基于植物功能性状的土壤侵蚀退化修复框架

第22卷　第2期2024年4月
中国水土保持科学
Science of Soil and Water Conservation
Vol.22　No.2Apr.2024
DOI :10.16843/j.sswc2023041
基于植物功能性状的土壤侵蚀退化修复框架
郝好鑫1,吕凤玲2,闫峰陵1,郭忠录3†
(1.长江水资源保护科学研究所,430051,武汉;2.武汉市农业技术推广中心,430014,武汉;
3.华中农业大学水土保持研究中心,430070,武汉)
摘要:当前,我国水土保持林草措施正在积极向提升生态功能为目标转变㊂为实现这一目标,管理者和工程师需要明确的生态学理论和方法作为指导㊂通过综述植物功能性状对侵蚀胁迫的响应和对土壤功能的影响机制,验证基于性状响应-影响的生态修复框架在土壤侵蚀治理中的2个关键理论:1)植物性状的变异体现植物对侵蚀退化环境的适应性;2)植物性状显著影响土壤生态过程㊂进而提出基于植物性状响应 影响的侵蚀退化修复框架㊂该框架将植物性状对环境变化响应纳入性状对生态系统影响的分析,根据恢复的水土保持服务目标,选择具有影响上述功能的特定植物性状,选择不受侵蚀(或物种间相互作用)胁迫或具有抗逆性的性状进行植物配置㊂该框架能扩宽当前土壤侵蚀植被治理的研究思路和实践领域,对提升我国的水土保持林草措施具有重要参考意义㊂关键词:侵蚀退化;植被恢复;植物功能性状;土壤保持;生态修复;性状响应 影响框架中图分类号:S157.1
文献标志码:A
文章编号:2096⁃2673(2024)02⁃0159⁃08
引用格式:郝好鑫,吕凤玲,闫峰陵,等.基于植物功能性状的土壤侵蚀退化修复框架[J].中国水土保持科学,2024,22(2):159-166.HAO Haoxin,LÜFengling,YAN Fengling,et al.A plant functional trait⁃based framework for ecological restoration of eroded land[J].Science of Soil and Water Conservation,2024,22(2):159-166.
收稿日期:20230223　修回日期:20230518
项目名称:国家自然科学基金 红壤丘陵区生物结皮驱动下土壤特性动态变化及侵蚀响应”(42077066);国家重大水利工程
建设基金(三峡后续工作)(12620202700221J001);云南省滇中引水工程环境保护科研项目 云南省滇中引水工程隧洞施工地下水位对植被结构功能影响及生态修复优化研究”(DZYS -ZH-HJBH-SJ -002)
第一作者简介:郝好鑫(1992 ),男,博士,高级工程师㊂主要研究方向:水土保持与生态恢复㊂E⁃mail:haohaoxin1992@ †通信作者简介:郭忠录(1980 ),男,博士,教授㊂主要研究方向:水土保持与生态恢复㊂E⁃mail:zlguohzau@
A plant functional trait⁃based framework for ecological restoration
of eroded land
HAO Haoxin 1,LÜFengling 2,YAN Fengling 1,GUO Zhonglu 3
(1.Changjiang Water Resources Protection Institute,430051,Wuhan,China;2.Wuhan Agricultural Technology Extension Centre,430014,Wuhan,China;
3.Research Center of Water and Soil Conservation,Huazhong Agricultural University,430070,Wuhan,China)
Abstract :[Background ]Vegetation restoration is the main eco⁃engineering measure for controlling soil erosion and restoring degraded soils.Recently,scientists have proposed that biological measures for soil and water conservation in China should be oriented to enhance ecological function and should combine the specific ecological problems and social economic factors to set the ecosystem service as the major goal of the ecological restoration.To achieve those goals,managers and engineers require clear framework and
tools.[Methods ]Here,we introduced a trait⁃based approach as a powerful tool to understand the mechanism of soil erosion controlled by vegetation restoration.A literature review was conducted to investigate how plant respond to erosion stresses and how plant influence soil function on plant trait level.
中国水土保持科学2024年[Results]Based on the literature review,two key hypothesizes for applying the trait⁃based response⁃and⁃effect framework to control soil erosion were verified.1)The intraspecific and interspecific variations in plant functional traits were significant heterogeneity.However,those variations were not entirely random, but rather influenced by environmental gradients.2)Plant traits significantly affect soil ecological processes.With this context,we proposed a trait⁃based response⁃and⁃effect framework for ecological restoration on eroded land.That is,plant traits should be selected according to target restored ecosystem service(e.g.,soil conservation,hydrological regulation and carbon storage).Firstly,the environmental stress factors of soil erosion and species interactions of the restored site should be clearly identified. Secondly,selected traits should not only affect the target ecosystem functions,but also not influenced by the environmental stress caused by erosional degradation(or inter⁃species interaction)or have stress tolerance.Furthermore,we summarized the erosion stress as drought,nutrient limitation,physical disturbance.[Conclusions]We believe that this framework could extend the current research ideas and practices of soil and water conservation vegetation measures,and have important practical implications for improving biological measures of soil and water conservation.
Keywords:erosional degradation;vegetation restoration;plant functional trait;soil conservation; ecological restoration;trait⁃based response⁃and⁃effect framework
土壤是人类赖以生存的基本自然资源,为人类可持续发展提供众多关键生态服务[1]㊂在气候变化和人类活动双重压力下,全球面临着广泛的土壤侵蚀加剧和功能退化问题[2]㊂植被恢复被认为是控制土壤侵蚀和修复土壤功能的重要措施[34]㊂然而,土壤侵蚀能显著胁迫植物的定植㊁存活㊁生长和演替,制约植被治理侵蚀和修复生态的效率[56]㊂近年来,有学者提出我国水土保持林草措施应以提升生态功能为导向,治理手段应从综合治理转向生态调控,并结合具体的生态问题和社会经济因素理论和方法作为指导[78]㊂为实现上述目标,管理者和工程师需要明确的生态学理论和方法作为指导[9]㊂
植物功能性状反映植物对环境变化的响应,同时强烈影响生态系统过程[1011]㊂当前基于植物性状响应-影响的框架(a trait⁃based response⁃and⁃effects framework,TRE)是连接环境变化和指导生态修复的前沿工具及热点研究方向[1214]㊂该框架由Suding等[12]在2008年提出,特点在于将性状对环境变化响应纳入性状对生态系统影响的分析,对于退化环境的修复具有强大的指导作用㊂在此基础上,Zirbel等[15]总结出基于TRE的生态修复框架(图1),该框架中植被恢复水平和被修复样地立地条件共同组成环境条件,环境条件直接及通过植物性状间接影响生态系统功能㊂对于用于生态修复的植物性状而言,影响特订生态功能的性状(影响性状)需不受环境条件的制约,即该性状必须能在当前环境条件下正常存活㊁生长和发挥其功能[15]㊂该框架诞生以来,已迅速运用在矿山生态修复㊁退化土壤修复和农业生态系统服务管理等方面[1618]㊂对于土壤侵蚀治理而言,侵蚀过程中径流等侵蚀营力干扰和侵蚀退化环境共同影响植物根系㊁叶片和冠层等性状;同时,这些性状又通过直接和间接作用影响侵蚀营力㊁土壤可蚀性和土壤有机质等性质,进而影响水土保持等众多土壤功能[19]㊂基于此,笔者通过文献综述方法,阐述植物功能性状对侵蚀胁迫的响应和对土壤功能的影响机制,提出基于植物性状以土壤保持为目标的生态修复框架,以期为提升我国水土保持林草措施提供参考㊂
1　基于性状响应—影响的生态修复框架1.1　植物功能性状对侵蚀胁迫响应
植物生长㊁发育和演替不同程度地受到侵蚀过程及其所塑造的土壤㊁地貌和景观等环境因素制约[2021]㊂土壤侵蚀对植被的胁迫可总结为原位和异位影响(图2):1)原位影响主要包含径流等侵蚀营力和泥沙搬运对种子和幼苗的物理扰动,及侵蚀引起的土地退化导致立地条件恶化和生境破碎;2)异位影响包含泥沙沉积对种子㊁幼苗和表层肥沃土壤的埋藏作用[2]㊂针对黄土丘陵区自然植被更新的限制因素,我国学者深入研究植被恢复对土壤侵蚀环境的响应机理,揭示径流等侵蚀扰动对种子和幼苗的影响[19,22]㊂从植物功能性状视角来看,性状对环境变化的响应被认为是植物对环境胁迫适应的
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　第2期
郝好鑫等:
基于植物功能性状的土壤侵蚀退化修复框架　a:性状对环境条件的响应过程㊂b:响应性状与影响性状之间
可能的关系㊂c:影响性状对生态系统功能产生影响的过程㊂d:环境条件对生态系统功能的独立影响,不受功能性状调节㊂a:The processes by which environmental conditions determine response traits.b:The possible relationship between response and effect traits.c:The process by which effect traits produce ecosystem traits.d:The independent effect of environmental conditions on ecosystem functions,
not mediated by functional traits
图1　基于性状响应 影响的生态修复框架[12]
Fig.1　A revised trait⁃based response⁃and⁃effects framework
for ecological restoration [12]
结果,体现环境对植物的可塑性[23]㊂已有研究表明,土壤和人为扰动主导小尺度上性状变异的大小 AWC:Available water content.SOM:Soil organic matter.
图2　侵蚀对植被的原位影响和异位影响
Fig.2　On⁃site and off⁃site effects of soil erosion on vegetation
和方向,例如具有养分吸收功能的细根性状随土壤养分含量降低而增加,而植被地上部分㊁枯落物和粗根则在资源丰富的环境中具有较高产量[2324];Hao 等[6]对丹江口库区不同侵蚀退化程度下对草地㊁灌木地和林地的研究发现,植物偏向采取高效投入 快速收益的光合产物分配模式以适应侵蚀退化土壤
的贫瘠胁迫,具有养分吸收功能的细根总体上随侵蚀退化程度加剧而增加,而粗根则相反㊂综上所述,植物性状在不同侵蚀环境下具有变异性,但变异不是随机的而是受环境梯度影响,体现植物对侵蚀退化环境的适应性㊂
1.2　植物功能性状对土壤功能影响
植物群落能塑造和影响生态系统功能,近些年有研究指出植物功能性状而非物种决定生态系统的
功能和过程,因而应从性状的视角理解植被对生态系统的影响[14,25]㊂已有研究表明,诸如物质生产㊁水分储存和过滤㊁养分存储和循环㊁栖息地维护和物理稳定等众多土壤功能可以通过与植物性状相关联
的一系列土壤生物化学或物理结构性质来表达,而这些土壤性质往往被少数简单易测的植物性状控制[23,26]㊂例如,Celentano 等[27]在巴西亚马逊退化森林生态系统中研究发现,树木高度和断面面积及草本生物量3个性状的退化可解释土壤孔隙㊁含水量㊁有机碳和养分等的变化,进而导致水土保持和生命支持等生态系统服务功能的退化;Lohbeck 等[28]
通过连接植被覆盖度和地上生物量等简单性状与土壤有机碳及侵蚀的关系,指导东非退化农业土壤的修复;Gould 等[29]在德国Jena 生物多样性长期观测实验基地对草地群落的研究发现,根长㊁根重㊁根径和比根长等根系形态性状解释植物多样性影响土壤团聚体稳定性㊁入渗和抗剪等一系列与土壤功能相关土壤性质的变化㊂
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中国水土保持科学2024年
1.3　基于性状响应—影响的生态修复框架
基于性状响应 影响的框架认为生态系统直接受生物和非生物环境因素控制,植被和环境及其反馈过程能共同塑造新的生态系统[11]㊂基于此,Zirbel 等[15]总结出基于性状响应 影响的生态修复框架(图1),该框架中植被恢复的管理水平和被修复的立地条件共同组成环境条件,环境条件直接或通过植物性状间接影响生态系统功能㊂对于筛选用于生态修复的植物性状而言,
具有特定生态功能影响的性状SOM:Soil organic matter.AS:Aggregate stability.
图3　基于植物性状响应 影响的土壤侵蚀退化修复框架
Fig.3　A trait⁃based response⁃and⁃effect framework for ecological restoration of eroded land
(影响性状)不受环境条件的制约,即响应性状和影响
性状发生重叠时,响应性状必须能在当前环境条件下正常存活㊁生长和发挥其功能㊂例如Navarro⁃Cano 等[17]在为修复尾矿筛选合适保育植物中发现,先锋物种虽然能在贫瘠土壤上迅速生长,但其不能促进其他植物生长和改善土壤条件,因而强调保育植物需同时满足能在极端环境压力下正常生存并且需具备修复土壤功能作用,并以此总结出适合尾矿修复的性状:较大植株形态和较长生命史等对生态系统功能促进作用大的性状,C4植物㊁主根与侧根比及叶片碳氮比低进而能在极端环境下生存性强的性状㊂
2　基于植物性状的土壤侵蚀退化修复框架
笔者阐述植物功能性状对侵蚀退化胁迫的响应,分析植物性状对一系列与土壤功能紧密联系的土壤性质影响,验证基于性状响应 影响的生态修复框架中2个关键理论:1)植物性状存在较大的种内和种间变异,但变异不是随机的而是受环境梯度影响,体现植物对侵蚀退化环境的适应性;2)植物性状显著影响土壤生态过程㊂综上,笔者提出一种以水土保持服务为目标的基于植物性状的生态修复框架(图3),该框架包含修复场地中胁迫因子和群落种间相互作用的识别,性状对上述胁迫因子和相互作用的响应,及性状对侵蚀营力和土壤可蚀性的影响3方面㊂
识别修复场地的侵蚀胁迫因子和群落种间相互作用关系是性状筛选的关键㊂图3中,浅蓝色菱形
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　第2期郝好鑫等:基于植物功能性状的土壤侵蚀退化修复框架
框表示筛选性状的步骤,其中的 /”表示筛选的性状需不受侵蚀环境因子(或植物种间相互作用)的胁迫, +”表示筛选的性状需对侵蚀环境胁迫(或植物种间相互作用)具有抗性, /”和 +”同时被箭头包涵时表示筛选的性状需不受影响( /”)或具有抗性( +”)㊂这一步的目的是保证植物在修复场地中的存活和生长进而保证其能发挥正常功能㊂表
1列举出常见侵蚀胁迫及抗胁迫性状:例如干旱是侵蚀造成常见的胁迫之一,植物抵抗干旱胁迫需要 高幼苗存活率,高水分利用效率,长的生命史和低蒸
腾面积等,相关性状特征可为大型植株,细根发达,
深根系及叶片角质层厚等[3033];人工林或种植园植被类型单一容易引起林下侵蚀,而能在林下正常存
活的植物的叶片需要具有耐阴等性状,并且植物种
间不存在异株克生的化感作用,此外林下植株应尽
量矮小以减少溅蚀发生[3435]㊂
选择对侵蚀营力和土壤可蚀性具有影响的性状进行组合是利用植被实现水土保持服务的关键㊂
表1　抗侵蚀胁迫相关植物性状特征及举例
Tab.1　Characteristics and examples of plant trait related to anti⁃erosion stress
侵蚀胁迫Erosion⁃related stresses
抗胁迫植物特征
Characteristics of stress⁃resistant plant相关性状举例
Examples of related traits参考文献
References
　干旱　Drought 高水分利用效率,长生命史,低蒸腾面积
　High water use efficiency,long life span
and low transpiration area
　大型植株,细根发达,深根系,小的表面积或体积比,叶片角
质层厚,等
　Large plant,high density of fine root,deep root system,less
surface or volume rate,thick leaf cuticle,etc.
[3033]
　林下侵蚀　Understory ero⁃sion 林下植被能与乔木共生
　Understory vegetation coexists with trees
　乔木枯落物易分解,林下植物耐阴,植株矮小,等
　Easily decompose of tree litter,shade⁃tolerant understory vege⁃
tation,dwarf plant,etc.
[3435]
　土壤贫瘠化　Soil nutrient de⁃pletion 高养分利用效率,权衡用于养分吸收和
生命活动的物质和能量分配
　High nutrient use efficiency,balance the
distribution of matter and energy for nutrient
absorption and life activities
　细根发达,比根长较高,器官氮含量较高㊁干物质低,具有固
氮能力,等
　High density of fine root,high specific root length,high nitro⁃
gen content with low dry mater in plant organ,fix nitrogen capaci⁃
ty,etc.
[24,3640]
　土壤密度增加　Increase soil bulk density 高幼苗存活率,扎根能力强
　High seedling colonization rate and strong
rooting ability
　高粗根比例,低根系木质部导管比例,营养繁殖,等
　High proportion of coarse roots,low proportion of root xylem
vessels,vegetative propagation,etc.
[41]
　砂粒含量增加　Increase the proportion of sand⁃y particle 高水分和养分利用效率
　High water use efficiency
　根长密度大,等
　High root length density,etc.[36]
　土壤埋藏　Burial of topsoil 种子萌发力强
　Strong seed germination capacity
　高芽位,地上芽,地面芽,种子萌发周期短,营养繁殖,等
　High aboveground buds,short germination period,vegetative
propagation,etc.
[21,42]
　径流扰动　Runoff disturb⁃ance 种子不易流失,根系拔出阻力大,粗壮或
柔性茎秆
　Seeds are not easy to lose,high uprooting
resistance,and strong or flexible stem.
　种子质量大㊁狭长形态㊁具有附属物㊁分泌粘液,植冠种子
库,深根系,侧根发达,柔性茎秆,等
　Seed with great mass,extreme elongated shape,appendages
and mucilage segregation,canopy seed bank,deep root system,
well⁃developed of lateral root,strong or flexible stem,etc.
[4244]
　细沟侵蚀　Rill erosion 种子不易流失
　Seeds are not easy to lose
　种子质量大㊁狭长形态㊁具有附属物㊁分泌粘液,等
　Seed with great mass,extreme elongated shape,appendages
and mucilage segregation,etc.
[4344]
　陡坡和沟蚀　Steep slope& gully erosion 高种子萌发率,高幼苗存活率,抗物理干
扰能力强
　High seed germination rate,high seedling
colonization rate and strong ability to resist
physical disturbance
　高地上芽位,种子萌发周期短,营养繁殖,大型植株,深根
系,等
　High aboveground buds,short germination period,vegetative
propagation,large plant,deep root system,etc.
[2,45]
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图3中, +”表示选择能增加土壤入渗㊁地表覆盖㊁土壤有机质㊁团聚体稳定性和黏聚力等控制侵蚀相关的植物性状㊂这一步目的是组合具有水土保持相关功能(如土壤保持㊁水文调节和碳存储等)性状以实现最终的目标服务㊂例如Hao等连接植物功能性状与土壤侵蚀的系列研究[6,3638]中发现高植物覆盖度㊁高枯落量㊁高细根长密度和高根际微生物等均显著促进土壤有机质㊁团聚体稳定性和降低土壤可蚀性,可以选择组合具有上述性状的植物构建群落从而实现土壤保持的生态修复目标㊂
3　讨论
笔者基于功能生态学㊁植物生态学和土壤侵蚀相关学科前沿进展,借鉴恢复生态学领域实践经验,提出基于性状响应 影响的土壤侵蚀退化修复框架㊂然而,实际操作中筛选不受侵蚀环境(或植物种间相互作用)胁迫或对侵蚀环境胁迫(或植物种间相互作用)具有抗性,同时又能影响土壤保持功能的性状具有一定挑战㊂已有的植被恢复策略选择往往依据专家经验,而本研究提出的生态修复框架不仅对专家友好,更衔接了众多生态学原理的应用㊂值得期待的是,随着功能生态学的迅速发展,已有大量研究报道植物的形态㊁生理㊁生化和解剖结构等性状特征及其对生物和非生物环境变化的响应,并且国际上已建立开放的性状数据库帮助研究者查询上述信息(TRY plant trait database,http:∥www.try⁃
)[46]㊂此外,Perez等[47]建立了用于控制土壤侵蚀和浅层滑坡的植物物种开放数据库(stability database,http:∥publish.plantnet⁃/project/ stability_en),这为研究者和工程师选择目标植物性状提供了更广泛和灵活的物种来源㊂但是,植物对环境的响应和植物对生态过程的影响均是极其复杂的过程,目前相关的认识仍较为有限㊂此外,一些研究还发现部分植物性状和生态理论预测的生境存在广泛的不匹配现象[48]㊂这说明无论采用专家经验或现有生态理论分析的方法来筛选性状都可能存在考虑不周的情况㊂因此,在开展大规模植被恢复前,有必要在修复场地内布设小型试验验证所选植物群落的存活能力㊂
4　结论
1)土壤侵蚀对植被的胁迫可总结为原位和异位影响,原位影响主要包含径流等侵蚀营力和泥沙搬运对种子和幼苗的物理扰动,及侵蚀引起的土地退化导致立地条件恶化和生境破碎;异位影响包含泥沙沉积对种子和幼苗的埋藏作用,及泥沙沉积对表层肥沃土壤的埋藏㊂
2)通过综述植物功能性状对侵蚀胁迫的响应和对土壤功能的影响机制,验证基于性状响应 影响的生态修复框架在土壤侵蚀治理中应用的2个关键理论:植物性状存在较大的种内和种间变异,体现植物对侵蚀退化环境的适应性;植物性状显著影响土壤生态过程㊂
3)提出基于植物性状响应 影响的土壤侵蚀退化修复框架㊂该框架特点在于将植物性状对环境变化响应纳入性状对生态系统影响的分析,即根据要恢复的水土保持服务目标(如土壤保持㊁水文调节和碳存储等)选择具有影响上述功能的特定植物性状,并明确修复场地的土壤侵蚀退化胁迫因子和物种间相互作用,选择不受侵蚀(或物种间相互作用)胁迫或具有抗逆性的性状进行植物配置㊂
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