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集约化互作体系植物根系高效获取土壤养分的策略与机制

花匠小妙招
2026-08-16 08:06

摘要:

目的植物根系的形态与生理变化是植物从土壤中高效获取养分资源的重要机制，由相同物种或不同物种组成的互作体系中植物根系对养分的吸收利用受相邻植物竞争的强烈影响，阐明互作体系不同竞争条件下植物根系获取养分的策略并揭示其作用机制，这是基于根系觅食行为探讨养分高效利用的根际调控途径与技术措施的重要理论基础。

主要进展根系属性的互补性有利于降低根系间对养分的竞争。根系构型的互补性，例如深根系与浅根系植物互作，促进个体植株对土壤剖面不同深度养分的吸收利用；由根系可塑性介导的水平方向上根系空间分布的互补性，提高了植物根系对同一土层不同空间位点土壤养分的挖掘；个体植株根系形态属性与相邻植物根际生理过程的互补性促进根系对不同形态养分的利用。互作体系根系获取养分的策略具有高度互补性，这有助于提高整个作物系统的养分利用效率，进而提高生产力。根系空间生态位的分离 (包括垂直与水平方向) 以及根际生物化学特征生态位的分离，是驱动互作体系根系高效获取养分资源的主要机制。合理的根层调控可以提高植物根系挖掘土壤养分的能力；优化互作体系物种的搭配能充分发挥根的互作效能，提高养分利用的生物潜力。

问题与展望今后应进一步针对集约化高投入作物体系，通过管理根层养分供应和物种间的互作效应，强化根际养分信号的调控作用，调节根系形态与生理特性，降低种间竞争，增强种间互利，以最大化根系和根际的生物学潜力，提高养分利用效率和作物产量，为实现以节肥增效为核心的可持续集约化作物生产提供重要的调控策略与途径。

Abstract:

ObjectivesRoot morphological and physiological plasticity is critical for plant nutrient acquisition from soil. The nutrient uptake ability of individual plant in mixture cropping system (intra-specific or inter-specific cropping systems) is strongly affected by neighboring plant presence. This paper focused on investigating the foraging behaviors and strategies of roots in mixture systems, understanding the underlying mechanism and suggesting major pathways and approaches of rhizosphere management for improving nutrient acquisition.

Major advancesThe complementarity of root traits of the neighboring plants could decrease root competition for nutrients. The difference in root architecture, i.e. deep-rooted plant intercropping with shallow-rooted plant, is beneficial to utilize soil nutrients in different soil layers. The root-placement pattern of neighboring plants driven by root plasticity improves nutrient use efficiency in different soil zones in the same soil layer. Root morphological plasticity may also exhibit a complementary effect with neighboring root physiological traits, resulting in the utilization of different fraction of nutrients in soil. Niche differentiation has been considered an important mechanism for species coexistence, including spatial niche differentiation in vertical and horizontal root distribution, and the biochemistry niche differentiation. Applying or controlling fertilizer nutrient supply at a proper level in root zone could enhance root ability for excavating soil nutrients; matching the intercropped species through optimizing plant combinations reasonably could improve nutrient use efficiency through maximizing the biological effect of root-root interactions.

Suggestions and expectationsAs for intensive farming systems, optimizing the combinations of intercropped species and the root-zone nutrient management will strengthen nutrient signaling-based rhizosphere processes and manipulation, and regulate root morphological and physiological traits, which will affect the root foraging behaviors reduce root competition and enhance root facilitation between plants and their neighbors. These strategies of root/rhizosphere interactions are critical to develop the approaches of improving nutrient use efficiency and productivity through maximizing root/rhizosphere efficiency in intensive cropping systems, and achieving " saving fertilizer and enhancing efficiency” sustainable intensification of grain production with less input and producing more.

图 1 基于根系高效获取土壤养分策略的物种共存机理概念模式图

Figure 1. Conceptual model for the mechanisms driving plant co-existence based on root foraging strategies for efficient nutrient use

[1]	张福锁, 申建波, 冯固. 根际生态学—过程与调控[M]. 北京: 中国农业大学出版社, 2009. Zhang F S, Shen J B, Feng G. Rhizosphere ecology: Processes and management [M]. Beijing: China Agricultural University Press, 2009.
[2]	Schenk H J. Root competition: beyond resource depletion[J]. Journal of Ecology, 2006, 94: 725–739.
[3]	Zhang F S, Li L, Sun J H. Contribution of above- and below-ground interactions to intercropping [A]. Horst W J, Schenk M K, Bürkert A, et al. Plant nutrition-food security and sustainability of agroecosystems [M]. Dordrecht: Kluwer Academic Publishers, 2001. 979–980.
[4]	Zhang F S, Shen J B, Zhang J L, et al. Rhizosphere processes and management for improving nutrient use efficiency and crop productivity: implications for China [A]. Sparks D L. Advances in agronomy [M]. Burlington: Academic Press, 2010. 1–32.
[5]	Lynch J P. Root architecture and nutrient acquisition [A]. BassiriRad H. Nutrient acquisition by plants [M]. Berlin/Heidelberg: Springer Berlin Heidelberg Publisher, 2005. 146–183.
[6]	Lambers H, Shane M W, Cramer M D, et al. Root structure and functioning for efficient acquisition of phosphorus: Matching morphological and physiological traits[J]. Annals of Botany, 2006, 98: 693–713.
[7]	White P J, Hammond J P. Phosphorus nutrition of terrestrial plants [A]. White P J, Hammond J P. The ecophysiology of plant phosphorus interaction [C]. Dordrecht: Springer, 2008. 51–81.
[8]	Rhichardson A E, Lynch J P, Ryan P R, et al. Plant and microbial strategies to improve the phosphorus efficiency of agriculture[J]. Plant and Soil, 2011, 349: 121–156.
[9]	Cahill J F, McNickle G G, Haag J J, et al. Plant integrate information about nutrients and neighbors[J]. Science, 2010, 328: 1657.
[10]	Mommer L, Van Ruijven J, Jansen C, et al. Interactive effects of nutrient heterogeneity and competition: implications for root foraging theory?[J]. Functional Ecology, 2012, 26: 66–73.
[11]	Zhang D S, Zhang C C, Tang X Y et al. Increased soil P availability induced by faba bean root exudation stimulates root growth and P uptake in neighbouring maize[J]. New Phytologist, 2016, 209: 823–831.
[12]	Shea K, Chesson P. Community ecology theory as a framework for biological invasion[J]. Trends in Ecology & Evolution, 2002, 17: 170–176.
[13]	Gause G F. The struggle for existence [M]. Baltimore: Williams and Wilkins, 1934.
[14]	Gross K L, Pregitzer K S, Burton A J. Spatial variation in nitrogen availability in three successional plant communities[J]. Journal of Ecology, 1995, 83: 357–367.
[15]	Marschner H. Mineral nutrition in higher plants [M]. London: Academic Press, 1995.
[16]	Drew M C. Comparison of the effects of localized supply of phosphorus, nitrate, ammonium and potassium on the growth of the seminal root system, and the shoot, in barley[J]. New Phytologist, 1975, 75: 479–490.
[17]	Jackson R B, Caldwell M M. The timing and degree of root proliferation in fertile-soil microsites for three cold-desert perennials[J]. Oecologia, 1989, 81: 149–153.
[18]	Hodge A, Robinson D, Griffiths B S, et al. Why plants bother: root proliferation results in increased nitrogen capture from an organic patch when two grasses compete[J]. Plant, Cell and Environment, 1999, 22: 811–820.
[19]	Farley R A, Fitter A H. The response of seven co-occurring woodland herbaceous perennials to localized nutrient-rich patches[J]. Journal of Ecology, 1999, 87: 849–859.
[20]	李洪波, 薛慕瑶, 林雅茹, 等. 土壤养分空间异质性与根系觅食作用: 从个体到群落[J]. 植物营养与肥料学报, 2013, 19: 995–1004. Li H B, Xue M Y, Lin Y R, et al. Spatial heterogeneity of soil nutrients and root foraging: from individual to community[J]. Journal of Plant Nutrition and Fertilizer, 2013, 19: 995–1004.
[21]	Hodge A. The plastic plant: root responses to heterogeneous supplies of nutrients[J]. New Phytologist, 2004, 162: 9–24.
[22]	Robinson D. The response of plants to non-uniform supplies of nutrients[J]. New Phytologist, 1994, 127: 635–674.
[23]	de Kroon H, Mommer L. Root foraging theory put to the test[J]. Trends in Ecology & Evolution, 2006, 21: 113–116.
[24]	Einsmann J C, Jones R H, Pu M, et al. Nutrient foraging traits in 10 co-occurring plant species of contrasting life forms[J]. Journal of Ecology, 1999, 87: 609–619.
[25]	Fransen B, Kroon H, Berendse F. Root morphological plasticity and nutrient acquisition of perennial grass species from habitats of different nutrient availability[J]. Oecologia, 1998, 115: 351–358.
[26]	Kembel S W, Cahill J F. Plant phenotypic plasticity belowground: a phylogenetic perspective on root foraging trade-offs[J]. American Naturalist, 2005, 166: 216–230.
[27]	Li H B, Ma Q H, Li H G, et al. Root morphological responses to localized nutrient supply differ among crop species with contrasting root traits[J]. Plant and Soil, 2014, 376: 151–163.
[28]	Jackson R B, Manwaring J H, Caldwell M M. Rapid physiological adjustment of roots to localized soil enrichment[J]. Nature, 1990, 344: 58–60.
[29]	Day K J, Hutchings M J, John E A. The effects of spatial pattern of nutrient supply on yields, structure and mortality in plant populations[J]. Journal of Ecology, 2003, 91: 541–553.
[30]	李洪波. 植物响应养分空间异质性分布的动态过程及调控根系获取养分的策略研究[D]. 北京: 中国农业大学博士学位论文, 2014. Li H B. The dynamic process of plant responses to heterogeneous nutrient environments and the strategies of root management for nutrient acquisition[D]. Beijing: PhD Dissertation of China Agricultural University, 2014.
[31]	Ma Q H, Zhang F S, Rengel Z, Shen J B. Localized application of NH4+-N plus P at the seedling and later growth stages enhances nutrient uptake and maize yield by inducing lateral root proliferation[J]. Plant and Soil, 2013, 372: 65–80.
[32]	McNickle G G, Cahill J F. Plant root growth and the marginal value theorem[J]. Proceedings of the National Academy of Sciences, 2009, 106: 4747–4751.
[33]	Cahill J F, McNickle G G. The behavioral ecology of nutrient foraging by plants[J]. Annual Review of Ecology Evolution and Systematics, 2011, 42: 289–311.
[34]	Gersani M, Abramsky Z, Falik O. Density-dependent habitat selection in plants[J]. Evolutionary Ecology, 1998, 12: 223–234.
[35]	Schenk H J, Callaway R M, Mahall B E. Spatial root segregation: are plants territorial?[J]. Advances in Ecological Research, 1999, 28: 145–168.
[36]	de Kroon H. How do roots interact?[J]. Science, 2007, 318: 1562–1563.
[37]	Kulmatiski A, Beard K H, Verweij R J T, et al. A depth controlled tracer technique measures vertical, horizontal and temporal patterns of water use by trees and grasses in a subtropical savanna[J]. New Phytologist, 2010, 188: 199–209.
[38]	Rubio G, Walk T, Ge Z Y, et al. Root gravitropism and below-ground competition among neighbouring plants: a modeling approach[J]. Annals of Botany, 2001, 88: 929–940.
[39]	Berendse F. Competition between plant-populations with different rooting depths. 1. Theoretical considerations[J]. Oecologia, 1979, 43: 19–26.
[40]	Li L, Sun J H, Zhang F S, et al. Root distribution and interactions between intercropped species[J]. Oecologia, 2006, 147: 280–290.
[41]	Espeleta J F, Donovan L A. Fine root demography and morphology in response to soil resources availability among xeric and mesic sand hill tree species[J]. Functional Ecology, 2002, 16: 113–121.
[42]	Comas L H, Bouma T J, Eissentat D M. Linking root traits to maximum potential growth rate among eleven mature temperate tree species[J]. Oecologia, 2002, 132: 34–43.
[43]	Hess L, de Kroon. Effects of rooting volume and nutrient availability as an alternative explanation for root self/non-self discrimination[J]. Journal of Ecology, 2007, 95: 241–251.
[44]	Genney D R, Alexander I J, Hartley S E. Soil organic matter distribution and below-ground competition between Calluna vulgaris and Nardus stricta[J]. Functional Ecology, 2002, 16: 664–670.
[45]	Robinson D, Hodge A, Griffiths B S, et al. Plant root proliferation in nitrogen-rich patches confers competitive advantage[J]. Proceedings of the Royal Society of London, 1999, 266: 431–435.
[46]	de Kroon H, Mommer L, Nishiwaki A. Root competition: Towards a mechanistic understanding[A]. de Kroon H, Visser E J W. Root ecology [M]. Berlin: Springer, 2003. 215–234.
[47]	Hutchings M J, John E A, Wijesinghe D K. Toward understanding the consequences of soil heterogeneity for plant populations and communities[J]. Ecology, 2003, 84: 2322–2334.
[48]	Brisson J, Reynolds J F. The effect of neighbors on root distribution in a creosote bush (Larrea tridentata) population[J]. Ecology, 1994, 75: 1693–1702.
[49]	Brisson J, Reynolds J F. Effects of compensatory growth on population processes: a simulation study[J]. Ecology, 1997, 78: 2378–2384.
[50]	Hinsinger P, Betencourt E, Bernard L, et al. P for two, sharing a scarce resource: soil phosphorus acquisition in the rhizosphere of intercropped species[J]. Plant Physiology, 2011, 156: 1078–1086.
[51]	Cu S T T, Huntson J, Schuller K A. Mixed culture of wheat (Triticum aestivum L.) with white lupin (Lupinusalbus L.) improves the growth and phosphorus nutrition of the wheat[J]. Plant and Soil, 2005, 272: 143–151.
[52]	Li L, Li S M, Sun J H, et al. Diversity enhances agricultural productivity via rhizosphere phosphorus facilitation on phosphorus-deficient soils[J]. Proceedings of the National Academy of Sciences, 2007, 104: 11192–11196.
[53]	Chase J M, Leibold M A. Ecological niches: Linking classical and contemporary approaches [M]. Chicago: University of Chicago Press, 2003.
[54]	Geno L, Geno B. Polyculture production-principles, benefits and risks of multiple cropping land management systems for Australia. A Report for the Rural Industries and Development Corporations [R]. CIRDC Publication No. 01/34, 2001.
[55]	Walter H. Ecology of tropical and subtropical vegetation [M]. Edinburgh: Oliver and Boyd, 1971.
[56]	Jackson R B, Canadell J, Ehleringer J R, et al. A global analysis of root distributions for terrestrial biomes[J]. Oecologia, 1996, 108: 389–411.
[57]	Schenk H J, Jackson R B. The global biogeography of roots[J]. Ecological Monographs, 2002, 72: 311–328.
[58]	Schenk H J, Jackson R B. Rooting depths, lateral root spreads and below-ground/above-ground allometries of plants in water-limited ecosystems[J]. Journal of Ecology, 2002, 90: 480–494.
[59]	Caldwell M M, Manwaring J H, Durham S L. The microscale distribution of neighbouring plant roots in fertile soil microsites[J]. Functional Ecology, 1991, 5: 765–772.
[60]	Caldwell M M, Manwaring J H, Durham S L. Species interactions at the level of fine roots in the field: influence of soil nutrient heterogeneity and plant size[J]. Oecologia, 1996, 106: 440–447.
[61]	Bloom A J, Chapin F S, Mooney H A. Resource limitation in plants-an economic analogy[J]. Annual Review of Ecology Evolution and Systematics, 1985, 16: 363–392.
[62]	Grime J P, Crick J C, Rincon J E. The ecological significance of plasticity[A]. Jennings D H, Trewavas A J. Plasticity in plants [C]. Cambridge, UK: The Company of Biologists, 1986.
[63]	Stephens D W, Krebs J R. Foraging theory [M]. Chichester, Sussex, UK: Princeton University Press, 1986.
[64]	张德闪. 玉米高效利用磷资源的根系互作过程及其作用机制 [D]. 北京: 中国农业大学博士论文, 2016. Zhang D S. Root-root interaction and mechanisms for improving phosphorus use efficiency by target maize when growing with different neighbours [D]. Beijing: PhD Dissertation of China Agricultural University, 2016.
[65]	Li B, Li Y Y, Wu H M, et al. Root exudates drive interspecific facilitation by enhancing nodulation and N2 fixation[J]. Proceedings of the National Academy of Sciences, 2016, 113: 6496–6501.
[66]	Shen J B, Li C J, Mi G H, et al. Maximizing root/rhizosphere efficiency to improve crop productivity and nutrient use efficiency in intensive agriculture of China[J]. Journal of Experimental Botany, 2013, 64: 1181–1192.
[67]	Shen J B, Yuan L X, Zhang J L, et al. Phosphorus dynamics: from soil to plant[J]. Plant Physiology, 2011, 156: 997–1005.
[68]	Zhu Y Y, Wang Y Y, Chen H R, et al. Conserving traditional rice through biodiversity management[J]. Nature, 2000, 406: 681–682.
[69]	Brooker R W, Bennett A E, Cong W F, et al. Improving intercropping: a synthesis of research in agronomy, plant physiology and ecology[J]. New Phytologist, 2014, 3: 1–11.
[70]	Grime J P. Plant strategies and vegetation processes [M]. London: Willey, 1997.
[71]	Tilman D. Resource competition and community structure [A]. Gavrilets S. Monographs in population biology [C]. Princeton, USA: Princeton University Press, 1982.
[72]	Rajaniemi T K, Allison V J, Goldberg D E. Root competition can cause a decline in diversity with increased productivity[J]. Journal of Ecology, 2003, 91: 407–416.
[73]	Tian Q Y, Chen F J, Liu J X, et al. Inhibition of maize root growth by high nitrate supply is correlated with reduced IAA levels in roots[J]. Journal of Plant Physiology, 2008, 165: 942–951.
[74]	Guo Y F, Mi G H, Chen F J, Zhang F S. Effect of NO3- supply on lateral root growth in maize plants[J]. Journal of Plant Physiology and Molecular Biology, 2005, 31: 90–96.
[75]	Boomsma C, Santini J, Tollenaar M, Vyn T. Maize per-plant and canopy-level morpho-physiological responses to the simultaneous stresses of intense crowding and low nitrogen availability[J]. Agronomy Journal, 2009, 101: 1426–1452.
[76]	Fransen B, de Kroon H, Berendse F. Soil nutrient heterogeneity alters competition between two perennial grass species[J]. Ecology, 2001, 82: 2534–2546.
[77]	Jing J Y, Rui Y, Zhang F S, et al. Localized application of phosphorus and ammonium improves growth of maize seedlings by stimulating root proliferation and rhizosphere acidification[J]. Field Crops Research, 2010, 119: 355–364.
[78]	Jing J Y, Zhang F S, Rengel Z, Shen J B. Localized fertilization with P plus N elicits an ammonium-dependent enhancement of maize root growth and nutrient uptake[J]. Field Crops Research, 2012, 133: 176–185.
[79]	Ma Q H, Wang X, Li H B, et al. Localized application of NH4+-N plus P enhances zinc and iron accumulation in maize via modifying root traits and rhizosphere processes[J]. Field Crops Research, 2014, 164: 107–116.
[80]	Ma Q H, Wang X, Li H B, et al. Comparing localized application of different N fertilizer species on maize grain yield and agronomic N-use efficiency on a calcareous soil[J]. Field Crops Research, 2015, 180: 72–79.