(1.1)--Belowground facilitation and tra耕作制度耕作学.pdf
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1、OpinionBelowground facilitation and trait matching:two or three to tango?Rui-Peng Yu,1Hans Lambers,2,3Ragan M.Callaway,4Alexandra J.Wright,5and Long Li1,*Highbiodiversityincreasesecosystemfunctions;however,belowgroundfacilita-tion remains poorly understood in this context.Here,we explore mechanismst
2、hat operate via givingreceiving feedbacks for belowground facilitation.These include direct effects via root exudates,signals,and root trait plasticity,and indirect biotic facilitation via the effects of root exudates on soil biota andfeedback from biota to plants.We then highlight that these two-or
3、 three-waymechanisms must affect biodiversityecosystem function relationships via spe-cific combinations of matching traits.To tango requires a powerful affinity andharmony between well-matched partners,and such matches link belowgroundfacilitation to the effect of biodiversity on function.Such matc
4、hing underpinsapplications in intercropping,forestry,and pasture systems,in which diversitycontributes to greater productivity and sustainability.Biodiversity,ecosystem function,and belowground facilitationMany experiments in natural ecosystems and agroecosystems have shown that experimentallyincrea
5、sed biodiversity enhances various ecosystem functions 14.The preeminent mechanismfor this effectof biodiversityis resource complementarity(see Glossary)and niche partitioning,which occurs when species in diverse polycultures have unique traits and requirements that leadto greater resource capture th
6、an that of any single species in monoculture.However,resourcepartitioning alone does not always explain biodiversityecosystem function(BEF)relation-ships 5,6.In addition to partitioning,facilitation has received increasing attention as anotherimportant mechanism driving BEF,and this occurs when one
7、or more species in polyculturesincrease a functional metric(e.g.,biomass)of one or more other species 3,7,8(Box 1).Todate,most studies interpret facilitation in BEF by calculating relative interaction intensities 9 ordiversity effects 10,for example,complementarity effect(consequences),in which faci
8、litationis framed,often appropriately,as a distinct alternative to resource complementarity.However,this integration does not shed light on mechanisms by which polycultures of species interact,and often cannot separate facilitation from resource complementarity(causes)4,10.More recently,researchers
9、have focused on belowground facilitative mechanisms underlyingpositive BEF relationships 3.For example,it is widely accepted that legumes may facilitateneighbours via dinitrogen(N2)fixation and increasing nitrogen(N)availability in soil 3,4,7.How-ever,beyond N facilitation,belowground facilitation h
10、as received little attention in BEF research.For example,a study showed that 40%more root biomass was produced in polycultures thanpredicted from monocultures,and this was not due to spatial partitioning.They argued thatroot responses were consistent with species-specific root recognition,but this w
11、as not demon-strated 11.Other studies also reported striking decreases in the occurrence of pathogenic fungiin plant polycultures(indirect bioticfacilitation),while the plant-beneficial microbepathogen inter-action was underexplored 12,13.HighlightsBelowground facilitation is a vital mecha-nism unde
12、rlying ecosystem function indiverse plant communities.A bottleneck to linking theory to applica-tion is that interspecific belowgroundfacilitation processes are incompletelyunderstood.Here,weexploredirectand indirectfacil-itation,which requires matching specificbelowground responses between thefacil
13、itated species and the facilitator.Understanding and integrating the pro-cesses of interspecific facilitation helpsto design sustainable diverse plant com-munities with compatible ecosystemfunctions.1Key Laboratory of Plant-Soil Interactions,Ministry of Education,Beijing KeyLaboratory of Biodiversit
14、y and OrganicFarming,College of Resources andEnvironmental Sciences,China AgriculturalUniversity,2 Yuan Ming Yuan West Road,Beijing 100193,PR China2School of Biological Sciences andInstitute of Agriculture,The University ofWestern Australia,35 Stirling Highway,Crawley,Perth,WA 6009,Australia3Nationa
15、l Academy of Agriculture GreenDevelopment,China Agricultural Univer-sity,2 Yuan Ming Yuan West Road,Beijing 100193,PR China4Division of Biological Sciences andInstitute on Ecosystems,University ofMontana,32 Campus Drive,Missoula,MT 59812,USA5Department of Biological Sciences,California State Univers
16、ity Los Angeles,5151 State University Drive,Los Angeles,CA 90032,USA*Correspondence:(L.Li).Trends in Plant Science,Month 2021,Vol.xx,No.xxhttps:/doi.org/10.1016/j.tplants.2021.07.0141 2021 Elsevier Ltd.All rights reserved.Trends inPlant ScienceTRPLSC 2167 No.of Pages 9Recent studies without a focus
17、on BEF have explored the role of belowground direct and indirectfacilitation in plantplant 14,plantmicrobiome 15 and crosskingdom interactions 16,providing opportunities to better conceptualise facilitative interactions that might underlie howbiodiversity increases ecosystem functions.Here,we synthe
18、sise these recent advances.Wealso highlight the importance of intimate matches among interacting species that enhance notonly direct abiotic facilitation via root exudates,signalling molecules,and species-specificroot phenotypic plasticity,but also indirect biotic facilitation via plantmicrobiome co
19、mbina-tions.As tango requires a powerful affinity and harmony between well-matched partners,so dothese belowgroundprocesses.However,tango is officially and logisticallylimited to two partners,but facilitative interactions that drive BEF are not.A deeper understanding of how theseprocesses allow faci
20、litation,and matches among species,can advance conceptual ecologicaltheory and bridge the gap between biodiversity theory and application.How does belowground facilitation work?First,we acknowledge that facilitation in highly diverse plant communities may operate via emer-gent effects associated wit
21、h unique properties of a more diverse community(direct abioticfacilitation orindirect biotic facilitationviasoilbiota)7,17,18(Box1).Forinstance,higherdiversityplant communities may have more complete occupation of belowground root space and this canlead to increased soil porosity and enhanced aerati
22、on of soils during a flooding event;as a result,all constituent species benefit from living in a higher diversity soil during a flood 17.Mycorrhizalfungi-and beneficial microbe-mediated indirect abiotic facilitation may play universal roles inincreasing plant nutrient uptake via common mycorrhizal n
23、etworks(e.g.,N and P),andbenefiting their hosts by enhancing their defence against pathogens in diverse plant communities13,1921.However,for the purpose of this review,we focus on specific two-and three-wayinteractions between neighbouring species(Box 1).In particular,we explore direct abiotic facil
24、ita-tion via nutrient-mobilising root exudates and signalling molecules,direct facilitation due to traitplasticity in response to neighbours,and indirect facilitation via rootsoil biota interactions.Direct facilitation via root exudates and signalling moleculesNutrient-mobilising root exudates and s
25、ignalling molecules may mediate belowground plantplant interactions via direct abiotic facilitation(Figure 1).In faba bean(Vicia faba)/maize(Zeamays)intercrops,rootexudatesfrommaizeincludeflavonoids,whicharesignallingcompoundsfor the rhizobial symbiosis with legume plants 22.These exudates increase
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