草原生态系统土壤多功能性及影响机制研究进展

doi:10.7522/j.issn.1000-694X.2023.00049

摘要/Abstract

摘要：

草原是陆地生态系统的关键组成部分，为维持区域生态安全发挥着极为重要的作用。土壤多功能性（soil multifunctionality，SMF）是衡量土壤系统提供多种功能的综合指标，对评估草原生态系统功能和区域可持续发展具有重要指导意义。然而，目前SMF研究仍处于初步阶段，研究对象多为农田生态系统，对于草原等自然生态系统的研究较少。此外，对草原生态系统SMF与植物和土壤微生物的关系以及全球变化对草原生态系统SMF影响机制的认识十分有限。因此，本文从SMF的概念和内涵、量化指标及方法、植物和土壤微生物与SMF的关系、全球变化对SMF的影响以及SMF的空间变异性5个方面论述SMF的研究进展，梳理了草原生态系统SMF的变化特征及影响机制。根据当前研究中存在的问题提出了今后研究应重点关注的内容：（1）统一规整SMF量化体系；（2）SMF在不同空间尺度上的转换机制；（3）不同类型草原生态系统SMF驱动机制；（4）土地利用变化、降水增加、气候变暖和氮沉降等全球变化交互作用对SMF的影响；（5）地上植物功能与SMF的联系和互馈机制；（6）根系功能性状对SMF的影响和植物在不同组织水平对SMF的贡献；（7）微生物稀有种群以及植物-土壤动物-土壤微生物多营养级联动对SMF的影响。

关键词: 草原生态系统, 全球变化, 土壤多功能性, 土壤质量

Abstract:

Grassland is a crucial part of terrestrial ecosystem and plays an important role in maintaining regional ecological security. Soil multifunctionality （SMF） is a comprehensive index to measure the provided soil multiple functions， which has imperative significance for assessing grassland ecosystem function and regional sustainable development. However， the current SMF studies are still in the initial stage， and the objects in studies are mostly farmland ecosystems， while there are relatively few studies on grassland and other natural ecosystems. In addition， there is few ecolgoists to elucidate the relationship among SMF of grassland ecosystems， plant and soil microorganisms， as well as the effect of global change on SMF in grassland ecosystems. Therefore， this paper discusses the study progress of SMF from five aspects： the concept of SMF， quantitative indicators and methods， the relationship among plants and soil microorganisms and SMF， the impact of global changes on SMF and the spatial variability of SMF， and reviews the change and mechanisms of SMF in grassland ecosystems. Based on the existing problems in the existing studies， the following topics should be focused on：（1） Unified SMF quantitative methods；（2） The transformation mechanism of SMF at different spatial scales；（3） SMF driving mechanism of different grassland ecosystems；（4） Impacts of global changes such as precipitation increase， climate warming and nitrogen deposition， and land use change on SMF；（5） The relationships and interactions between above-ground ecological functions and SMF；（6） Effects of root functional traits on SMF and contributions of plants to SMF at different scale levels；（7） The contribution of rare species of microorganisms and plant-soil animal-soil microbial multitrophic linkage on SMF.

Key words: grassland ecosystem, global change, soil multifunctionality, soil quality

中图分类号:

S154.1

宋兆斌, 岳平, 李香云, 胡亚, 乔静娟, 左小安. 草原生态系统土壤多功能性及影响机制研究进展[J]. 中国沙漠, 2023, 43(6): 151-165.

Zhaobin Song, Ping Yue, Xiangyun Li, Ya Hu, Jingjuan Qiao, Xiaoan Zuo. Variation and influencing mechanism of soil multifunctionality in grassland ecosystem[J]. Journal of Desert Research, 2023, 43(6): 151-165.

图/表 5

参考文献 131

1	Wall D H， Nielsen U N， Six J.Soil biodiversity and human health［J］.Nature，2015，528（7580）：69-76.
2	Karlen D L， Ditzler C A， Andrews S S.Soil quality：why and how？［J］.Geoderma，2003，114：145-156.
3	Brussaard L.Biodiversity and ecosystem functioning in soil［J］.Ambio，1997，26（8）：563-570.
4	Bennett J A， Klironomos J.Mechanisms of plant-soil feedback：interactions among biotic and abiotic drivers［J］.New Phytologist，2019，222（1）：91-96.
5	Xi N， Adler P B， Chen D，et al.Relationships between plant-soil feedbacks and functional traits［J］.Journal of Ecology，2021，109（9）：3411-3423.
6	Bouma J， Mc Bratney A.Framing soils as an actor when dealing with wicked environmental problems［J］.Geoderma，2013，200：130-139.
7	Hartemink A E， Mc Bratney A.A soil science renaissance［J］.Geoderma，2008，148：123-129.
8	Creamer R E， Barel J M， Bongiorno G，et al.The life of soils：Integrating the who and how of multifunctionality［J］.Soil Biology and Biochemistry，2022，166：108561.
9	Manning P， van der Plas F， Jonathan S，et al.Redefining ecosystem multifunctionality［J］.Nature Ecology & Evolution，2018，2（3）：427-436.
10	Lefcheck J S， Byrnes J E K， Isbell F，et al.Biodiversity enhances ecosystem multifunctionality across trophic levels and habitats［J］.Nature Communications，2015，6（1）：1-7.
11	Zhang J Z， Li T T， Jia J Y，et al.Bacterial taxa and fungal diversity are the key factors determining soil multifunctionality in different cropping systems［J］.Land Degradation & Development，2021，32：5012-5022.
12	Sanderson M A， Skinner R H， Barker D J，et al.Plant species diversity and management of temperate forage and grazing land ecosystems［J］.Crop Science，2004，44（4）：1132-1144.
13	Hector A， Bagchi R.Biodiversity and ecosystem multifunctionality ［J］.Nature，2007，448（7150）：188-190.
14	Haygarth P M， Ritz K.The future of soils and land use in the UK：soil systems for the provision of land-based ecosystem services［J］.Land Use Policy，2009，26：187-197.
15	Schulte R P O， Creamer R E， Donnellan T，et al.Functional land management：a framework for managing soil-based ecosystem services for the sustainable intensification of agriculture［J］.Environmental Science & Policy，2014，38：45-58.
16	Hu W， Ran J， Dong L，et al.Aridity-driven shift in biodiversity-soil multifunctionality relationships［J］.Nature Communications，2021，12（1）：1-15.
17	Bai Y， Cotrufo M F.Grassland soil carbon sequestration：current understanding，challenges，and solutions［J］.Science，2022，377（6606）：603-608.
18	Liu Y R， Delgado-Baquerizo M， Trivedi P，et al.Identity of biocrust species and microbial communities drive the response of soil multifunctionality to simulated global change［J］.Soil Biology and Biochemistry，2017，107：208-217.
19	Fry E L， Savage J， Hall A L，et al.Soil multifunctionality and drought resistance are determined by plant structural traits in restoring grassland［J］.Ecology，2018，99（10）：2260-2271.
20	Valencia E， Gross N， Quero J L，et al.Cascading effects from plants to soil microorganisms explain how plant species richness and simulated climate change affect soil multifunctionality［J］.Global Change Biology，2018，24（12）：5642-5654.
21	Guo Y， Xu T， Cheng J，et al.Above-and belowground biodiversity drives soil multifunctionality along a long-term grassland restoration chronosequence［J］.Science of The Total Environment，2021，772：145010.
22	Mausel P W.Soil quality in Illinois：an example of a soils geography resource analysis［J］.The Professional Geographer，1971，23（2）：127-136.
23	Doran J W， Parkin T B.Defining and assessing soil quality［J］.Environmental Science，1994，35：1-21.
24	Andrews S S， Karlen D L， Cambardella C A.The soil management assessment framework［J］.Soil Science Society of America Journal，2004，68（6）：1945-1962.
25	Bünemann E K， Bongiorno G， Bai Z，et al.Soil quality：a critical review［J］.Soil Biology and Biochemistry，2018，120：105-125.
26	李奕赞，张江周，贾吉玉，等.农田土壤生态系统多功能性研究进展［J］.土壤学报，59（5）：1187-1195.
27	Wen Z， Zheng H， Zhao H，et al.Land-use intensity indirectly affects soil multifunctionality via a cascade effect of plant diversity on soil bacterial diversity［J］.Global Ecology and Conservation，2020，23：e01061.
28	Wang H， Bu L， Tian J，et al.Particular microbial clades rather than total microbial diversity best predict the vertical profile variation in soil multifunctionality in desert ecosystems［J］.Land Degradation & Development，2021，32（6）：2157-2168.
29	Su Y G， Liu J， Zhang Y M，et al.More drought leads to a greater significance of biocrusts to soil multifunctionality［J］.Functional Ecology，2021，35：989-1000.
30	Dooley A， Isbell F， Kirwan L，et al.Testing the effects of diversity on ecosystem multi-functionality using a multivariate model［J］.Ecology Letters，2015，18：1242-1251.
31	Maestre F T， Delgado-Baquerizo M， Jeffries T C，et al.Increasing aridity reduces soil microbial diversity and abundance in global drylands［J］.Proceedings of the National Academy of Sciences，2015，112（51）：15684-15689.
32	Byrnes J E K， Gamfeldt L， Isbell F，et al.Investigating the relationship between biodiversity and ecosystem multifunctionality：challenges and solutions［J］.Methods in Ecology and Evolution，2014，5（2）：111-124.
33	徐炜，井新，马志远，等.生态系统多功能性的测度方法［J］.生物多样性，2016，24（1）：72-84.
34	Li J W， Liu Y L， Hai X Y，et al.Dynamics of soil microbial C∶N∶P stoichiometry and its driving mechanisms following natural vegetation restoration after farmland abandonment［J］.Science of Total Environment，2019，693：133613.
35	Maestre F T， Quero J L， Gotelli N J，et al.Plant species richness and ecosystem multifunctionality in global drylands［J］.Science，2012，335：214-218.
36	Delgado-Baquerizo M， Maestre F T， Reich P B，et al.Microbial diversity drives multifunctionality in terrestrial ecosystems［J］.Nature Communications，2016，7（1）：1-8.
37	Jing X， Sanders N J， Shi Y，et al.The links between ecosystem multifunctionality and above-and belowground biodiversity are mediated by climate［J］.Nature Communications，2015，6（Sep）.
38	Orwin K H， Buckland S M， Johnson D，et al.Linkages of plant traits to soil properties and the functioning of temperate grassland［J］.Journal of Ecology，2010，98（5）：1074-1083.
39	Dijkstra F A， Carrillo Y， Pendall E，et al.Rhizosphere priming：a nutrient perspective［J］.Frontiers in Microbiology，2013，4：216.
40	Baets S D， Poesen J， Knapen A，et al.Impact of root architecture on the erosion‐reducing potential of roots during concentrated flow［J］.Earth Surface Processes and Landforms，2007，32（9）：1323-1345.
41	Stokes A， Atger C， Bengough A G，et al.Desirable plant root traits for protecting natural and engineered slopes against landslides［J］.Plant and Soil，2009，324（1）：1-30.
42	Kuzyakov Y.Priming effects：interactions between living and dead organic matter［J］.Soil Biology and Biochemistry，2010，42（9）：1363-1371.
43	Graf F， Frei M.Soil aggregate stability related to soil density，root length，and mycorrhiza using site-specific Alnus incana and Melanogaster variegatus s.l.［J］.Ecological Engineering，2013，57：314-323.
44	Eisenhauer N， Lanoue A， Strecker T，et al.Root biomass and exudates link plant diversity with soil bacterial and fungal biomass［J］.Scientific Reports，2017，7：1-8.
45	Yan Y， Zhang Q， Buyantuev A，et al.Plant functional β diversity is an important mediator of effects of aridity on soil multifunctionality［J］.Science of the Total Environment，2020，726：138529.
46	Hautier Y， Isbell F， Borer E T，et al.Local loss and spatial homogenization of plant diversity reduce ecosystem multifunctionality［J］.Nature Ecology & Evolution，2018，2（1）：50-56.
47	Kardol P， Cregger M A， Campany C E，et al.Soil ecosystem functioning under climate change：plant species and community effects［J］.Ecology，2010，91（3）：767-781.
48	Cardinale B J， Duffy J E， Gonzalez A，et al.Biodiversity loss and its impact on humanity［J］.Nature，2012，486（7401）：59-67.
49	Tilman D， Wedin D， Knops J.Productivity and sustainability influenced by Phylotype diversity within soil fungal functional biodiversity in grassland ecosystems［J］.Nature，1996，379（6567）：718-720.
50	Inkotte J， Bomfim B， da Silva S C，et al.Linking soil biodiversity and ecosystem function in a Neotropical savanna ［J］.Applied Soil Ecology，2022，169：104209.
51	Liu S， García-Palacios P， Tedersoo L，et al.Phylotype diversity within soil fungal functional groups drives ecosystem stability［J］.Nature Ecology & Evolution，2022：1-10.
52	Fanin N， Moorhead D， Bertrand I.Eco-enzymatic stoichiometry and enzymatic vectors reveal differential C，N，P dynamics in decaying litter along a land-use gradient［J］.Biogeochemistry，2016，129（1）：21-36.
53	García‐Palacios P， Maestre F T， Kattge J，et al.Climate and litter quality differently modulate the effects of soil fauna on litter decomposition across biomes［J］.Ecology letters，2013，16（8）：1045-1053.
54	Delgado‐Baquerizo M， Eldridge D J， Ochoa V，et al.Soil microbial communities drive the resistance of ecosystem multifunctionality to global change in drylands across the globe［J］.Ecology Letters，2017，20（10）：1295-1305.
55	Wagg C， Bender S F， Widmer F，et al.Soil biodiversity and soil community composition determine ecosystem multifunctionality［J］.Proceedings of the National Academy of Sciences，2014，111（14）：5266-5270.
56	Falkowski P G， Fenchel T， Delong E F.The microbial engines that drive Earth's biogeochemical cycles［J］.Science，2008，320（5879）：1034-1039.
57	Fierer N.Embracing the unknown：disentangling the complexities of the soil microbiome［J］.Nature Reviews Microbiology，2017，15（10）：579-590.
58	Philippot L， Spor A， Hénault C，et al.Loss in microbial diversity affects nitrogen cycling in soil［J］.The ISME Journal，2013，7（8）：1609-1619.
59	Wang J， Wang X， Liu G，et al.Bacterial richness is negatively related to potential soil multifunctionality in a degraded alpine meadow［J］.Ecological Indicators，2021，121：106996.
60	Loreau M， Hector A.Partitioning selection and complementarity in biodiversity experiments［J］.Nature，2011，412：72-76.
61	Hooper D U， Chapin F S III， Ewel J J，et al.Effects of biodiversity on ecosystem functioning，a consensus of current knowledge［J］.Ecological Monographs，2005，75：3-35.
62	Delgado-Baquerizo M， Trivedi P， Trivedi C，et al.Microbial richness and composition independently drive soil multifunctionality［J］.Functional Ecology，2017，31（12）：2330-2343.
63	Ma L， Zhang C， Xu X，et al.Different facets of bacterial and fungal communities drive soil multifunctionality in grasslands spanning a 3，500 km transect［J］.Functional Ecology，36（12）：3120-3133.
64	Li Z， Liu X， Zhang M，et al.Plant diversity and fungal richness regulate the changes in soil multifunctionality in a semi-arid grassland［J］.Biology，2022，11（6）：870.
65	Clemmensen K E， Bahr A， Ovaskainen O，et al.Roots and associated fungi drive long-term carbon sequestration in boreal forest［J］.Science，2013，339（6127）：1615-1618.
66	Clemmensen K E， Finlay R D， Dahlberg A，et al.Carbon sequestration is related to mycorrhizal fungal community shifts during long-term succession in boreal forests［J］.New Phytologist，2015，205（4）：1525-1536.
67	Li J， Delgado-Baquerizo M， Wang J T，et al.Fungal richness contributes to multifunctionality in boreal forest soil［J］.Soil Biology and Biochemistry，2019，136：107526.
68	Xu H， Yu M， Cheng X.Abundant fungal and rare bacterial taxa jointly reveal soil nutrient cycling and multifunctionality in uneven-aged mixed plantations［J］.Ecological Indicators，2021，129：107932.
69	Mahmoudi N， Caeiro M F， Mahdhi M，et al.Arbuscular mycorrhizal traits are good indicators of soil multifunctionality in drylands［J］.Geoderma，2021，397：115099.
70	Qiao Y， Bai Y， She W，et al.Arbuscular mycorrhizal fungi outcompete fine roots in determining soil multifunctionality and microbial diversity in a desert ecosystem［J］.Applied Soil Ecology，2022，171：104323.
71	Zhang Z， Lu Y， Wei G，et al.Rare species-driven diversity-ecosystem multifunctionality relationships are promoted by stochastic community assembly［J］.mBio，2022：e0044922.
72	Xiong C， He J Z， Singh B K，et al.Rare taxa maintain the stability of crop mycobiomes and ecosystem functions［J］.Environmental Microbiology，2021，23（4）：1907-1924.
73	Elshahed M S， Youssef N H， Spain A M，et al.Novelty and uniqueness patterns of rare members of the soil biosphere［J］.Applied and Environmental Microbiology，2008，74（17）：5422-5428.
74	Wu Y， Chen D， Saleem M，et al.Rare soil microbial taxa regulate the negative effects of land degradation drivers on soil organic matter decomposition［J］.Journal of Applied Ecology，2021，58（8）：1658-1669.
75	Jousset A， Bienhold C， Chatzinotas A，et al.Where less may be more：how the rare biosphere pulls ecosystems strings［J］.The ISME Journal，2017，11（4）：853-862.
76	Van Den Hoogen J， Geisen S， Routh D，et al.Soil nematode abundance and functional group composition at a global scale［J］.Nature，2019，572（7768）：194-198.
77	Yeates G W.Nematodes as soil indicators：functional and biodiversity aspects［J］.Biology and Fertility of Soils，2003，37：199-210.
78	Zhang C， Wang J， Ren Z，et al.Root traits mediate functional guilds of soil nematodes in an ex-arable field［J］.Soil Biology and Biochemistry，2020，151：108038.
79	Gibelin A L， Déqué M.Anthropogenic climate change over the Mediterranean region simulated by a global variable resolution model［J］.Climate Dynamics，2003，20（4）：327-339.
80	Huntington T G.Evidence for intensification of the global water cycle：Review and synthesis［J］.Journal of Hydrology，2006，319（1）：83-95.
81	Meron E， Gilad E， Von Hardenberg J，et al.Vegetation patterns along a rainfall gradient［J］.Chaos，Solitons & Fractals，2004，19（2）：367-376.
82	Durán J， Delgado‐Baquerizo M， Dougill A J，et al.Temperature and aridity regulate spatial variability of soil multifunctionality in drylands across the globe［J］.Ecology，2018，99（5）：1184-1193.
83	Singh A K， Rai A， Banyal R，et al.Plant community regulates soil multifunctionality in a tropical dry forest［J］.Ecological Indicators，2018，95：953-963.
84	De Vries F T， Shade A.Controls on soil microbial community stability under climate change［J］.Frontiers in Microbiology，2013，4：265.
85	Kaisermann A， De Vries F T， Griffiths R I，et al.Legacy effects of drought on plant-soil feedbacks and plant-plant interactions［J］.New Phytologist，2017，215（4）：1413-1424.
86	Rousk J， Smith A R， Jones D L.Investigating the long-term legacy of drought and warming on the soil microbial community across five European shrubland ecosystems［J］.Global Change Biology，2013，19（12）：3872-3884.
87	Dacal M， Garcia-Palacios P， Asensio S，et al.Climate change legacies contrastingly affect the resistance and resilience of soil microbial communities and multifunctionality to extreme drought［J］.Functional Ecology，2022，36：908-920.
88	Steinauer K， Tilman D， Wragg P D，et al.Plant diversity effects on soil microbial functions and enzymes are stronger than warming in a grassland experiment［J］.Ecology，2015，96（1）：99-112.
89	Schrama M， Bardgett R D.Grassland invasibility varies with drought effects on soil functioning［J］.Journal of Ecology，2016，104（5）：1250-1258.
90	Toberman H， Evans C D， Freeman C，et al.Summer drought effects upon soil and litter extracellular phenol oxidase activity and soluble carbon release in an upland Calluna heathland［J］.Soil Biology and Biochemistry，2008，40（6）：1519-1532.
91	Lozano Y M， Aguilar-Trigueros C A， Onandia G，et al.Effects of microplastics and drought on soil ecosystem functions and multifunctionality［J］.Journal of Applied Ecology，2021，58：988-996.
92	Vadez V.Root hydraulics：the forgotten side of roots in drought adaptation［J］.Field Crops Research，2014，165：15-24.
93	Fowler D， Coyle M， Skiba U，et al.The global nitrogen cycle in the twenty-first century［J］.Philosophical Transactions of the Royal Society B：Biological Sciences，2013，368（1621）：20130164.
94	Galloway J N， Townsend A R， Erisman J W，et al.Transformation of the nitrogen cycle：recent trends，questions，and potential solutions［J］.Science，2008，320：889-892.
95	Steffen W， Richardson K， Rockström J，et al.Planetary boundaries：guiding human development on a changing planet［J］.Science，2015，347（6223）：1259855.
96	Elser J J， Bracken M E S， Cleland E E，et al.Global analysis of nitrogen and phosphorus limitation of primary producers in freshwater，marine and terrestrial ecosystems［J］.Ecology Letters，2007，10：1135-1142.
97	Vitousek P M， Aber J D， Howarth R W，et al.Human alteration of the global nitrogen cycle：sources and consequences［J］.Ecological Applications，1997，7：737-750.
98	Saiya-Cork K R， Sinsabaugh R L， Zak D R.The effects of long term nitrogen deposition on extracellular enzyme activity in an Acer saccharum forest soil［J］.Soil Biology and Biochemistry，2002，34：1309-1315.
99	Wei C， Yu Q， Bai E，et al.Nitrogen deposition weakens plant-microbe interactions in grassland ecosystems［J］.Global Change Biology，2013，19（12）：3688-3697.
100	Liu X， Lamb E G， Zhang S.Nitrogen addition impacts on soil microbial stoichiometry are driven by changes in plant resource stoichiometry not by the composition of main microbial groups in an alpine meadow［J］.Biology and Fertility of Soils，2020，56（2）：261-271.
101	Yuan Z， Ali A， Ruiz‐Benito P，et al.Above‐and below‐ground biodiversity jointly regulate temperate forest multifunctionality along a local‐scale environmental gradient［J］.Journal of Ecology，2020，108（5）：2012-2024.
102	Xu Z， Li M H， Zimmermann N E，et al.Plant functional diversity modulates global environmental change effects on grassland productivity［J］.Journal of Ecology，2018，106（5）：1941-1951.
103	Yuan X， Niu D， Gherardi L A，et al.Linkages of stoichiometric imbalances to soil microbial respiration with increasing nitrogen addition：evidence from a long-term grassland experiment［J］.Soil Biology and Biochemistry，2019，138：107580.
104	Liu X， Shi X， Zhang S.Soil abiotic properties and plant functional diversity co-regulate the impacts of nitrogen addition on ecosystem multifunctionality in an alpine meadow［J］.Science of the Total Environment，2021，780：146476.
105	Bell C， Carrillo Y， Boot C M，et al.Rhizosphere stoichiometry：are C∶N∶P ratios of plants，soils，and enzymes conserved at the plant species-level？［J］.New Phytologist，2014，201（2）：505-517.
106	Liu W， Liu L， Yang X，et al.Long-term nitrogen input alters plant and soil bacterial，but not fungal beta diversity in a semiarid grassland［J］.Global Change Biology，2021，27（16）：3939-3950.
107	Bai Y， Wu J， Clark C M，et al.Tradeoffs and thresholds in the effects of nitrogen addition on biodiversity and ecosystem functioning：evidence from Inner Mongolia grasslands［J］.Globe Change Biology，2010，16：358-372.
108	Liu X， Zhang S.Nitrogen addition shapes soil enzyme activity patterns by changing pH rather than the composition of the plant and microbial communities in an alpine meadow soil［J］.Plant and Soil，2019，440：11-24.
109	Cui H Y， Sun W， Delgado-Baquerizo M，et al.Phosphorus addition regulates the responses of soil multifunctionality to nitrogen over-fertilization in a temperate grassland［J］.Plant and Soil，2022，473：73-87.
110	de Gea A B， Hautier Y， Geisen S.Interactive effects of global change drivers as determinants of the link between soil biodiversity and ecosystem functioning［J］.Global Change Biology，2023，29（2）：296-307.
111	Crain C M， Kroeker K， Halpern B S.Interactive and cumulative effects of multiple human stressors in marine systems［J］.Ecology Letters，2008，11（12）：1304-1315.
112	Shovon T A， Rozendaal D M A， Gagnon D，et al.Plant communities on nitrogen-rich soil are less sensitive to soil moisture than plant communities on nitrogen-poor soil［J］.Journal of Ecology，2020，108（1）：133-144.
113	Klaus V H， Kleinebecker T， Busch V，et al.Land use intensity，rather than plant species richness，affects the leaching risk of multiple nutrients from permanent grasslands［J］.Global Change Biology，2018，24（7）：2828-2840.
114	Ren H， Eviner V T， Gui W，et al.Livestock grazing regulates ecosystem multifunctionality in semi arid grassland［J］.Functional Ecology，2018，32（12）：2790-2800.
115	Newbold T， Hudson L N， Contu S，et al.Widespread winners and narrow-ranged losers：land use homogenizes biodiversity in local assemblages worldwide［J］.PLoS Biology，2018，16（12）：e2006841.
116	Ding L， Wang P.Afforestation suppresses soil nitrogen availability and soil multifunctionality on a subtropical grassland［J］.Science of the Total Environment，2021，761：143663.
117	Chen K， Zhou H， Lu B，et al.Single-species artificial grasslands decrease soil multifunctionality in a temperate steppe on the Qinghai-Tibet Plateau［J］.Agronomy，2021，11（11）：2092.
118	Liu Q， Zhang Q， Jarvie S，et al.Ecosystem restoration through aerial seeding：interacting plant-soil microbiome effects on soil multifunctionality［J］.Land Degradation & Development，2021，32（18）：5334-5347.
119	Jackson R B， Caldwell M M.The scale of nutrient eeterogeneity around individual plants and its quantification with geostatistics［J］.Ecology，1993，74：612-614.
120	Farley R A， Fitter A H.Temporal and spatial variation in soil resources in a deciduous woodland［J］.Journal of Ecology，1999，87：688-696.
121	Zuo X A， Zhao X Y， Zhao H L，et al.Spatial pattern and heterogeneity of soil organic carbon and nitrogen in sand dunes related to vegetation change and geomorphic position in Horqin Sandy Land，Northern China ［J］.Environmental Monitoring and Assessment，2010，164：29-42.
122	Zheng Q， Hu Y， Zhang S，et al.Soil multifunctionality is affected by the soil environment and by microbial community composition and diversity［J］.Soil Biology and Biochemistry，2019，136：107521.
123	Ding J， Eldridge D J.Climate and plants regulate the spatial variation in soil multifunctionality across a climatic gradient［J］.Catena，2021，201：105233.
124	Gilbert B， Macdougall A S， Kadoya T，et al.Climate and local environment structure asynchrony and the stability of primary production in grasslands［J］.Global Ecology and Biogeography，2020，29：1177-1188.
125	宋兆斌，辛智鸣，朱雅娟.内蒙古荒漠-草原过渡带灌木群落特征［J］.中国沙漠，2022，42（2）：104-112.
126	Rillig M C， van der Heijden M G A， Berdugo M，et al.Increasing the number of stressors reduces soil ecosystem services worldwide［J］.Nature Climate Change，2023，13（5）：478-483.
127	Yang G， Ryo M， Roy J，et al.Multiple anthropogenic pressures eliminate the effects of soil microbial diversity on ecosystem functions in experimental microcosms［J］.Nature Communications，2022，13：4260.
128	Zhou X， Gu X， Smaill S J.Rethinking experiments that explore multiple global change factors［J］.Trends in Ecology & Evolution，2023，38（5）：399-401.
129	Hong P， Schmid B， Laender De F，et al.Biodiversity promotes ecosystem functioning despite environmental change［J］.Ecology Letters，2022，25：555-569.
130	Wagg C， Schlaeppi K， Banerjee S，et al.Fungal-bacterial diversity and microbiome complexity predict ecosystem functioning［J］.Nature Communications，2019，10：4841.
131	Eisenhauer N， Hines J， Isbell F，et al.Plant diversity maintains multiple soil functions in future environments［J］.Elife，2018，7：e41228.

指标内容	指标类型	养分储量	养分状态	养分转化率	生态过程
养分循环	碳循环	有机碳^*、土壤微生物碳…	可溶性有机碳^*…	α葡糖苷酶^、β葡糖苷酶^、β木糖苷酶^、纤维素二糖水解酶^…	土壤呼吸、CO₂通量、凋落物分解速率、代谢熵…
	氮循环	土壤全氮^*、土壤微生物氮…	铵态氮^、硝态氮^…	β-1, 4-N-乙酰氨基葡萄糖苷酶^、亮氨酸氨基肽酶^…	CH₄通量、N₂O通量…
	磷循环	土壤全磷^*、土壤微生物磷…	速效磷^*…	酸/中/碱性磷酸酶^*…
	硫循环	土壤硫含量…		硫酸酯酶…
水分涵养	土壤水稳性团聚体、土壤持水量、土壤含水量、土壤水力传导度…
物理性质	pH、土壤团聚体、孔隙度、阳离子交换量…

指标内容	指标类型	养分储量	养分状态	养分转化率	生态过程
养分循环	碳循环	有机碳^*、土壤微生物碳…	可溶性有机碳^*…	α葡糖苷酶^、β葡糖苷酶^、β木糖苷酶^、纤维素二糖水解酶^…	土壤呼吸、CO₂通量、凋落物分解速率、代谢熵…
	氮循环	土壤全氮^*、土壤微生物氮…	铵态氮^、硝态氮^…	β-1, 4-N-乙酰氨基葡萄糖苷酶^、亮氨酸氨基肽酶^…	CH₄通量、N₂O通量…
	磷循环	土壤全磷^*、土壤微生物磷…	速效磷^*…	酸/中/碱性磷酸酶^*…
	硫循环	土壤硫含量…		硫酸酯酶…
水分涵养	土壤水稳性团聚体、土壤持水量、土壤含水量、土壤水力传导度…
物理性质	pH、土壤团聚体、孔隙度、阳离子交换量…

[1]	黄梦真, 鲁瑞洁, 赵瑾, 马罗. 柴达木盆地典型风蚀区土壤质量评价[J]. 中国沙漠, 2023, 43(3): 199-209.
[2]	鲍婧婷, 孙靖尧, 王进. 生物土壤结皮中微生物群落特征综述[J]. 中国沙漠, 2022, 42(6): 33-43.
[3]	陈云, 李玉强, 王旭洋, 姚彩萍, 牛亚毅. 中国生态脆弱区全球变化风险及应对技术途径和主要措施[J]. 中国沙漠, 2022, 42(3): 148-158.
[4]	马良, 朱再春, 曾辉. NPP评估过程模型应用研究进展[J]. 中国沙漠, 2017, 37(6): 1250-1260.
[5]	王芳;肖洪浪;苏永中;范桂萍. 临泽边缘绿洲区盐化草甸开垦后土壤质量演变[J]. 中国沙漠, 2011, 31(3): 723-728.
[6]	高天鹏. 菌根真菌对CO2浓度升高和N沉降的响应[J]. 中国沙漠, 2009, 29(1): 131-135.
[7]	靳鹤龄;李明启;苏志珠;董光荣;赵晖. 220 ka以来萨拉乌苏河流域地层磁化率与气候变化[J]. 中国沙漠, 2006, 26(5): 680-686.
[8]	李明启;靳鹤龄;董光荣;张洪;孙忠. 萨拉乌苏河流域微量元素揭示的气候变化[J]. 中国沙漠, 2006, 26(2): 172-179.
[9]	刘树林, 王涛. 浑善达克沙地地区的气候变化特征[J]. 中国沙漠, 2005, 25(4): 557-562.
[10]	陈仁升, 康尔泗, 杨建平, 张济世. 内陆河流域分布式水文模型——以黑河干流山区建模为例[J]. 中国沙漠, 2004, 24(4): 416-424.
[11]	樊恒文, 贾晓红, 张景光, 马凤云, 李新荣. 干旱区土地退化与荒漠化对土壤碳循环的影响[J]. 中国沙漠, 2002, 22(6): 525-533.
[12]	杨小平. 巴丹吉林沙漠及其毗邻地区的景观类型及其形成机制初探[J]. 中国沙漠, 2000, 20(2): 166-170.
[13]	冯兆东, 陈发虎, 张虎才, 马玉贞. 末次冰期-间冰期蒙古高原与黄土高原对全球变化的重要贡献[J]. 中国沙漠, 2000, 20(2): 171-177.
[14]	. 中国沙漠形成演化与气候变化研究——获中国科学院自然科学一等奖[J]. 中国沙漠, 1998, 18(4): 297-298.