学物质(如木质素、单宁、萜类化合物和酚醛类物质)需要专一化程度更高的微生物群落进行分解(Milcu & Manning, 2011)。如果“客场”环境中缺少这些特定类群的微生物, 凋落物的分解速率就会降低。因此, 低质量凋落物的分解速率在“主场”和“客场”之间就更容易出现差异, 展现出更强的主场优势(Strickland et al., 2009; Milcu & Manning, 2011; Austin et al., 2014; 立天宇等, 2015; Yeung et al., 2019)。例如, Chomel等(2015)在对Picea glauca和辽杨(Populus maximowiczii)的凋落物互置实验中发现只有低质量的Picea glauca凋落物表现出明显的主场优势。可见, 凋落物自身质量对主场优势的影响尚无定论。
除凋落物自身质量(化学成分等)外, “主场”和“客场”凋落物质量的差异, 也可影响主场优势的大小(Veen et al., 2015a; Li et al., 2017)。在农田、森林及草地生态系统开展的凋落物分解互置实验中均发现凋落物间的质量差异越大, 凋落物分解的主场优势越高(Aponte et al., 2012; 立天宇等, 2015; Fanin et al., 2016; 杨红玲等, 2019)。Hobbie等(2006)对波兰14种常见树种的凋落物分解实验证明了凋落物的主场优势主要受“主客场”凋落物间木质素含量差异的调控。这些研究结果与Freschet等(2012)提出的凋落物质量与环境相互作用假说(SMI, substrate quality-matrix quality interaction)相一致。即“客场”凋落物质量与“主场”凋落物质量之间的差异越大, “客场”凋落物分解的速率越低, 进而产生更大的主场优势。这一假说得到相关研究的支持(Veen et al., 2015a; Li et al., 2017), 但也有研究并未证实这一假说(Perez et al., 2013; Jewell et al., 2015)。可见, “主客场”凋落物质量的差异对主场优势的影响也存在争议。因此, 在将来开展凋落物分解主场优势研究时, 可选择多个凋落物质量成梯度变化的物种进行凋落物分解互置实验, 以进一步明确凋落物质量之间的差异对主场优势的影响。
2.2 土壤生物的影响
2.2.1 土壤微生物
土壤微生物是影响凋落物分解的主场优势的重要因素(de Toledo Castanho & de Oliveira, 2008; 查同刚等, 2012; 立天宇等, 2015; 杨红玲等, 2019; Li et al., 2020)。这归因于凋落物和土壤微生物间存在长期相互作用, 凋落物与其相对应的土壤微环境之间建立了亲和效应(Vivanco & Austin, 2008)。因此, 土壤中的微生物分解凋落物具有专一性(Ayres et al., 2009b; Kagata & Ohgushi, 2013)。本地土壤微生物群落可快速定植在本地凋落物上并对某些特定化学组分形成偏好利用(Lin et al., 2019), 尤其是木质素(Negrete-Yankelevich et al., 2008)。Veen等(2019)发现凋落物分解的主场优势效应与特定的优势真菌显著相关。另外, 温带草原凋落物分解实验也发现小尺寸的真菌和细菌黄丝孢目(Xanthomonadales)与皮丝孢酵母(Cutaneotrichosporon)可分解菊蒿(Tanacetum vulgare)和疆千里光(Jacobaea vulgaris)凋落物的特定成分, 从而促进主场优势(Li et al., 2020)。然而, 当凋落物被放置在“客场”时, “客场”土壤微生物群落和凋落物之间缺乏亲和关系, 其分解速率会降低(Palozzi & Lindo, 2017)。
尽管土壤微生物在凋落物分解产生主场优势的过程中扮演着重要角色, 但有部分研究认为土壤微生物群落调节主场优势的作用有限(Fanin et al., 2016)。这是由于微生物具有功能冗余性, 且微生物能够在数周或者数月内迅速适应新的凋落物输入, 降低其对主场优势效应的影响(Gießelmann et al., 2011; St. John et al., 2011; Austin et al., 2014; Lin et al., 2019)。例如, Li等(2020)发现土壤微生物群落对主场优势的影响只发生在分解实验的前6个月。除此之外, 林下昆虫和食草动物的活动使土壤微生物难以形成和维持特定的群落, 这削弱了土壤微生物对凋落物分解的主场优势的影响(Kagata & Ohgushi, 2013)。而且气候条件也可掩盖土壤微生物群落对凋落物分解的独立影响。如恶劣的气候能限制微生物活动来削弱主场优势效应(Keiser & Bradford, 2017)。可见, 土壤微生物对凋落物分解主场优势的影响还有待明确, 未来需进一步明确土壤微生物与环境条件的相互作用对主场优势的影响。
2.2.2 土壤动物
土壤动物通过破碎、摄食等行为影响凋落物分解, 也是调节主场优势的重要因素(Hättenschwiler et al., 2011), 尤其是在凋落物量大的生态系统中(Milcu & Manning, 2011)。相对于土壤微生物, 土壤动物尤其是土壤中的螨虫对栖息地的要求更加专一化, 当凋落物主客场发生变化时, 它可增强对凋落物分解的影响, 进而改变凋落物分解的主场优势(St. John et al., 2011; 查同刚等, 2012)。针对14种常见树种凋落物分解实验发现, 土壤中的蚯蚓群落决定凋落物的分解速率及主场优势效应(Hobbie et al., 2006)。然而, 土壤动物也可能对土壤微生物进行选择性取食(Bardgett et al., 1993), 降低土壤微生物对凋落物分解的影响(Newell, 1984; Bradford et al., 2002; Crowther et al., 2012), 削弱土壤微生物群落差异引起的主场优势。但Lin等(2019)在亚热带森林开展的凋落物分解实验发现, 土壤中的大型或中型动物对主场优势无显著影响。当前, 关于土壤动物对凋落物分解主场优势的影响还存在争议, 相关的机制有待进一步明确。
2.3 叶际微生物的影响
叶际微生物是生活在植物叶片表面和内部的细菌、真菌等微生物类群(Lindow & Brandl, 2003), 它在促进植物生长、防止病原菌侵害以及植物碳氮循环中起着重要的作用(Laforest-Lapointe et al., 2017; Liu et al., 2020)。近年来有研究发现叶际微生物在凋落物分解的早期起着重要的作用(Fanin et al., 2021)。叶片在即将凋落时通常会释放大量的易降解化合物(Ibrahima et al., 1995), 叶际微生物可利用这些养分并在叶片表面和内部生长繁殖(Osono, 2006; Vacher et al., 2016), 成为凋落物早期分解的加速者。除此之外, 部分特定的叶际微生物可以分解凋落物中的特定成分(Wolfe & Ballhorn, 2020)。有研究表明木樨科植物的内生菌能够分解其宿主叶片中的木质素, 从而提高叶片凋落物分解速度(Osono & Takeda, 1999; Osono, 2002)。凋落物落到地表后, 其表面的叶际微生物群落会逐渐被土壤微生物群落替代, 对凋落物分解的影响减弱(图2)。
图2
图2
凋落物分解过程中叶际微生物和土壤微生物的相对变化示意图(改自Fanin等(2021))。在叶片掉落前夕(①), 叶际微生物快速生长, 然后掉落在地表(②); 在凋落物分解早期(③), 叶际微生物和土壤微生物共存; 在凋落物分解中晚期(④), 土壤微生物占主导。
Fig. 2
Schematic diagram of the relative changes of phyllosphere microbes and soil microbes during litter decomposition (adapted from Fanin et al. (2021)). On the eve of leaf drop (①), phyllosphere microbes grew and colonized rapidly and then leaf litter fell to the ground (②); in the early stage of litter decomposition (③), phyllosphere microbes and soil microbes coexisted; in the middle and late stages of litter decomposition (④), soil microbes dominated.
叶际微生物不仅能提高叶片凋落物的分解速率, 也能显著增强叶片凋落物分解的主场优势(Fanin et al., 2021)。相对于灭菌处理的叶片凋落物, 未灭菌的凋落物主场优势更高, 其增幅达8.2% (Fanin et al., 2021)。叶际微生物对凋落物早期的快速分解能在调节土壤微生物和土壤动物引起的主场优势效应中发挥关键作用。一方面, 叶际微生物在凋落物分解的早期可快速生长繁殖, 改变微生物活动的生态位(Fanin et al., 2021), 从而影响土壤微生物群落在凋落物上的定植和后续演替(Voříšková & Baldrian, 2013; Fukami, 2015)。叶际微生物可优先利用易分解的凋落物成分, 与能利用这些易分解成分的土壤微生物类群形成竞争, 限制它们的定植和生长。根据资源互补效应, 能利用难分解成分的土壤微生物类群的生长则可得到显著的促进, 从而加速叶片凋落物的分解, 进而增强主场优势。另一方面, 叶际微生物在凋落物中的早期分解作用也可调节土壤中大型和中型动物对凋落物的分解(Gessner et al., 1999), 潜在影响主场优势的强度。此外, 叶际微生物与环境条件存在长期的适应过程, 在自身适应的“主场”环境中加速凋落物的分解, 而在“客场”环境中对凋落物分解的影响减弱(Strickland et al., 2015), 进而引起主场优势。但是, 叶际微生物在不同环境间适应性的差异可能会随着时间延长而减弱, 从而降低对主场优势的影响。综上, 叶际微生物可通过多种途径影响凋落物分解的主场优势, 但是各途径的作用强度和相对贡献还不明晰。
2.4 气候条件的影响
气候条件是影响凋落物分解的主要因素(Parton et al., 2007; Zhang et al., 2008), 也可影响凋落物分解的主场优势。在瑞典北部的亚北极地区沿着海拔梯度开展的凋落物分解实验中, Veen等(2015b)发现气温影响凋落物分解的主场优势, 且主场优势随着气温升高而增加。除此之外, 气温和降水能显著改变土壤微生物和动物的活动, 这将改变它们对凋落物的分解过程, 进而间接影响主场优势(Gießelmann et al., 2011)。虽然现有的研究认为凋落物分解的主场优势在全球大多数生态系统中广泛存在(Veen et al., 2015a), 但气候条件, 尤其是温度或降水对主场优势的影响及其作用机理还知之甚少。
2.5 其他影响因素
凋落物分解的主场优势有时也取决于凋落物分解阶段(Ayres et al., 2009b; Keiser et al., 2014; Fanin et al., 2016; Veen et al., 2018)。Lin等(2020)在亚热带森林中开展的凋落物分解实验发现毛竹(Phyllostachys edulis)凋落物的主场优势仅出现在分解早期(第3-7个月), 杉木(Cunninghamia lanceolata)凋落物在第7个月才显示显著的负主场优势, 且毛竹和杉木凋落物均是随着分解阶段先出现正主场优势再出现负主场优势, 而甜槠(Castanopsis eyrei)凋落物在所有的分解阶段均为正主场优势。同样, Ayres等(2009a)在室内针对Populus tremuloides、Pinus contorta和Picea engelmannii凋落物的互置分解实验发现, Pinus contorta的主场优势随着分解时间先不断增加然后趋于稳定, Populus tremuloides的主场优势在分解的第4-6个月中保持高峰水平, 然后开始下降, 而Picea engelmannii却没表现出类似规律。随着凋落物分解时间的延长, 凋落物化学特征及微生物群落的改变可能是造成主场优势随分解阶段变化的重要原因(Ayres et al., 2009a; Lin et al., 2020)。
3 主场优势效应的产生机制
针对叶片凋落物分解的主场优势这一生态学现象, 有多个假说被提出来解释它的产生机制, 其中主要有以下三种假说:
凋落物分解的主场优势受到了土壤微生物的驱动(de Toledo Castanho & de Oliveira, 2008; 查同刚等, 2012; 立天宇等, 2015; 杨红玲等, 2019; Li et al., 2020)。因此, 分解者控制假说被提出。该假说认为“主场”的土壤微生物适应了其对应的植物凋落物, 是分解者控制了主场优势的大小(Wardle et al., 2004; Ayres et al., 2009b)。因为“主场”的土壤微生物分解凋落物具有专一性(Ayres et al., 2009b; Kagata & Ohgushi, 2013), 它能快速定植本地凋落物(Lin et al., 2019), 同时也能快速分解凋落物中难以分解的成分(Negrete-Yankelevich et al., 2008)。这就使得凋落物在“主客场”互置时, 凋落物在“主场”的分解速率显著高于“客场”, 造成主场优势(Palozzi & Lindo, 2017)。
虽然凋落物化学性质趋同假说、分解者控制假说及凋落物质量与环境相互作用假说都能在某种程度上解释主场优势的产生和驱动机制, 但它们仍不完善。凋落物分解的快慢不仅取决于凋落物质量(化学成分等), 也取决于“主场”环境中特定土壤微生物类群的作用。因为凋落物中难降解的化学物质需要较为专一化的微生物类群进行分解(Milcu & Manning, 2011)。在实际研究中如果仅考虑凋落物质量, 而不考虑其与“主场”土壤微生物群落的协同作用, 得到的结果就可能存在一定的偏差。因此, 我们认为凋落物分解的主场优势可能主要是由凋落物和土壤微生物群落的协同作用所驱动。凋落物质量和土壤微生物群落两者缺一不可, 对它们两者的协同作用进行深入研究能更全面地揭示凋落物分解的主场优势的产生和驱动机制。
4 总结和展望
除气候条件、凋落物质量(化学成分等)和分解者特征外, 凋落物分解的主场优势已成为影响凋落物分解的重要因素。尽管近些年对凋落物分解的主场优势开展了大量研究, 但仍然存在着一些问题和挑战。因此, 针对当前主场优势研究存在的不足, 本文提出相关的解决方案,并且指出了未来需要重点关注的发展方向。
4.1 进一步深入探究相关因素对主场优势的影响
凋落物分解的主场优势受到了凋落物质量(化学成分等)、土壤生物、气候条件等因素的影响, 但受实验条件、实验设计、数据量等限制因素的影响, 无法准确判断各因素对主场优势的相对贡献。为了解决上述问题, 需开展标准化的凋落物分解互置实验, 实验设计要同时考虑凋落物质量、土壤生物群落、叶际微生物等主要因素对主场优势的影响, 从而更加深入地理解各个因素对主场优势的作用机理及其相对贡献。除此, 在未来也应该开展相关的野外控制实验, 尤其是在全球变化背景下需要关注气温升高、氮沉降、降水变化等如何影响凋落物分解的主场优势。
4.2 进一步揭示叶际微生物对主场优势的影响
叶际微生物对凋落物分解及其主场优势的影响近年来被逐渐关注(Austin et al., 2014; Fanin et al., 2021), 但其作用机制和强度还需进一步明确(Fanin et al., 2021)。首先, 以往的大部分研究都采用灭菌的方法探究叶际微生物存在与否对主场优势的影响, 但灭菌过程对凋落物本身的影响往往被忽略。未来可通过对叶际微生物移植和接种等方法, 进一步明确叶际微生物对主场优势的影响。其次, 叶际微生物群落受微环境(St. John et al., 2011)、凋落物化学性质(Freschet et al., 2012)和土壤生物(Milcu & Manning, 2011)的影响。在凋落物分解实验中, 需要进一步揭示叶际微生物和其他生物及非生物因子之间的关系。最后, 叶际微生物在凋落物分解过程中的相对丰度、群落组成和功能会发生改变。因此, 在未来的研究中应从叶际微生物群落组成和功能的角度出发, 探究其在凋落物分解的主场优势中的作用机制(Voříšková & Baldrian, 2013)。
4.3 进一步拓展主场优势的研究范围
当前关于凋落物分解主场优势的大多数研究集中在热带、亚热带和温带区域, 寒带区域的研究很少(Veen et al., 2015a)。不同气候区的凋落物性质、土壤生物区系、气候条件存在显著差异, 可能会影响凋落物分解的主场优势的强度。因此, 有必要在不同气候区分别开展凋落物分解的主场优势研究, 探明不同气候条件对主场优势效应的影响。此外, 已有的关于主场优势的研究大部分集中在森林生态系统中(Ayres et al., 2009b), 而在草地、农田生态系统中关注较少(Fanin et al., 2016; Schmitt & Perfecto, 2021)。因此, 需要加强不同生态系统中凋落物分解的主场优势的研究, 进而更为全面地揭示主场优势的产生机理。最后, 当前的主场优势研究几乎都是关注于叶片凋落物, 而涉及植物根系分解的研究屈指可数。在陆地生态系统中有20% (热带雨林)到70% (温带草原)的生物量分配在根中(Poorter et al., 2012), 根系的分解是土壤有机碳和养分的重要来源。探究根系分解的主场优势, 对揭示地下生态系统的养分循环过程有重要的指导意义(Lin et al., 2020)。
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Responses of soil microbial carbon metabolism to the leaf litter composition in Liaohe River Nature Reserve of northern Hebei Province, China
冀北辽河源自然保护区土壤微生物碳代谢对阔叶林叶凋落物组成的响应
利用凋落物袋法研究了冀北辽河源地区阔叶混交林内山杨、白桦、蒙古栎叶凋落物单一分解及混合分解对0~5、5~10和10~20 cm表层土壤微生物生物量碳、微生物呼吸和微生物代谢熵的影响.结果表明: 0~20 cm土层对照、白桦、山杨和蒙古栎处理的土壤微生物生物量碳平均含量分别为124.84、325.29、349.79和319.02 mg·kg<sup>-1</sup>;微生物呼吸平均速率分别为0.66、1.12、1.16和1.10 μg·g<sup>-1</sup>·h<sup>-1</sup>.0~20 cm土层单一凋落物处理、两种叶凋落物混合处理、3种叶凋落物混合处理的土壤微生物生物量碳平均含量分别为331.37、418.52和529.34 mg·kg<sup>-1</sup>;微生物呼吸平均速率分别为1.13、1.30和1.46 μg·g<sup>-1</sup>·h<sup>-1</sup>.土壤微生物代谢熵则呈现出与微生物生物量碳、微生物呼吸相反的变化趋势.说明凋落物质量不同,其土壤微生物碳代谢特征不同,表现为高质量凋落物土壤微生物生物量碳、微生物呼吸速率以及微生物对土壤中有机质的利用效率较高,低质量凋落物则与之相反.植物叶凋落物混合能够增强土壤微生物活性,增加土壤微生物对土壤碳的利用效率,促进土壤微生物代谢途径的多样化,有利于林地土壤质量的维护和提高.
Home-field advantages of litter decomposition increase with increasing N deposition rates: a litter and soil perspective
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“Home” and “away” litter decomposition depends on the size fractions of the soil biotic community
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Home-field advantage of litter decomposition differs between leaves and fine roots
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Fungi participate in driving home-field advantage of litter decomposition in a subtropical forest
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Microbiology of the phyllosphere
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Linking the phyllosphere microbiome to plant health
DOI:S1360-1385(20)30199-0
PMID:32576433
[本文引用: 1]
The phyllosphere harbors diverse microbial communities that influence ecosystem functioning. Emerging evidence suggests that plants impaired in genetic networks harbor an altered microbiome and develop dysbiosis in the phyllosphere, which pinpoints plant genetics as a key driver of the phyllosphere microbiome assembly and links the phyllosphere microbiome to plant health.Copyright © 2020 Elsevier Ltd. All rights reserved.
All size classes of soil fauna and litter quality control the acceleration of litter decay in its home environment
DOI:10.1111/more.2011.120.issue-9 URL [本文引用: 8]
Species-specific characteristics of trees can determine the litter macroinvertebrate community and decomposition process below their canopies
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Interaction between two decomposer basidiomycetes and a collembolan under Sitka spruce: grazing and its potential effects on fungal distribution and litter decomposition
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Phyllosphere fungi on leaf litter of Fagus crenata: occurrence, colonization, and succession
DOI:10.1139/b02-028
URL
[本文引用: 1]
Phyllosphere fungi occur on various litters, but the ecology of these fungi on leaf litter has received little attention. To investigate the occurrence, colonization, and succession of phyllosphere fungi on leaf litter of Fagus crenata Blume, fungi were isolated from living, senescent, freshly fallen, and decomposing leaves by surface sterilization and washing methods. A total of 18 and 47 fungal species were isolated from the interior and surface of living and senescent leaves, respectively, and 15 frequent species were regarded as phyllosphere fungi. These fungi were divided into three groups according to their frequency on freshly fallen and decomposing leaves. Nine species (Group I) occurred frequently on decomposing leaves, two species (Group II) on freshly fallen leaves only, and four species (Group III) were frequent on living or senescent leaves only. Colonization of sterilized, freshly fallen leaves by phyllosphere fungi was investigated to test their ability to infect litter directly after litter fall. Frequencies of four species were lower on sterilized leaves than on unsterilized leaves, whereas frequencies of other species did not differ between sterilized and unsterilized leaves. Successional trends of endophytes and epiphytes were observed during decomposition from freshly fallen to decomposing leaves. The sum of frequencies of endophytes decreased temporarily on freshly fallen leaves and increased on decomposing leaves. The sum of frequencies of epiphytes decreased from freshly fallen to decomposing leaves.Key words: beech, decomposition, endophyte, epiphyte, Xylariaceae.
Role of phyllosphere fungi of forest trees in the development of decomposer fungal communities and decomposition processes of leaf litter
The ecology of endophytic and epiphytic phyllosphere fungi of forest trees is reviewed with special emphasis on the development of decomposer fungal communities and decomposition processes of leaf litter. A total of 41 genera of phyllosphere fungi have been reported to occur on leaf litter of tree species in 19 genera. The relative proportion of phyllosphere fungi in decomposer fungal communities ranges from 2% to 100%. Phyllosphere fungi generally disappear in the early stages of decomposition, although a few species persist until the late stages. Phyllosphere fungi have the ability to utilize various organic compounds as carbon sources, and the marked decomposing ability is associated with ligninolytic activity. The role of phyllosphere fungi in the decomposition of soluble components during the early stages is relatively small in spite of their frequent occurrence. Recently, the roles of phyllosphere fungi in the decomposition of structural components have been documented with reference to lignin and cellulose decomposition, nutrient dynamics, and accumulation and decomposition of soil organic matter. It is clear from this review that several of the common phyllosphere fungi of forest trees are primarily saprobic, being specifically adapted to colonize and utilize dead host tissue, and that some phyllosphere fungi with marked abilities to decompose litter components play important roles in decomposition of structural components, nutrient dynamics, and soil organic matter accumulation.
Decomposing ability of interior and surface fungal colonizers of beech leaves with reference to lignin decomposition
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Pure and mixed litters of Sphagnum and Carex exhibit a home-field advantage in Boreal peatlands
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Global-scale similarities in nitrogen release patterns during long-term decomposition
DOI:10.1126/science.1134853
PMID:17234944
[本文引用: 1]
Litter decomposition provides the primary source of mineral nitrogen (N) for biological activity in most terrestrial ecosystems. A 10-year decomposition experiment in 21 sites from seven biomes found that net N release from leaf litter is dominantly driven by the initial tissue N concentration and mass remaining regardless of climate, edaphic conditions, or biota. Arid grasslands exposed to high ultraviolet radiation were an exception, where net N release was insensitive to initial N. Roots released N linearly with decomposition and exhibited little net N immobilization. We suggest that fundamental constraints on decomposer physiologies lead to predictable global-scale patterns in net N release during decomposition.
Home-field advantage: a matter of interaction between litter biochemistry and decomposer biota
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Biomass allocation to leaves, stems and roots: meta-analyses of interspecific variation and environmental control
DOI:10.1111/j.1469-8137.2011.03952.x
PMID:22085245
[本文引用: 1]
We quantified the biomass allocation patterns to leaves, stems and roots in vegetative plants, and how this is influenced by the growth environment, plant size, evolutionary history and competition. Dose-response curves of allocation were constructed by means of a meta-analysis from a wide array of experimental data. They show that the fraction of whole-plant mass represented by leaves (LMF) increases most strongly with nutrients and decreases most strongly with light. Correction for size-induced allocation patterns diminishes the LMF-response to light, but makes the effect of temperature on LMF more apparent. There is a clear phylogenetic effect on allocation, as eudicots invest relatively more than monocots in leaves, as do gymnosperms compared with woody angiosperms. Plants grown at high densities show a clear increase in the stem fraction. However, in most comparisons across species groups or environmental factors, the variation in LMF is smaller than the variation in one of the other components of the growth analysis equation: the leaf area : leaf mass ratio (SLA). In competitive situations, the stem mass fraction increases to a smaller extent than the specific stem length (stem length : stem mass). Thus, we conclude that plants generally are less able to adjust allocation than to alter organ morphology.© 2011 The Authors. New Phytologist © 2011 New Phytologist Trust.
Decomposition of broadleaf and needle litter in forests of British Columbia: influences of litter type, forest type, and litter mixtures
DOI:10.1139/x00-097
URL
[本文引用: 1]
We measured rates of decomposition at three sites representing the major mixedwood forest types of British Columbia: (i) boreal forests of white spruce (Picea glauca (Moench) Voss) and trembling aspen (Populus tremuloides Michx.); (ii) coastal forests of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and red alder (Alnus rubra Bong.); and (iii) a wet interior forest of Douglas-fir, paper birch (Betula papyrifera Marsh.), and lodgepole pine (Pinus contorta Doug. ex Loud.). Mass loss of litter of each species (both pure and in combination with the other species) was measured for 2-5 years in forests of each species to determine (i) if broadleaf litter decomposed faster than needle litter, (ii) if litter decomposed faster in broadleaf or mixedwood forests than in coniferous forests, and (iii) if mixing with broadleaf hastened decomposition of needle litter. The broadleaf litters decomposed faster than needles during the first year but, thereafter, decomposed more slowly, so differences were small after 3 years. Litter tended to decompose faster in the broadleaf forests than in the coniferous forests. There was either no effect or a slight suppression of decomposition when litters were mixed; thus, there was no evidence that addition of broadleaf litter hastened decomposition of needle litter. The results clearly indicate that the mixing of needle litter with broadleaf litter is unlikely to hasten decomposition in mixedwood forests of British Columbia. The main influence of broadleaves was more rapid decomposition in broadleaf or mixedwood forest floors, which does not appear to be simply an effect of litter quality or litter mixing.
An exotic tree alters decomposition and nutrient cycling in a Hawaiian montane forest
DOI:10.1007/s10021-004-0009-y URL [本文引用: 1]
Coffee leaf litter decomposition: short term home-field advantage in shaded coffee agro- ecosystems
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No “home” versus “away” effects of decomposition found in a grassland- forest reciprocal litter transplant study
DOI:10.1016/j.soilbio.2011.03.022 URL [本文引用: 6]
Contrasting effects of ectomycorrhizal fungi on early and late stage decomposition in a boreal forest
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Climate history shapes contemporary leaf litter decomposition
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Litter quality is in the eye of the beholder: initial decomposition rates as a function of inoculum characteristics
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The phyllosphere: microbial jungle at the plant- climate interface
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Decomposition of labile and recalcitrant litter types under different plant communities in urban soils
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Litter quality and environmental controls of home-field advantage effects on litter decomposition
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Variation in home-field advantage and ability in leaf litter decomposition across successional gradients
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Relationships between fungal community composition in decomposing leaf litter and home-field advantage effects
DOI:10.1111/fec.v33.8 URL [本文引用: 1]
Environmental factors and traits that drive plant litter decomposition do not determine home-field advantage effects
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Tree species identity alters forest litter decomposition through long-term plant and soil interactions in Patagonia, Argentina
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Fungal community on decomposing leaf litter undergoes rapid successional changes
DOI:10.1038/ismej.2012.116 [本文引用: 2]
Litter chemistry changes more rapidly when decomposed at home but converges during decomposition-transformation
DOI:10.1016/j.soilbio.2012.09.027 URL [本文引用: 1]
Home-field advantage of litter decomposition and nitrogen release in forest ecosystems
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Ecological linkages between aboveground and belowground biota
DOI:10.1126/science.1094875
PMID:15192218
[本文引用: 1]
All terrestrial ecosystems consist of aboveground and belowground components that interact to influence community- and ecosystem-level processes and properties. Here we show how these components are closely interlinked at the community level, reinforced by a greater degree of specificity between plants and soil organisms than has been previously supposed. As such, aboveground and belowground communities can be powerful mutual drivers, with both positive and negative feedbacks. A combined aboveground-belowground approach to community and ecosystem ecology is enhancing our understanding of the regulation and functional significance of biodiversity and of the environmental impacts of human-induced global change phenomena.
Do foliar endophytes matter in litter decomposition?
DOI:10.3390/microorganisms8030446 [本文引用: 1]
Home-field effects of leaf litter decomposition of dominate sand-fixing shrubs in the Horqin Sandy Land
DOI:10.7522/j.issn.1000-694X.2018.00085
[本文引用: 3]
Under the influence of climate change and human activities, sandy grassland in Horqin gradually degenerates into shrubs. In this study, we collected leaf litters of two dominant sand-fixing shrubs,<i>Caraganamicrophylla</i> and <i>Artemisia halodendron</i>,to carry out the interactive transplantation experiments by placing single litter and mixed litter to "home" or "away" environment.wedescirbed CO<sub>2</sub> release and dry matter loss and also compared the difference between measured and predicted CO<sub>2</sub> emissions from mixed litter. We studied the causes of home-field effect and its driving mechanism. Hence, it can provide a theoretical basis for incorporating the home field effect into the litter decomposition model. The results showed that compared with the high-quality leaf litters of <i>C.microphylla</i>, the decomposition of <i>A.halodendron</i> litter had a stronger home field effect. Also, the special effects of soil microbes rather than the transport or storage behavior of soil animals lead to thehome field effect of leaf litter decomposition. In addition, the home field effect of mixed litter is closely related to the similarity of quality of long-term input litter in the decomposing habitat. The greater the quality similarity, the stronger of the home field effect. This study provides a theoretical basis for incorporating the main field effect into the litter decomposition model and improving the simulation accuracy.
科尔沁沙地优势固沙灌木叶片凋落物分解的主场效应
DOI:10.7522/j.issn.1000-694X.2018.00085
[本文引用: 3]
在气候变化和人类活动的影响下,科尔沁沙质草地中灌木植物种增加,导致沙质草地逐渐向灌木地转变。选取该地区优势固沙灌木差不嘎蒿(Artemisia halodendron)和小叶锦鸡儿叶(Caragana microphylla)凋落物及其混合凋落物开展交互移置培养试验,分析了培养过程中CO<sub>2</sub>释放和干物质损失量以及混合凋落物CO<sub>2</sub>释放量实测值与预测值的差异,辨析主场效应产生的原因及其驱动机制,以期为将主场效应纳入到凋落物分解模型提供理论基础。结果表明:与高质量的小叶锦鸡儿叶凋落物相比,质量较低的差不嘎蒿叶凋落物分解具有更强的主场效应;其次,引起叶凋落物分解的主场效应归因于土壤微生物的特化作用,而不是土壤动物的搬运或贮藏行为。此外,混合凋落物主场效应与其分解生境中长期输入的凋落物的质量相似性紧密相关,质量相似性越大,主场效应越强,这也是本研究中混合凋落物分解在差不嘎蒿灌丛土壤下具有较强主场效应的原因。
Stronger effects of litter origin on the processing of conifer than broadleaf leaves: a test of home-field advantage of stream litter breakdown
DOI:10.1111/fwb.13367
[本文引用: 1]
Ecological shifts that enhance the efficiency of resource acquisition by consumers can affect the fate of resource subsidies in recipient ecosystems. To date, findings have been mixed about whether plant litter breakdown by stream decomposers is faster in locations where the litter originates from (i.e. home region) compared with other locations (away region). This phenomenon, known as home-field advantage (HFA), may be influenced by litter quality and particularly decomposer groups (shredders versus microbes), rather than being an inherent consequence of the breakdown of locally native litter. We studied the effects of HFA on litter breakdown and litter-associated communities in streams within mid- to late-successional forests. Two pairs of riparian broadleaf and one pair of conifer litter species with contrasting chemical quality were reciprocally incubated across two temperate regions (similar to 3,000 km apart). Species in each pair are congeneric and allopatric. The HFA was assessed using within-pair litter comparisons in home and away regions. Coarse- and fine-mesh litterbags were used to separate the contributions to litter breakdown by microbial decomposition and shredder feeding. Conifer litter species used in this study were more recalcitrant than broadleaf litter species. Consequently, there was a significant positive HFA for the microbial decomposition of conifer litter. In contrast, there was no HFA for either microbial or shredder-mediated breakdown of broadleaf litter, which might have been overridden by catchment-scale differences in biophysical variables affecting breakdown. The effects of HFA on shredder colonisation and feeding on broadleaf litter were more variable among streams. Numerically abundant shredder taxa tended to have higher rates of colonisation and consumption on higher-quality litter than on low-quality litter, irrespective of litter origin. Our results suggest that prior (evolutionary and/or contemporary) litter exposure could strongly influence the litter-processing capacity of microbial decomposers, but not shredders, at the community level. In particular, microbial decomposers specialised in degrading conifer litter probably had lower resource-use plasticity than those processing broadleaf litter. Inter-stream differences in riparian vegetation and habitat conditions could influence the HFA through altering the litter exposure history of decomposer communities. Our findings highlight the importance of prior litter-microbe interactions in determining the rates of microbial decomposition of native and exotic litter species.
Nitrogen addition enhances home-field advantage during litter decomposition in subtropical forest plantations
DOI:10.1016/j.soilbio.2015.07.026 URL [本文引用: 1]
Litter decomposition in fenced and grazed grasslands: a test of the home-field advantage hypothesis
DOI:10.1016/j.geoderma.2019.07.034 [本文引用: 1]
Home-field advantage of litter decomposition and its soil biological driving mechanism: a review
DOI:10.5846/stxb URL [本文引用: 4]
凋落物分解主场效应及其土壤生物驱动
Rates of litter decomposition in terrestrial ecosystems: global patterns and controlling factors
DOI:10.1093/jpe/rtn002 URL [本文引用: 1]
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