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Journal of Integrative Agriculture
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Biochar-driven redistribution of soil nitrogen pools enhances nitrogen uptake in wheat under elevated CO2

Qi Yang1, 2, Lujie Li1, Wanhong Jin1, Ping Li1, Zhiqiang Gao1, Dongsheng Zhang1, Xinrui Shi1, Xingyu Hao1, 2#, Yuzheng Zong1, 2#

1 College of Agriculture, Shanxi Agricultural University, Taigu 030801, China

2 Shanxi Provincial Key Laboratory of Crop Ecology and Efficient Water & Nutrient Use, Taigu 030801, China

 Highlights 

· Biochar increases wheat N uptake under e[CO2] by improving both fertilizer-N and native soil-N use.

· Biochar shifts soil N to labile pools and retains it within micro-aggregates.

· Biochar mitigates elevated CO2-induced NO3--N accumulation.

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摘要  

全球气候变化背景下,大气CO2浓度升高对农田生态系统氮循环带来严峻挑战。本研究通过为期三年(2019-2022年)的小麦盆栽试验揭示了生物炭在CO2浓度升高条件下调控土壤氮转化与植物氮获取的机制。结合15N同位素示踪宏基因组测序技术,我们分析了两种CO2浓度(正常CO2浓度: 400 μmol mol⁻¹,高CO2浓度: 600 μmol mol⁻¹)与添加2%(质量分数)生物炭之间的交互效应。结果表明,在CO2浓度升高条件下,施用生物炭使肥料源氮重组有机氮中的储存降低39.4%,同时使土壤源氮在轻组有机氮中的含量提高34.0%。此外,生物炭促进了微团聚体(<0.25 mm)和颗粒有机氮的形成,增幅分别为37.3%和13.2%。宏基因组分析表明,在CO2浓度升高条件下,生物炭抑制了在正常CO2条件下上调的氮循环关键基因(包括同化、硝化和硝酸盐还原)的相对丰度。上述物理化学过程与微生物调控共同作用,导致土壤NO3⁻-N积累量降低52.6%并显著提高了植株地上部氮吸收。结构方程模型进一步表明,生物炭能够抵消CO2浓度升高对微团聚体稳定性和氮循环基因丰度的负面影响。在这种协同作用下,生物炭在CO2浓度升高条件下使肥料氮土壤氮的吸收分别30.8%111.4%使籽粒氮积累量增加55.4%研究表明,生物炭作为一种有效的土壤改良剂,通过将氮从惰性活性库再分配、增强氮生物有效性,缓解CO2浓度升引起的氮限制,从而为未来农业生态系统作物生产力的维持提供支持



Abstract  

Under accelerating climate change, elevated atmospheric CO2 (e[CO2]) presents a substantial challenge to nitrogen (N) cycling in agricultural systems. This study elucidated the mechanistic role of biochar in regulating soil N transformations and plant N acquisition under e[CO2] conditions through a three-year pot experiment (2019-2022) with wheat. Using 15N isotope tracing combined with metagenomic sequencing, we examined the interactions between two CO2 concentrations (a[CO2] 400 μmol mol1 vs. e[CO2] 600 μmol mol−1) and 2% (w/w) biochar amendment. Our results demonstrated that under e[CO2], biochar application reduced the incorporation of fertilizer-derived N into the recalcitrant heavy fraction organic N (HFON) by 39.4%, while enhancing the content of native soil-derived N in the light fraction N (LFON) by 34.0%. Concurrently, biochar promoted the formation of micro-aggregates (<0.25 mm) and particulate organic N (PON) by 37.3 and 13.2%, respectively. Metagenomic analysis revealed that biochar under e[CO2] suppressed the relative abundance of key N-cycling genes (involved in assimilation, nitrification, and nitrate reduction) that were upregulated under a[CO2] condition. These physicochemical processes, coupled with microbial modulation, resulted in a 52.6% reduction in soil NO3--N accumulation and a significant increase in aboveground N uptake. Structural equation modeling indicated that biochar counteracted the adverse effects of e[CO2] on micro-aggregate stability and N-cycling gene abundance. Synergistically, biochar enhanced the uptake of fertilizer-N and native soil-N by 30.8% and 111.4%, respectively, under e[CO2], leading to a 55.4% increase in grain N accumulation. Our findings demonstrate that biochar is an effective amendment for mitigating e[CO2]-induced N limitation by redistributing N from recalcitrant to labile pools, enhancing N bioavailability, and ultimately supporting crop productivity in future CO2-enriched agroecosystems.

Keywords:  biochar       nitrogen transformation              elevated CO2 concentration              gene abundance  
Online: 12 March 2026  
Fund: 

The research was supported by the Shanxi Scholarship Council of China (2024-070), the Youth Talents Support Program of Shanxi Agricultural University, China (BJRC201602), the National Natural Science Foundation of China (32572456), the Graduate Education Reform and Quality Improvement Program of College of Agriculture, Shanxi Agricultural University, China (2023YCX16).

About author:  Qi Yang, E-mail: yangxiaoqi517@163.com; #Correspondence Xingyu Hao, E-mail: haoxingyu1976@126.com; Yuzheng Zong, E-mail: zongyuzheng@163.com

Cite this article: 

Qi Yang, Lujie Li, Wanhong Jin, Ping Li, Zhiqiang Gao, Dongsheng Zhang, Xinrui Shi, Xingyu Hao, Yuzheng Zong. 2026. Biochar-driven redistribution of soil nitrogen pools enhances nitrogen uptake in wheat under elevated CO2. Journal of Integrative Agriculture, Doi:10.1016/j.jia.2026.03.033

Ahmad S, Sehrish A K, Umair M, Mirino M W, Ali S, Guo H. 2024. Effect of biochar amendment on bacterial community and their role in nutrient acquisition in spinach (Spinacia oleracea L.) grown under elevated CO2. Chemosphere, 364, 143098.

Baggs E M, Richter M, Cadisch G, Hartwig U A. 2003. Denitrification in grass swards is increased under elevated atmospheric CO2. Soil Biology and Biochemistry, 35, 729-732.

Ball P N, Mackenzie M D, Deluca T H, Holben W E. 2010. Wildfire and charcoal enhance nitrification and ammonium-oxidizing bacterial abundance in dry montane forest soils. Journal of Environmental Quality, 39, 1243-1253.

Barnard R, Leadley P W, Lensi R, Barthes L. 2005. Plant, soil microbial and soil inorganic nitrogen responses to elevated CO2: A study in microcosms of Holcus lanatus. Acta Oecologica, 27, 171-178.

Bloom A J, Burger M, Asensio J S R, Cousins A B. 2010. Carbon dioxide enrichment inhibits nitrate assimilation in wheat and Arabidopsis. Science, 328, 899-903.

Choudhury T M A, Khanif Y M. 2001. Evaluation of effects of nitrogen and magnesium fertilization on rice yield and fertilizer nitrogen efficiency using 15N tracer technique. Journal of Plant Nutrition, 24, 855-871.

Drigo B, Pijl A S, Duyts H, Kielak A M, Gamper H A, Houtekamer M J, Boschker H T S, Bodelier P L E, Whiteley A S, Veen J A V, Kowalchuk G A. 2010. Shifting carbon flow from roots into associated microbial communities in response to elevated atmospheric CO2. Proceedings of the National Academy of Sciences of the United States of America, 107, 10938-10942.

Fu X, Shao M, Wei X, Horton R. 2010. Soil organic carbon and total nitrogen as affected by vegetation types in northern Loess Plateau of China. Geoderma, 155, 31-35.

Gao K, Mao Z, Meng E, Li J, Liu X, Zhang Y, Zhang L, Wang G, Liu Y. 2022. Effects of elevated CO2 and warming on the root-associated microbiota in an agricultural ecosystem. Environmental Microbiology, 24, 6252-6266.

Gou Z, Zheng H, He Z, Su Y, Chen S, Chen H, Chen G, Ma N L, Sun Y. 2023. The combined action of biochar and nitrogen-fixing bacteria on microbial and enzymatic activities of soil N cycling. Environmental Pollution, 317, 120790.

Hao X Y, Li P, Li H Y, Zong Y Z, Zhang B, Zhao J Z, Han Y H. 2017. Elevated CO2 increased photosynthesis and yield without decreasing stomatal conductance in broomcorn millet. Photosynthetica, 55, 176-183.

Hovenden M J, Newton P C D, Carran R A, Theobald P, Wills K E, Vander Schoor J K, Williams A L, Osanai Y. 2008. Warming prevents the elevated CO2-induced reduction in available soil nitrogen in a temperate, perennial grassland. Global Change Biology, 14, 1018-1024.

IPCC (Intergovernmental Panel on Climate Change). 2021. Summary for policymakers. In: Masson-Delmotte V, Zhai P, Pirani A, Connors S L, P´ean C, Berger S, Caud N, Chen Y, Goldfarb L, Gomis M I, Huang M, Leitzell K, Lonnoy E, Matthews J B R , Maycock T K, Waterfield T, Yelekçi O, Yu R, Zhou B, eds., Climate Change 2021: The Physical Science Basis. Contribution of Working Group I to the Sixth Assessment Report of the Intergovernmental Panel on Climate Change. Cambridge University Press, UK.

Janzen H H, Campbell C A, Brandt S A, Lafond G P, Townley-Smith L. 1992. Light-fraction organic matter in soils from long-term crop rotations. Soil Science Society of America Journal, 56, 1799-1806.

Jin J, Krohn C, Franks A E, Wang X, Wood J L, Petrovski S, Mccaskill M, Batinovic S, Xie Z, Tang C. 2022. Elevated atmospheric CO2 alters the microbial community composition and metabolic potential to mineralize organic phosphorus in the rhizosphere of wheat. Microbiome, 10, 1-17.

Kim H, Lieffering M, Kobayashi K, Okada M, Miura S. 2003. Seasonal changes in the effects of elevated CO2 on rice at three levels of nitrogen supply: A free air CO2 enrichment (FACE) experiment. Global Change Biology, 9, 826-837.

Laird D A, Fleming P, Baiqun W, Horton R, Karlen D L. 2010. Biochar impact on nutrient leaching from a Midwestern agricultural soil. Geoderma, 158, 436-442.

Lehmann J, Da Silva J P, Steiner C, Nehls T, Zech W, Glaser B. 2003. Nutrient availability and leaching in an archaeological Anthrosol and a Ferralsol of the Central Amazon basin: Fertilizer, manure and charcoal amendments. Plant and Soil, 249, 343-357.

Li S, Wang Z, Stewart B A. 2013. Responses of crop plants to ammonium and nitrate N. In: Advances in Agronomy. Elsevier, USA.

Li Z, Tian D, Wang B, Wang J, Wang S, Chen H Y H, Xu X, Wang C, He N, Niu S. 2019. Microbes drive global soil nitrogen mineralization and availability. Global Change Biology, 25, 1078-1088.

Liu Q, Zhang Y, Liu B, Amonette J E, Lin Z, Liu G, Ambus P, Xie Z. 2018. How does biochar influence soil N cycle? A meta-analysis. Plant and Soil, 426, 211-225.

Luo Y, Su B O, Currie W S, Dukes J S, Finzi A, Hartwig U, Hungate B, Mc Murtrie R E, Oren R, Parton W J, Pataki D E, Shaw M R, Zak D R, Field C B. 2004. Progressive nitrogen limitation of ecosystem responses to rising atmospheric carbon dioxide. Bioscience, 54, 731-739.

Mark F, Thomas K K, Lynne M M, Todd M M, Daniel V M, Phillip A H, Bhupinder P S, Karen B, Evelyn S K, Jeff A B. 2013. Microbial utilization of biochar-derived carbon. Science of the Total Environment, 465, 288-297.

Müller C, Rütting T, Abbasi M K, Laughlin R J, Kammann C, Clough T J, Sherlock R R, Kattge J, Jäger H, Watson C J, Stevens R J. 2009. Effect of elevated CO2 on soil N dynamics in a temperate grassland soil. Soil Biology & Biochemistry, 41, 1996-2001.

Mulvaney R L. 1996. Nitrogen-Inorganic Forms. Soil Science Society of America, Madison.

Nelissen V, Ruysschaert G, Manka’Abusi D, D’Hose T, De Beuf K, Al-Barri B, Cornelis W, Boeckx P. 2015. Impact of a woody biochar on properties of a sandy loam soil and spring barley during a two-year field experiment. European Journal of Agronomy, 62, 65-78.

Nelson D M, Cann I K, Mackie R I. 2010. Response of archaeal communities in the rhizosphere of maize and soybean to elevated atmospheric CO2 concentrations. PLoS ONE, 5, e15897.

Norby R J, Warren J M, Iversen C M, Medlyn B E, Mcmurtrie R E, Schlesinger W H. 2010. CO2 enhancement of forest productivity constrained by limited nitrogen availability. Proceedings of the National Academy of Sciences of the United States of America, 107, 19368-19373.

Norgbey E, Murava R T, Rajasekar A, Huang Q, Zhou J, Robinson S. 2022. Effects of anthropogenic nitrogen additions and elevated CO2 on microbial community, carbon and nitrogen content in a replicated wetland. Environmental Monitoring and Assessment, 194, 575.

Obia A, Mulder J, Martinsen V, Cornelissen G, Børresen T. 2016. In situ effects of biochar on aggregation, water retention and porosity in light-textured tropical soils. Soil & Tillage Research, 155, 35-44.

Ontl T A, Cambardella C A, Schulte L A, Kolka R K. 2015. Factors influencing soil aggregation and particulate organic matter responses to bioenergy crops across a topographic gradient. Geoderma, 255-256, 1-11.

Phillips R P, Finzi A C, Bernhardt E S. 2011. Enhanced root exudation induces microbial feedbacks to N cycling in a pine forest under long-term CO2 fumigation. Soil Ecology Letters, 14, 187-194.

Ray D K, Mueller N D, West P C, Foley J A. 2013. Yield trends are insufficient to double global crop production by 2050. PLoS ONE, 8, e66428.

Rees F, Germain C, Sterckeman T, Morel J. 2015. Plant growth and metal uptake by a non-hyperaccumulating species (Lolium perenne) and a Cd-Zn hyperaccumulator (Noccaea caerulescens) in contaminated soils amended with biochar. Plant and Soil, 395, 57-73.

Ren J, Huang H, Zhang Z, Xu X, Zhao L, Qiu H, Cao X. 2023. Enhanced microbial reduction of Cr (VI) in soil with biochar acting as an electron shuttle: Crucial role of redox-active moieties. Chemosphere, 328, 138601.

Sharifi M, Zebarth B J, Burton D L, Grant C A, Cooper J M. 2007. Evaluation of some indices of potentially mineralizable nitrogen in soil. Soil Science Society of America Journal, 71, 1233-1239.

Sheng M, Ai X, Huang B, Zhu M, Liu Z, Ai Y. 2023. Effects of biochar additions on the mechanical stability of soil aggregates and their role in the dynamic renewal of aggregates in slope ecological restoration. Science of the Total Environment, 898, 165478.

Shrestha G, Traina S J, Swanston C W. 2010. Black carbon’s properties and role in the environment: A comprehensive review. Sustainability, 2, 294-320.

Six J, Bossuyt H, Degryze S, Denef K. 2004. A history of research on the link between (micro)aggregates, soil biota, and soil organic matter dynamics. Soil & Tillage Research, 79, 7-31.

Sui Q, Di F, Zhong H, Chen M, Wei Y. 2024. Molecular insight into the allocation of organic carbon to heterotrophic bacteria: Carbon metabolism and the involvement in nitrogen and phosphorus removal. Science of the Total Environment, 933, 173302.

Tian L, Wang Y, Jin D, Zhou Y, Mukhamed B, Liu D, Feng B. 2025. The application of biochar and organic fertilizer substitution regulates the diversities of habitat specialist bacterial communities within soil aggregates in proso millet farmland. Biochar, 7, 1-18.

Uzoma K C, Inoue M, Andry H, Fujimaki H, Zahoor A, Nishihara E. 2011. Effect of cow manure biochar on maize productivity under sandy soil condition. Soil Use and Management, 27, 205-212.

Wang L, Li M, Liu X. 2025. Role of urea in the retention of DON in soil by clay minerals: Analysis based upon molecular weight. Journal of Environmental Sciences, 150, 362-372.

Xu M, He Z, Deng Y, Wu L, van Nostrand J D, Hobbie S E, Reich P B, Zhou J. 2013. Elevated CO2 influences microbial carbon and nitrogen cycling. BMC Microbiology, 13, 124.

Yan D, Wang D, Yang L. 2007. Long-term effect of chemical fertilizer, straw, and manure on labile organic matter fractions in a paddy soil. Biology and Fertility of Soils, 44, 93-101.

Yu C, Chen H, Tang B, Liu H, Liu X, Zhang H, Wang G. 2026. Biochar integrate dicyandiamide modified soil aggregates and optimized nitrogen supplying to boosting the soybean-wheat yield in saline-alkali soil. Soil & Tillage Research, 257, 106922.

Yu Z, Li Y, Wang G, Liu J, Liu J, Liu X, Herbert S J, Jin J. 2016. Effectiveness of elevated CO2 mediating bacterial communities in the soybean rhizosphere depends on genotypes. Agriculture, Ecosystems & Environment, 231, 229-232.

Zak D R, Pregitzer K S, Curtis P S, Teeri J A, Fogel R, Randlett D L. 1993. Elevated atmospheric CO2 and feedback between carbon and nitrogen cycles. Plant and Soil, 151, 105-117.

Zhu K, Ma S, Chen N, Dai Y, Wang T, Guo X, Jia H. 2024. Robust reactive oxygen species production in interfacial reaction between organic acids and biochar: The combined effect of electron acceptance and electron conduction. Journal of Hazardous Materials, 464, 132960.

Van Zwieten L, Kimber S, Morris S, Chan K Y, Downie A, Rust J, Joseph S, Cowie A. 2010. Effects of biochar from slow pyrolysis of papermill waste on agronomic performance and soil fertility. Plant and Soil, 327, 235-246.

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