Contrasting residual effects of different biochar types on maize nitrogen uptake, biomass accumulation, water and nitrogen use efficiency under alternate partial root-zone drying irrigation

Abstract

Although the immediate benefits of biochar in enhancing nitrogen cycling and crop productivity are well documented, its residual effects across different biochar types and irrigation regimes over successive growing seasons have not been fully elucidated. Here, we assessed the residual effects of softwood (SWB) and wheat-straw (WSB) biochar on soil–plant nitrogen (N) dynamics and maize (Zea mays L.) productivity over two growing seasons following a one-time application. Experiments were conducted in 2021 and 2022 under full (FI), deficit (DI), and alternate partial root-zone drying (APRI) irrigation. In both years, despite limited changes in water consumption and total N uptake, WSB-APRI combination improved total dry biomass (+ 13.5%), harvest index (+ 4.4%), water use efficiency (+ 26.7%), and N use efficiency (+ 10.3%). These improvements were linked to enhanced microbial activity (+ 26.8–51.2%) and soil N availability (+ 4.8–13.2%), which stimulated root growth (+ 7.4–22.7%) and N uptake (+ 7.0–17.8%) under water stress. However, under reduced irrigation in 2021, SWB markedly suppressed microbial respiration (− 42.4%) and N availability (− 29.2%), which in turn led to compromised crop performance, particularly under DI. Partial least squares path modeling revealed that microbial activity and root traits indirectly affected maize water and N use efficiency by influencing water consumption, N uptake, and biomass accumulation. Notably, excessive N uptake reduced N use efficiency, whereas biomass accumulation enhanced it. Considering the residual effects of biochar, APRI combined with WSB offers a promising approach to continuously enhance water-nitrogen coordination and maintain maize productivity under limited irrigation.

Keywords

Biochar / Alternate partial root-zone drying irrigation / Root traits / Nitrogen availability / Water use efficiency / Nitrogen use efficiency

Highlight

Highlights

•	Residual effects of softwood and wheat-straw biochar on soil-plant N dynamics were evaluated across two growing seasons.
•	Soil N availability and root morphology were mainly driven by soil C:N stoichiometry and microbial activity.
•	Residual softwood biochar suppressed soil N availability and reduced maize productivity under water-limited conditions.
•	Wheat-straw biochar combined with alternate partial root-zone drying irrigation improved both N and water use efficiencies.

References

[1] Abrisqueta J, Mounzer O, Álvarez S, Conejero W, García-Orellana Y, Tapia L, Vera J, Abrisqueta I, Ruiz-Sánchez M. Root dynamics of peach trees submitted to partial root-zone drying and continuous deficit irrigation. Agric Water Manage, 2008, 95: 959-967.

[2] Akhtar SS, Li G, Andersen MN, Liu F. Biochar enhances yield and quality of tomato under reduced irrigation. Agric Water Manag, 2014, 138: 37-44.

[3] Akhtar SS, Andersen MN, Liu F, Rizwan M. Residual effects of biochar on improving growth, physiology and yield of wheat under salt stress. Agric Water Manag, 2015, 158: 61-68.

[4] Ameloot N, Sleutel S, Das KC, Kanagaratnam J, De Neve S. Biochar amendment to soils with contrasting organic matter level: effects on N mineralization and biological soil properties. GCB Bioenergy, 2015, 7: 135-144.

[5] Archbold DD, MacKown CT. Seasonal and cropping effects on total and fertilizer nitrogen use in June-bearing and day-neutral strawberries. J Am Soc Hortic Sci, 1995, 120: 403-408.

[6] Baronti S, Magno R, Maienza A, Montagnoli A, Ungaro F, Vaccari FP. Long-term effect of biochar on soil-plant water relation and fine roots: results after 10 years of vineyard experiment. Sci Total Environ, 2022, 851. 158225

[7] Beesley L, Moreno-Jiménez E, Gomez-Eyles JL, Harris E, Robinson B, Sizmur T. A review of biochars’ potential role in the remediation, revegetation and restoration of contaminated soils. Environ Pollut, 2011, 159: 3269-3282.

[8] Birch H. The effect of soil drying on humus decomposition and nitrogen availability. Plant Soil, 1958, 10: 9-31.

[9] Bornø ML, Müller-Stöver DS, Liu F. Contrasting effects of biochar on phosphorus dynamics and bioavailability in different soil types. Sci Total Environ, 2018, 627: 963-974.

[10] Brewer CE, Hu YY, Schmidt-Rohr K, Loynachan TE, Laird DA, Brown RC. Extent of pyrolysis impacts on fast pyrolysis biochar properties. J Environ Qual, 2012, 41: 1115-1122.

[11] BRC U (2009) UK Biochar Research Centre. Reducing and removing CO2 while improving soils: a significant and sustainable response to climate change.

[12] Butnan S, Deenik JL, Toomsan B, Antal MJ, Vityakon P. Biochar characteristics and application rates affecting corn growth and properties of soils contrasting in texture and mineralogy. Geoderma, 2015, 237: 105-116.

[13] Casamali B, van Iersel MW, Chavez DJ. Nitrogen partitioning in young “Julyprince” peach trees grown with different irrigation and fertilization practices in the southeastern United States. Agronomy, 2021, 11: 350.

[14] Chaves MM, Maroco JP, Pereira JS. Understanding plant responses to drought—from genes to the whole plant. Funct Plant Biol, 2003, 30: 239-264.

[15] Chen X, Yang SH, Jiang ZW, Ding J, Sun X. Biochar as a tool to reduce environmental impacts of nitrogen loss in water-saving irrigation paddy field. J Clean Prod, 2021, 290. 125811

[16] Chen Y, Du Z, Weng Z, Sun K, Zhang Y, Liu Q, Yang Y, Li Y, Wang Z, Luo Y. Formation of soil organic carbon pool is regulated by the structure of dissolved organic matter and microbial carbon pump efficacy: a decadal study comparing different carbon management strategies. Glob Change Biol, 2023, 29: 1321-1336.

[17] Cheng H, Jones DL, Hill P, Bastami MS, Tu C. Influence of biochar produced from different pyrolysis temperatures on nutrient retention and leaching. Arch Agron Soil Sci, 2018, 64: 850-859.

[18] Chew J, Joseph S, Chen G, Zhang Y, Zhu L, Liu M, Taherymoosavi S, Munroe P, Mitchell DR, Pan G. Biochar-based fertiliser enhances nutrient uptake and transport in rice seedlings. Sci Total Environ, 2022, 826. 154174

[19] Comas LH, Becker SR, Cruz VMV, Byrne PF, Dierig DA. Root traits contributing to plant productivity under drought. Front Plant Sci, 2013, 4: 442.

[20] Cong M, Hu Y, Sun X, Yan H, Yu G, Tang G, Chen S, Xu W, Jia H. Long-term effects of biochar application on the growth and physiological characteristics of maize. Front Plant Sci, 2023, 14: 1172425.

[21] Delgado-Baquerizo M, Reich PB, Khachane AN, Campbell CD, Thomas N, Freitag TE, Abu Al-Soud W, Sørensen S, Bardgett RD, Singh BK. It is elemental: soil nutrient stoichiometry drives bacterial diversity. Environ Microbiol, 2017, 19: 1176-1188.

[22] Dieleman WIJ, Vicca S, Dijkstra FA, Hagedorn F, Hovenden MJ, Larsen KS, Morgan JA, Volder A, Beier C, Dukes JS. Simple additive effects are rare: a quantitative review of plant biomass and soil process responses to combined manipulations of CO₂ and temperature. Glob Change Biol, 2012, 18: 2681-2693.

[23] Dodd IC. Rhizosphere manipulations to maximize ‘crop per drop’ during deficit irrigation. J Exp Bot, 2009, 60: 2454-2459.

[24] Dodd IC, Theobald JC, Bacon MA, Davies WJ. Alternation of wet and dry sides during partial root-zone drying irrigation alters root-to-shoot signalling of abscisic acid. Funct Plant Biol, 2006, 33: 1081-1089.

[25] Dodd IC, Puértolas J, Huber K, Pérez-Pérez JG, Wright HR, Blackwell MS. The importance of soil drying and re-wetting in crop phytohormonal and nutritional responses to deficit irrigation. J Exp Bot, 2015, 66: 2239-2252.

[26] Dong T, Duan B, Zhang S, Korpelainen H, Niinemets Ü, Li C. Growth, biomass allocation and photosynthetic responses are related to intensity of root severance and soil moisture conditions in the plantation tree Cunninghamia lanceolata. Tree Physiol, 2016, 36: 807-817.

[27] Du Z, Weng E, Jiang L, Luo Y, Xia J, Zhou X. Carbon–nitrogen coupling under three schemes of model representation: a traceability analysis. Geosci Model Dev, 2018, 11: 4399-4416.

[28] Eissenstat DM, Yanai RD. The ecology of root lifespan. Adv Ecol Res, 1997, 27: 1-60.

[29] 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. Biogeochemistry, 2016, 129: 21-36.

[30] FAOIFADUNICEFWFPWHOThe state of food security and nutrition in the world 2020: transforming food systems for affordable healthy diets, 2020RomeFAO.

[31] Freschet GT, Cornelissen JHC, van Logtestijn RSP, Aerts R. Evidence of the ‘plant economics spectrum’ in a subarctic flora. J Ecol, 2010, 98: 362-373.

[32] Gao S, Wang D, Dangi SR, Duan Y, Pflaum T, Gartung J, Qin R, Turini T. Nitrogen dynamics affected by biochar and irrigation level in an onion field. Sci Total Environ, 2020, 714. 136432

[33] Garnett T, Appleby MC, Balmford A, Bateman IJ, Benton TG, Bloomer P, Burlingame B, Dawkins M, Dolan L, Fraser D. Sustainable intensification in agriculture: premises and policies. Science, 2013, 341: 33-34.

[34] Gaskin JW, Steiner C, Harris K, Das KC, Bibens B. Effect of low-temperature pyrolysis conditions on biochar for agricultural use. Trans ASABE, 2008, 51: 2061-2069.

[35] Gul S, Whalen JK, Thomas BW, Sachdeva V, Deng H. Physico-chemical properties and microbial responses in biochar-amended soils: mechanisms and future directions. Agric Ecosyst Environ, 2015, 206: 46-59.

[36] Guo L, Bornø ML, Niu W, Liu F. Biochar amendment improves shoot biomass of tomato seedlings and sustains water relations and leaf gas exchange under different irrigation and nitrogen regimes. Agric Water Manag, 2021, 245. 106580

[37] Hair JFA primer on partial least squares structural equation modeling (PLS-SEM), 2014Thousand OaksSAGE Publications.

[38] Hodge A, Fitter AH. Microbial mediation of plant competition and community structure. Funct Ecol, 2013, 27: 865-875.

[39] Hou J, Liu X, Zhang J, Wei Z, Ma Y, Wan H, Liu J, Cui B, Zong Y, Chen Y. Combined application of biochar and partial root-zone drying irrigation improves water relations and water use efficiency of cotton plants under salt stress. Agric Water Manag, 2023, 290. 108584

[40] Hou J, Wan H, Liang K, Cui B, Ma Y, Chen Y, Liu J, Wang Y, Liu X, Zhang J. Biochar amendment combined with partial root-zone drying irrigation alleviates salinity stress and improves root morphology and water use efficiency in cotton. Sci Total Environ, 2023, 904. 166978

[41] Hou J, Zhang J, Liu X, Ma Y, Wei Z, Wan H, Liu F. Effect of biochar addition and reduced irrigation regimes on growth, physiology and water use efficiency of cotton plants under salt stress. Ind Crops Prod, 2023, 198. 116702

[42] Ibrahim MM, Tong C, Hu K, Zhou B, Xing S, Mao Y. Biochar-fertilizer interaction modifies N-sorption, enzyme activities and microbial functional abundance regulating nitrogen retention in rhizosphere soil. Sci Total Environ, 2020, 739. 140065

[43] Jenkinson DS, Powlson DS. The effects of biocidal treatments on metabolism in soil—V: a method for measuring soil biomass. Soil Biol Biochem, 1976, 8: 209-213.

[44] Jonasson S, Michelsen A, Schmidt IK, Nielsen EV, Callaghan TV. Microbial biomass C, N and P in two arctic soils and responses to addition of NPK fertilizer and sugar: implications for plant nutrient uptake. Oecologia, 1996, 106: 507-515.

[45] Kalu S, Simojoki A, Karhu K, Tammeorg P. Long-term effects of softwood biochar on soil physical properties, greenhouse gas emissions and crop nutrient uptake in two contrasting boreal soils. Agric Ecosyst Environ, 2021, 316. 107454

[46] Kang SZ, Zhang JH. Controlled alternate partial root-zone irrigation: its physiological consequences and impact on water use efficiency. J Exp Bot, 2004, 55: 2437-2446.

[47] Kuzyakov Y, Xu X. Competition between roots and microorganisms for nitrogen: mechanisms and ecological relevance. New Phytol, 2013, 198: 656-669.

[48] Larimer AL, Bever JD, Clay K. The interactive effects of plant microbial symbionts: a review and meta-analysis. Symbiosis, 2010, 51: 139-148.

[49] Lehmann J, Gaunt J, Rondon M. Bio-char sequestration in terrestrial ecosystems—a review. Mitig Adapt Strateg Glob Change, 2006, 11: 403-427.

[50] Lehmann J, Rillig MC, Thies J, Masiello CA, Hockaday WC, Crowley D. Biochar effects on soil biota—a review. Soil Biol Biochem, 2011, 43: 1812-1836.

[51] Li FL, Bao WK. New insights into leaf and fine-root trait relationships: implications of resource acquisition among 23 xerophytic woody species. Ecol Evol, 2015, 5: 5344-5351.

[52] Li W, Hou Y, Long M, Wen X, Han J, Liao Y. Long-term effects of biochar application on rhizobacteria community and winter wheat growth on the Loess Plateau in China. Geoderma, 2023, 429. 116250

[53] Liao N, Li Q, Zhang W, Zhou G, Ma L, Min W, Ye J, Hou Z. Effects of biochar on soil microbial community composition and activity in drip-irrigated desert soil. Eur J Soil Biol, 2016, 72: 27-34.

[54] Liu F, Shahnazari A, Andersen MN, Jacobsen SE, Jensen CR. Effects of deficit irrigation (DI) and partial root drying (PRD) on gas exchange, biomass partitioning, and water use efficiency in potato. Sci Hortic, 2006, 109: 113-117.

[55] Liu F, Shahnazari A, Andersen MN, Jacobsen SE, Jensen CR. Physiological responses of potato (Solanum tuberosum L.) to partial root-zone drying: ABA signalling, leaf gas exchange, and water use efficiency. J Exp Bot, 2006, 57: 3727-3735.

[56] Liu X, Fan Y, Long J, Wei R, Kjelgren R, Gong C, Zhao J. Effects of soil water and nitrogen availability on photosynthesis and water use efficiency of Robinia pseudoacacia seedlings. J Environ Sci, 2013, 25: 585-595.

[57] Liu X, Ren J, Zhang Q, Liu C. Long-term effects of biochar addition and straw return on N₂O fluxes and the related functional gene abundances under wheat-maize rotation system in the North China Plain. Appl Soil Ecol, 2019, 135: 44-55.

[58] Liu X, Wei Z, Ma Y, Liu J, Liu F. Effects of biochar amendment and reduced irrigation on growth, physiology, water-use efficiency and nutrients uptake of tobacco (Nicotiana tabacum L.) on two different soil types. Sci Total Environ, 2021, 770: 144738.

[59] Liu X, Wei Z, Manevski K, Liu J, Ma Y, Andersen MN, Liu F. Partial root-zone drying irrigation increases water-use efficiency of tobacco plants amended with biochar. Ind Crops Prod, 2021, 166. 113486

[60] Liu X, Ma Y, Manevski K, Andersen MN, Li Y, Wei Z, Liu F. Biochar and alternate wetting-drying cycles improving rhizosphere soil nutrients availability and tobacco growth by altering root growth strategy in Ferralsol and Anthrosol. Sci Total Environ, 2022, 806. 150517

[61] Loladze I. Rising atmospheric CO₂ and human nutrition: Toward globally imbalanced plant stoichiometry?. Trends Ecol Evol, 2002, 17: 457-461.

[62] Lu T, Wang Y, Zhu H, Wei X, Shao M. Drying-wetting cycles consistently increase net nitrogen mineralization in 25 agricultural soils across intensity and number of drying-wetting cycles. Sci Total Environ, 2020, 710. 135574

[63] Luo X, Chen L, Zheng H, Chang J, Wang H, Wang Z, Xing B. Biochar addition reduced net N mineralization of a coastal wetland soil in the yellow river delta, china. Geoderma, 2016, 282: 120-128.

[64] Ma R, Guan S, Dou S, Wu D, Xie S, Ndzelu BS. Different rates of biochar application change ¹⁵N retention in soil and ¹⁵N utilization by maize. Soil Use Manage, 2020, 36: 773-782.

[65] Markesteijn L, Poorter L. Seedling root morphology and biomass allocation of 62 tropical tree species in relation to drought- and shade-tolerance. J Ecol, 2009, 97: 311-325.

[66] Mašek O, Buss W, Roy-Poirier A, Lowe W, Peters C, Brownsort P, Mignard D, Pritchard C, Sohi S. Consistency of biochar properties over time and production scales: a characterisation of standard materials. J Anal Appl Pyrolysis, 2018, 132: 200-210.

[67] Michelsen A, Graglia E, Schmidt IK, Jonasson S, Sleep D, Quarmby C. Differential responses of grass and a dwarf shrub to long-term changes in soil microbial biomass C, N and P following factorial addition of NPK fertilizer, fungicide and labile carbon to a heath. New Phytol, 1999, 143: 523-538.

[68] Muller B, Pantin F, Génard M, Turc O, Freixes S, Piques M, Gibon Y. Water deficits uncouple growth from photosynthesis, increase C content, and modify the relationships between C and growth in sink organs. J Exp Bot, 2011, 62: 1715-1729.

[69] Nartey OD, Zhao B. Biochar preparation, characterization, and adsorptive capacity and its effect on bioavailability of contaminants: an overview. Adv Mater Sci Eng, 2014, 2014. 715398

[70] Nguyen C. Rhizodeposition of organic c by plants: mechanisms and controls. Agronomie, 2003, 23: 375-396.

[71] Oladele S, Adeyemo A, Awodun M. Influence of rice husk biochar and inorganic fertilizer on soil nutrients availability and rain-fed rice yield in two contrasting soils. Geoderma, 2019, 336: 1-11.

[72] Pandit NR, Mulder J, Hale SE, Martinsen V, Schmidt HP, Cornelissen G. Biochar improves maize growth by alleviation of nutrient stress in a moderately acidic low-input Nepalese soil. Sci Total Environ, 2018, 625: 1380-1389.

[73] Passioura J. Increasing crop productivity when water is scarce—from breeding to field management. Agric Water Manage, 2006, 80: 176-196.

[74] Pausch J, Kuzyakov Y. Carbon input by roots into the soil: quantification of rhizodeposition from root to ecosystem scale. Glob Change Biol, 2018, 24: 1-12.

[75] Peng X, Ye L, Wang C, Zhou H, Sun B. Temperature- and duration-dependent rice straw-derived biochar: characteristics and its effects on soil properties of an Ultisol in southern China. Soil Tillage Res, 2011, 112: 159-166.

[76] Peng Y, Prentice IC, Bloomfield KJ, Campioli M, Guo Z, Sun Y, Tian D, Wang X, Vicca S, Stocker BD. Global terrestrial nitrogen uptake and nitrogen use efficiency. Nat Plants, 2022, 8: 933-940.

[77] Qi D, Hu T. Effects of nitrogen application rates and irrigation regimes on root growth and nitrogen-use efficiency of maize under alternate partial root-zone irrigation. J Soil Sci Plant Nutr, 2022, 22: 2793-2804.

[78] Qin W. Recent developments in using advanced sequencing technologies for the genomic studies of lignin and cellulose degrading microorganisms. Int J Biol Sci, 2016, 12: 156-171.

[79] Rahman MM, Uddin S, Jahangir MMR, Solaiman ZM, Alamri S, Siddiqui MH, Islam MR. Integrated nutrient management enhances productivity and nitrogen use efficiency of crops in acidic and charland soils. Plants, 2021, 10: 2547.

[80] Rewald B, Godbold DL, Falik O, Rachmilevitch S. Root and rhizosphere processes—high time to dig deeper. Front Plant Sci, 2014, 5. 486

[81] Sanchez GPLS path modeling with R, 2013BerkeleyTrowchez Editions383

[82] Schardl CL, Leuchtmann A, Spiering MJ. Symbioses of grasses with seedborne fungal endophytes. Annu Rev Plant Biol, 2004, 55: 315-340.

[83] Shen J, Li C, Mi G, Li L, Yuan L, Jiang R, Zhang F. Maximizing root/rhizosphere efficiency to improve crop productivity and nutrient use efficiency in intensive agriculture of China. J Exp Bot, 2013, 64: 1181-1192.

[84] Silva JLA, Souza AF, Caliman A, Voigt EL, Lichston JE. Weak whole-plant trait coordination in a seasonally dry South American stressful environment. Ecol Evol, 2018, 8: 4-12.

[85] Sinclair TR, Muchow RC. Radiation Use Efficiency. Adv Agron, 1999, 65: 215-265.

[86] Song X, Razavi BS, Ludwig B, Zamanian K, Zang H, Kuzyakov Y, Dippold MA, Gunina A. Combined biochar and nitrogen application stimulates enzyme activity and root plasticity. Sci Total Environ, 2020, 735. 139393

[87] Stefanowicz AM, Stanek M, Nobis M, Zubek S. Species-specific effects of plant invasions on activity, biomass, and composition of soil microbial communities. Biol Fertil Soils, 2016, 52: 841-852.

[88] Tammeorg P, Brandstaka T, Simojoki A, Helenius J. Nitrogen mineralisation dynamics of meat bone meal and cattle manure as affected by the application of softwood chip biochar in soil. Earth Environ Sci Trans R Soc Edinb, 2012, 103: 19-30.

[89] Tomczyk A, Sokołowska Z, Boguta P. Biochar physicochemical properties: pyrolysis temperature and feedstock kind effects. Rev Environ Sci Biotechnol, 2020, 19: 191-215.

[90] Turan V. Confident performance of chitosan and pistachio shell biochar on reducing Ni bioavailability in soil and plant plus improved the soil enzymatic activities, antioxidant defense system and nutritional quality of lettuce. Ecotoxicol Environ Saf, 2019, 183. 109594

[91] Walkley A, Black IA. An examination of the Degtjareff method for determining soil organic matter, and a proposed modification of the chromic acid titration method. Soil Sci, 1934, 37: 29-38.

[92] Wan H, Chen Y, Cui B, Liu X, Hou J, Wei Z, Liu J, Liu F. Biochar amendment increases C and N retention in the soil–plant systems: its implications in enhancing plant growth and water-use efficiency under reduced irrigation regimes of maize (Zea mays L.). J Soil Sci Plant Nutr, 2023, 23: 1-13.

[93] Wan H, Liu X, Shi Q, Chen Y, Jiang M, Zhang J, Cui B, Hou J, Wei Z, Hossain MA, Liu F. Biochar amendment alters root morphology of maize plant: its implications in enhancing nutrient uptake and shoot growth under reduced irrigation regimes. Front Plant Sci, 2023, 14. 1122742

[94] Wan H, Hou J, Wei Z, Liu F. Contrasting maize responses to soil phosphorus and potassium availability driven by biochar under reduced irrigation. Plant Soil, 2024, 492: 1-24.

[95] Wan H, Wei Z, Liu C, Yang X, Wang Y, Liu F. Biochar amendment modulates xylem ionic constituents and ABA signaling: its implications in enhancing water-use efficiency of maize (Zea mays L.) under reduced irrigation regimes. J Integr Agric, 2025, 24: 132-146.

[96] Wang Y, Liu F, de Neergaard A, Jensen LS, Luxhøi J, Jensen CR. Alternate partial root-zone irrigation induced dry/wet cycles of soils stimulate N mineralization and improve N nutrition in tomatoes. Plant Soil, 2010, 337: 167-177.

[97] Wang Y, Hu Y, Zhao X, Wang S, Xing G. Comparisons of biochar properties from wood material and crop residues at different temperatures and residence times. Energy Fuels, 2013, 27: 5890-5899.

[98] Wang C, Liu J, Shen J, Chen D, Li Y, Jiang B, Wu J. Effects of biochar amendment on net greenhouse gas emissions and soil fertility in a double rice cropping system: a 4-year field experiment. Agric Ecosyst Environ, 2018, 262: 83-96.

[99] Wang L, Ok YS, Tsang DCW, Alessi DS, Rinklebe J, Wang H, Mašek O, Hou R, O’Connor D, Hou D. New trends in biochar pyrolysis and modification strategies: feedstock, pyrolysis conditions, sustainability concerns and implications for soil amendment. Soil Use Manag, 2020, 36: 358-386.

[100] Wang Z, Wang Z, Luo Y, Zhan YN, Meng YL, Zhou ZG. Biochar increases ¹⁵N fertilizer retention and indigenous soil N uptake in a cotton–barley rotation system. Geoderma, 2020, 357. 113944

[101] Wang XC, Liu R, Luo JN, Zhu PF, Wang YS, Pan XC, Shu LZ. Effects of water and NPK fertigation on watermelon yield, quality, irrigation-water, and nutrient use efficiency under alternate partial root-zone drip irrigation. Agric Water Manag, 2022, 271. 107785

[102] Wei Z, Fang L, Li X, Liu J, Liu F. Endogenous ABA level modulates the effects of CO₂ elevation and soil water deficit on growth, water and nitrogen use efficiencies in barley and tomato plants. Agric Water Manag, 2021, 249. 106808

[103] Wetzels M, Odekerken-Schröder G, Van Oppen C. Using PLS path modeling for assessing hierarchical construct models: guidelines and empirical illustration. MIS Q, 2009, 33: 177-195.

[104] Xiang Y, Deng Q, Duan H, Guo Y. Effects of biochar application on root traits: a meta-analysis. GCB Bioenergy, 2017, 9: 1563-1572.

[105] Xu N, Wang R, Liu J, Lu P, Guo W. Hierarchy of plasticity traits in responses of Quercus aliena to light conditions and water availability. Dendrobiology, 2015, 74: 49-59.

[106] Yan B, Ji Z, Fan B, Wang X, He G, Shi L, Liu G. Plants adapted to nutrient limitation allocate less biomass into stems in an arid-hot grassland. New Phytol, 2016, 211: 1232-1240.

[107] Yang J, Zhang J. Crop management techniques to enhance harvest index in rice. J Exp Bot, 2010, 61: 3177-3189.

[108] Yang X, Bornø ML, Wei Z, Liu F. Combined effect of partial root drying and elevated atmospheric CO₂ on the physiology and fruit quality of two genotypes of tomato plants with contrasting endogenous ABA levels. Agric Water Manag, 2021, 254. 106970

[109] Zegada-Lizarazu W, Monti A. Deep root growth, ABA adjustments and root water uptake response to soil water deficit in giant reed. Ann Bot, 2019, 124: 605-615.

[110] Zhang Q, Song Y, Wu Z, Yan X, Gunina A, Kuzyakov Y, Xiong Z. Effects of six-year biochar amendment on soil aggregation, crop growth, and nitrogen and phosphorus use efficiencies in a rice–wheat rotation. J Clean Prod, 2020, 242. 118435

[111] Zhao X, Bi G, Harkess RL, Varco JJ, Blythe EK. Spring nitrogen uptake, use efficiency, and partitioning for growth in Iris germanica ‘Immortality’. HortScience, 2016, 51(5): 563-566.

[112] Zhu L, Luan L, Chen Y, Wang X, Zhou S, Zou W, Han X, Duan Y, Zhu B, Li Y. Community assembly of organisms regulates soil microbial functional potential through dual mechanisms. Glob Change Biol, 2024, 30. e17160

Funding

Inner Mongolia Agricultural University Basic Research Project(BR22-13-04)

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