doi:10.11838/sfsc.1673-6257.23539 有机肥增强植物磷素吸收利用机制研究进展 戈应同，王宇蕴，徐 翔，李 兰，高江飞，徐 智 (云南农业大学资源与环境学院，云南 昆明 650201) 摘要:磷素对植物的正常生长发育极为重要，但由于土壤中固有的磷形态生物有效性低，植物磷素吸收能力弱，导致磷素利用率低。有机肥施用能有效增强植物的磷素吸收利用能力，但其背后的原因错综复杂。综述了有机肥施用如何增加土壤磷素有效性以及增强植物对磷素的吸收能力的机制，并重点强调了在此过程中微生物的重要作用。有机肥通过改变土壤吸附特性、团聚体形成以及微生物代谢活动增加土壤磷素有效性；通过调节根际微生物与植物根系间的关系促进根系生长、改变根系形态，从而增强磷素吸收能力，是以“碳”为核心的植物- 微生物- 土壤互作的连续体。 关键词:有机肥；磷吸收能力；磷素有效性；植物- 微生物- 土壤连续体 收稿日期:2023-09-05；录用日期:2023-11-04 基金项目:国家重点研发项目(2022YFD1901500，2022YFD1901504)； 国家自然科学基金(32160744)； 云南省自然科学基金(202101AT070252)； 云南省高层次人才培养计划(YNWR-QNBJ-2019-249)。 作者简介:戈应同(1999-)，硕士研究生，主要研究方向为养分资源管理。E-mail: gyt991126@126.com。 通讯作者:徐智，E-mail: xuzhi9910@126.com。 参考文献: [1] George TS，Gi les CD，M enezes-blackburn，e tal.Org anic phosphorus in the terrestrialenvironment:a perspective on the state of the artand future priorities[J].Plant andSoil，2018， 427(1/2):191-208. [2] He XJ，AugustoL，Goll DS，etal.Glo bal patterns and drivers of soil total phosphorusconcentration[J].Earth System Science Data，2021，13(12):5831-5846. [3] Atere CT，GeT，Z huZ，etal.As similate allocation by rice and carbon stabilisation insoil: effect of water management and phosphorus fertilisation[J].Pla ntSoil，2019，445:153-167. [4] EtesamiH.Enhanced phosphorus fertilizer use efficiency with microorganisms[M]// MeenaR.(eds) Nutrient dynamics for sustainablecrop production.Spri nger,Singapore，2020:215-245. [5] HonvaultN，HoubenD，NobileC，etal.T radeoffs among phosphorus-acquisition root traits of crop species for agroecological intensification[J].Pla nt andSoil，2020，461:137-150. [6] Raven JA，La mbersH，Smi th SE，e tal.C osts of acquiring phosphorus byvascular land plants: patterns and implications for plantcoexistence[J].NewPhytologist，2018，217(4): 1420-1427. [7] ZhiH，HongB，QiS，etal.Tradeoffs among rootmorphology， exudation and mycorrhizal symbioses for phosphorus-acquisition strategiesof 16 crop species.[J].New Phytologist，2019，223 (2):882-895. [8] WangC，LukasT，Dippold MA，etal.Reductive dissolution of iron phosphate modifies rice root morphology in phosphorus-deficient paddy soils[J].Soil Biology and Biochemistry，2023， 177:108904. [9] LambersH，Raven JA，Shaver GR，etal.P lant nutrient- acquisition strategieschange with soil age[J].Trends in Ecology & Evolution，2007，23(2):95-103. [10] Mai WX，Xue XR，FengG，etal.Arbuscular mycorrhizal fungi-15-Fold enlargement of the soil volume of cotton roots for phosphorus uptake in intensive planting conditions[J].EuropeanJournal of Soil Biology，2019，90:31-35. [11]Smith S E，AndersonI C，Smith FA.Mycorrhizal associations and phosphorusacquisition:from ells toecosystems，PhosphorusMetabolism in Plants[M].Wiley-Blackwell，2015:409-439. [12] ZhouYP，SarkerU，NeumannG，etal.The LaCEP1 peptide modulates cluster root morphology in Lupinusalbus[J].Physiologia Plantarum，2018，166(2):525-537. [13] Shi JM，StrackD，Albornoz FE，etal.Differences in investment and functioning of cluster roots account for different distributions of Banksiaattenuata and B. sessilis，with contrasting lifehistory[J].Plant andSoil:An International Journal onPlant-Soil Relationships，2020，447(2):85-98. [14] Wang BS，WangY，SunY，etal.Watermelon responds to organic fertilizer by enhancing root-associated acid phosphatase activity toimprove organic phosphorus utilization[J].Journal ofPlant Physiology，2022，279:153838. [15] ErroJ，Zamarreno AM，Garcia-mina JM.Ability of various water-insoluble fertilizers to supply available phosphorus in hydroponics to plant species with diverse phosphorus-acquisitionefficiency:Involvement of organic acid accumulation in plant tissues and rootexudates[J].Journal of Plant Nutrition and SoilScience，2010，173(5):772-777. [16] ThonarC，Lekfeldt J DS，CozzolinoV，etal.Potential of three microbial bio-effectors to promote maize growth and nutrient acquisition from alternative phosphorous fertilizers in contrastingsoils[J].Chemical and Biological Technologiesin Agriculture，2017，4(1):7. [17] Alori ET，Glick BR，Babalola OO.Microbial phosphorus solubilization and its potential for usein sustainable agriculture[J]. Frontiersin Microbiology，2017，8:971. [18] LiuD.Root developmental responses to phosphorusnutrition[J]. Journal of Integrative PlantBiology，2021，63(6):1065-1090. [19] ZhaoY，Li YL，YangF.Critical review on soil phosphorus migration and transformation under freezing-thawing cycles and typical regulatorymeasurements[J].Science of the TotalEnvironment，2021，751:141614. [20] ZhuJ，LiM，WhelanM.Phosphorus activators contribute tolegacy phosphorus availability in agricultural soils: areview[J]. Scienceof the Total Environment，2018，612:522-537. [21] MengelK.Agronomic measures for better utilization of soil and fertilizer phosphates[J].European Journal ofAgronomy，1997，7(1-3):221-233. [22] Rubaek GH，KristensenK，Olesen SE，etal.Phosphorus accumulation and spatial distribution in agricultural soils inDenmark[J].Geoderma，2013，209:241-250. [23]严玉鹏，王小明，刘凡，等 .有机磷与土壤矿物相互作用及其环境效应研究进展[J].土壤学报，2019，56(6):1290-1299. [24] Ding WL，Cong WF，LambersH.Plant phosphorus-acquisition and -use strategies affect soil carboncycling[J].Trends in Ecology & Evolution，2021，36(10):899-906. [25] Bello SK.An overview of themorphological，genetic and metabolic mechanisms regulating phosphorus efficiency via roottraits in soybean[J].Journal of Soil Scienceand Plant Nutrition，2021，21(2):1013-1029. [26] Weng LP，Vega FA，Van Riemsdijk WH.Competitive and synergistic effects in pH dependent phosphate adsorptionin soils: LCDmodeling[J].Environmental Science andTechnology，2011，45(19):8420-8428. [27]章永松，林咸永，倪吾钟.有机肥对土壤磷吸附 -解吸的直接影响[J].植物营养与肥料学报，1996(3):200-205. [28] Li RL，Zhang SR，ZhangM，etal.Phosphorus fractions and adsorption-desorption in aggregates in coastal saline-alkaline paddy soil with organic fertilizerapplication[J].Journal of SoilsandSediments，2021，21:3084-3097. [29] Nobile CM，Bravin MN，BecquerT，etal.Phosphorus sorption and availability in an andosol after a decade of organic or mineral fertilizerapplications:Importance of pH and organic carbon modifications in soil as compared to phosphorus accumulation[J].Chemosphere，2020，239(C):124709. [30] FischerP，PthigR，VenohrM.The degree of phosphorus saturationof agricultural soils inGermany:current and future risk of diffuse P loss and implications for soil P management in Europe[J].Science of the TotalEnvironment，2017，599:1130-1139. [31] Jin JW，Fang YY，HeS，etal.Improved phosphorus availability and reduced degree of phosphorus saturation by biochar-blended organic fertilizer addition to agricultural fieldsoils[J].Chemosphere，2023，317:137809. [32]Li KJ，Bi QF，Liu XP，etal.Unveiling the role of dissolved organic matter on phosphorus sorption and availability in a 5-year manure amended paddysoil[J].The Science of the TotalEnvironment，2022，838(P1):155892. [33]PizzeghelloD，BertiA，NardiS，etal.Phosphorus forms and P-sorption properties in three alkaline soils after long-term mineral and manure applications in north-easternItaly[J].Agriculture， Ecosystemsand Environment，2011，141(1):58-66. [34]Barrow NJ.The effects of pH on phosphate uptake from the soil[J].Plant andSoil，2017，410(1/2):401-410. [35] Yan ZJ，ChenS，DariB，etal.Phosphorus transformation response to soil properties changes induced by manure application in a calcareoussoil[J].Geoderma，2018，322:163-171. [36] Zhang YJ，GaoW，Luan HA，etal.Long-term organic substitution management affects soil phosphorus speciation and reduces leaching in greenhouse vegetable production[J].Journal of CleanerProduction，2021，327:129464. [37]魏朝富，谢德体，李保国.土壤有机无机复合体的研究进展[J].地球科学进展，2003(2):221-227.[38]周亦靖，牛犇，李欢，等.长期施肥对旱地红壤微团聚体磷素有效性的影响[J].土壤通报，2023，54(1):89-99. [39] TinasheM，ManojM，HarrietB，etal.Preferential wheat(Triticum aestivum. L cv.Fielder)root growth in different sizedaggregates[J].Soil & TillageResearch，2021，212:105054.[40] Yang JJ，Shi JX，Jiang LH，etal.Co-occurrence network in core microorganisms driving the transformation of phosphorous fractionations during phosphorus recovery product used as soil fertilizer [J].Science of the TotalEnvironment，2023，871:162081. [41] Richardson AE，Simpson RJ.Soil microorganisms mediatingphosphorus availability[J].Plant Physiology，2011，156(3): 989-996. [42] Alori ET，Glick BR，Babalola OO.Microbial phosphorus solubilization and its potential for usein sustainable agriculture[J]. Frontiersin Microbiology，2017，8:971. [43] ColemanD，ReidC，ColeC.Biological strategies of nutrient cycling in soilsystems[1]. Advances inEcologicalResearch，1983，13:1-55. [44]Huang LM，Jia XX，Zhang GL，etal.Soil organic phosphorus transformation during ecosystemdevelopment:areview[J].Plant andSoil，2017，417(1/2):17-42. [45] BiQF，Li KJ，Zheng BX，etal.Partial replacement ofinorganic phosphorus(P)by organic manure reshapes phosphate mobilizing bacterial community and promotes P bioavailability in apaddy soil[J].Science of the Total Environment，2020，703(C):134977. [46] WeiK，Chen ZH，JiangN，etal.Effects of mineral phosphorus fertilizer reduction and maize straw incorporation onsoil phosphorusavailability，acid phosphataseactivity，and maize grain yield in northeastChina[J].Archives of Agronomy andSoil Science，2021，67(1):66-78. [47] Enebe MC，Babalola OO.The Influence of soil fertilization on the distribution and diversity of phosphorus cycling genes and microbes community of maize rhizosphere using shotgun metagenomics[J].Genes，2021，12(7):1022. [48] Zhang YJ，GaoW，Luan HA，etal.Effects of a decade of organic fertilizer substitution on vegetable yield and soil phosphoruspools，phosphataseactivities，and the microbial community in a greenhouse vegetable productionsystem[J].Journal of IntegrativeAgriculture，2022，21(7):2119-2133. [49]Hu JL，Lin XG，Wang JH，etal.Population size and specific potential of P-mineralizing and -solubilizing bacteria under long-term P-deficiency fertilization in a sandy loamsoil[J].Pedobiologia，2009，53(1):49-58. [50] GyaneshwarP，Kumar GN，Parekh LJ，etal.Poole. Role of soil microorganisms in improving P nutritionof plants[J].PlantandSoil，2002，245(1):133-143. [51]OburgerE，Jones DL，Wenzel WW.Phosphorus saturation and pH differentially regulate the efficiency of organic acid anion-mediated P solubilizationmechanisms in soil[J].Plant andSoil，2011，341(1/2):363-382. [52] Jones DL，OburgerE.Solubilization of phosphorus by soilmicroorganisms[M]//B nemann EK.Phosphorus inaction: biological processes in soil phosphorus cycling.Berlin:Springer-Verlag Berlin，2011. [53]Rawat P，DasS，ShankhdharD，etal.Phosphate-solubilizingmicroorganisms:mechanism and their role in phosphate solubilization anduptake[J].Journal of Soil Science and PlantNutrition，2020，21(1):49-68. [54]Zhang YJ，GaoW，MaL，etal.Long-term partial substitution of chemical fertilizer by organic amendments influences soil microbial functional diversity of phosphorus cycling and improves phosphorusavailability in greenhouse vegetable production[J].Agriculture，Ecosystemsand Environment，2023，341:108193. [55] Wu JS，HuangM，Xiao HA，etal.Dynamics in microbial immobilization and transformations of phosphorus in highly weathered subtropical soil following organicamendments[J].Plant andSoil，2007，290(1/2):333-342. [56]KunitoT，HirutaN，MiyagishiY，etal.Changes in phosphorus fractions caused by increased microbial activity in forest soil in a short-term incubationstudy[J].Chemical Speciation andBioavailability，2018，30(1):9-13. [57] KathuriaS，Martiny AC.Prevalence of a calcium-based alkaline phosphatase associated with the marine cyanobacterium prochlorococcus and other oceanbacteria[J].EnvironmentalMicrobiology，2011，13(1):74-83. [58] HuangYL，Dai ZM，Lin JH，etal.Labile carbon facilitated phosphorus solubilization as regulated by bacterial and fungal communities in zeamays[J].Soil Biology andBiochemistry，2021a，163:108465. [59] Huang YL，Dai ZM，Lin JH，etal.Contrasting effects of carbon source recalcitrance on soil phosphorus availability and communities of phosphorus solubilizingmicroorganisms[J].Journal of EnvironmentalManagement，2021b，298:113426. [60] ZhangL L，Niu J F，Lu XW，etal.Dosage effects of organic manure on bacterial community assemblage and phosphorus transformation profiles in greenhousesoil[J].Frontiers inMicrobiology，2023，14:1188167. [61] LambersH.Phosphorus acquisition and utilization in plants[J].Annual Reviewof Plant Biology，2021，73:17-42. [62] Bzdyk RM，OlchowikJ，StudnickiM，etal.The impact of effectivemicroorganisms(EM)and organic and mineral fertilizers on the growth and mycorrhizal colonization of fagus sylvatica and quercus robur seedlings in a bare-root nurseryexperiment[J].Forests，2018，9(10):597. [63]SongG，ChenR，XiangW，etal.Contrasting effects of long-term fertilization on the community of saprotrophic fungi and arbuscular mycorrhizal fungi in a sandy loamsoil[J].Plant，Soil and Environment，2015，64(3):127-136. [64] Li XX，Zeng RS，LiaoH.Improving crop nutrient efficiency through root architecture modifications[J].Journal of IntegrativePlant Biology，2016，58(3):193-202. [65]WipfD，KrajinskiF，Van TuinenD，etal.Trading on the arbuscular mycorrhizamarket:from arbuscules to commonmycorrhizal networks[J].The Newphytologist，2019，223(3): 1127-1142. [66] ParniskeM.Arbuscularmycorrhiza:the mother of plant rootendosymbiosis[J].Nature ReviewsMicrobiology，2008，6(10):763-775. [67] Cavagnaro TR.Impacts of compost application on the formationand functioning of arbuscular mycorrhizas[J].Soil Biology andBiochemistry，2014，78:38-44. [68] QinH，Lu KP，Strong PJ，etal.Long-term fertilizer application effects on thesoil，root arbuscular mycorrhizal fungi and community composition in rotationagriculture[J].AppliedSoil Ecology，2015，89:35-43. [69]Hammer EC，NasrH，WallanderH.Effects of different organic materials and mineral nutrients on arbuscular mycorrhizal fungal growth in a mediterranean salinedryland[J].Soil Biology andBiochemistry，2011，43(11):2332-2337. [70]Linderman RG，Davis EA.Evaluation of commercial inorganic and organic fertilizer effects on arbuscular mycorrhizae formed by glomusintraradices[J].HortTechnology，2004，14(2):196-202. [71] Bello SK.An overview of themorphological，genetic and metabolic mechanisms regulating phosphorus efficiency via roottraits in soybean[J].Journal of Soil Scienceand Plant Nutrition，2021，21(2):1013-1029. [72]Yang QL，Zheng FX，Jia XC，etal.The combined application of organic and inorganic fertilizers increases soil organic matter and improves soil microenvironment in wheat-maizefield[J].Journal of Soils andSediments，2020，20(5):2395-2404. [73] Falk KG，Jubery TZ，O’Rourke JA，etal.Soybean root system architecture traitstudy throughgenotypic，phenotypic， andshape-based clusters[J].Plant Phenomics，2020，2020: 1925495. [74] Dobbss LB，Canellas LP，Olivares FL，etal.Bioactivity of chemically transformed humic matter from vermicompost on plant rootgrowth[J].Journal of Agricultural and FoodChemistry，2010，58(6):3681-3688. [75] Li SJ，Wang BS，WangY，etal.Soluble organic nutrients induce ClaPhys expression to enhance phytase activity of watermelonroots[J].Annals ofAppliedBiology，2022，181(1):80-92. [76] Kang YL，Ma YW，An XR，etal.Effects on the root morphology and mircostructure of youngpear(Pyrus pyrifolia) tree by split-root supply of bioorganic and chemical fertilizer[J].Rhizosphere，2022，22:100504. [77]Kudoyarova GR，Vysotskaya LB，Arkhipova TN，etal.Effect of auxin producing and phosphate solubilizing bacteria on mobility ofsoil phosphorus，growthrate，and P acquisition by wheat plants[J].Acta Physiologiae Plantarum，2017，39(11):253. [78] Li YB，Liu XM，Hao TY，etal.Colonization and maize growth promotion induced by phosphate solubilizing bacterial isolates[J].International Journal of MolecularSciences，2017，18(7):1253. [79]Pathan SI，Větrovsk yT，GiagnoniL，etal.Microbial expression profiles in the rhizosphere of two maize lines differing in N useefficiency[J].Plant andSoil，2018，433(1/2):401-413. [80]HaidarB，FerdousM，FatemaB，etal.Population diversity of bacterial endophytes fromjute( Corchorus olitorius)and evaluation of their potential role asbioinoculants[J].MicrobiologicalResearch，2018，208:43-53. [81] Bal HB，NayakL，DasS，etal.Isolation of ACC deaminase producing PGPR from rice rhizosphere and evaluating their plant growth promoting activity under saltstress[J].Plant andSoil，2013，366(1/2):93-105. [82] Wang JF，Zhang YQ，LiY，etal.Endophytic microbes Bacillus sp. LZR216-regulated root development is dependent on polar auxin transport in Arabidopsisseedlings[J].Plant CellReports，2015，34(6):1075-1087. [83] BargazA，ElhaissoufiW，KhourchiS，etal.Benefits of phosphate solubilizing bacteria on belowground crop performance for improved crop acquisitionof phosphorus[J].MicrobiologicalResearch，2021，252:126842. [84]于雯霏，王佩佩，刘俊娥，等.黄土高原典型植被根系对土壤团聚体及其有机碳组分的影响[J].水土保持学报，2023，37(6):246-254. [85]谭文峰，许运，史志华，等.胶结物质驱动的土壤团聚体形成过程与稳定机制[J].土壤学报，2023，60(5):1000-1007. Research progress on the mechanism of phosphorus absorption and utilization enhanced by organic fertilizer GE Ying-tong，WANG Yu -yun，XU Xiang，LI Lan，GAO Jiang -fei，XU Zhi (College of Resources and Environmental Science，Yunnan AgriculturalUniversity，Kunming Yunnan650201) Abstracts:Phosphorus is extremely important for the normal growth and development ofplants，but due to the low bioavailability of phosphorus forms inherent in thesoil，the weak phosphorus uptake capacity of plants leads to low phosphorus utilization. Organic fertilizer application can effectively enhance the phosphorus uptake and utilization capacity ofplants，but the reasons behind this are intricate. This review examines the mechanisms of how organic fertilizer applicationincreases soil phosphorus effectiveness and enhances plant phosphorus uptakecapacity，and highlights the important role of microorganisms in this process. Organic fertilizers increase soil phosphorus effectiveness by altering soil adsorptionproperties， aggregate formation and microbial metabolicactivities；promote root growth and enhance phosphorus uptake by regulating the relationship between inter-root microorganisms and plantroots；and alter root morphology to enhance phosphorusuptake， which is a continuum of plant-microorganism-soil interactions with“carbon”as the nucleus.