Plant competition reshapes nitrogen utilisation in native Vachellia nilotica (Fabaceae)
Anathi Magadlela

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Abstract
Vachellia nilotica (Benth.) Kyal. & Boatwr is a leguminous species subjected to competition from co-occurring grassland species in mesic grasslands, including Themeda triandra Forssk. In South Africa, Vachellia nilotica enhances soil nitrogen in nutrient-deficient mesic savanna. The addition of phosphorus and competition between the grass and legume may assist in maintaining balance between the two plant groups.
Improved P supply in nutrient-poor soils favours growth of legumes over grasses. This study aimed to assess how nutrient deficiency, P fertilisation, and competition with a dominant grass species, particularly Themeda triandra, affects the aboveground and belowground growth, N nutrition, and nodulation potential of Vachellia nilotica in the Ukulinga Grassland nutrient experiment.
Soils used in the study were collected from superphosphate-fertilised trials (+P, i.e. P-enriched) and non-superphosphate trials (−P, i.e. no P enrichment). V. nilotica seedlings were grown independently and in competition with T. triandra seedlings.
Our findings confirmed that soil in +P treatment had greater P and calcium concentrations than did the −P soils. Competition from T. triandra decreased the P concentration, specific nitrogen assimilation rates and the %NDFA in V. nilotica, but the opposite effect was observed for specific nitrogen utilisation rates.
Zero nodule formation in soils with low P concentrations and competition affected the N source reliance of V. nilotica.
The addition of P might maximise N inputs through increased legume growth performance and weaken the competitive effect of a grass species.
Keywords: acidic soils, competition, kangaroo grass, Leguminosae, nitrogen fixation, nutrient deficiency, phosphorus, Ukulinga Grassland nutrient experiment.
References
Agren GI, Franklin O (2003) Root:shoot ratios, optimization and nitrogen productivity. Annals of Botany 92, 795-800.
| Crossref | Google Scholar | PubMed |
Aydin I, Uzun F (2005) Nitrogen and phosphorus fertilization of rangelands affects yield, forage quality and the botanical composition. European Journal of Agronomy 23(1), 8-14.
| Crossref | Google Scholar |
Bhadouria R, Singh R, Srivastava P, Tripathi S, Raghubanshi AS (2017) Interactive effect of water and nutrient on survival and growth of tree seedlings of four dry tropical tree species under grass competition. Tropical Ecology 58, 611-621.
| Google Scholar |
Bhatti AA, Haq S, Bhat RA (2017) Actinomycetes benefaction role in soil and plant health. Microbial Pathogenesis 111, 458-467.
| Crossref | Google Scholar | PubMed |
Cramer MD, Chimphango SBM, Van Cauter A, Waldram MS, Bond WJ (2007) Grass competition induces N2 fixation in some species of African Acacia. Journal of Ecology 95, 1123-1133.
| Crossref | Google Scholar |
Cramer MD, Van Cauter A, Bond WJ (2010) Growth of N2-fixing African savanna Acacia species is constrained by below-ground competition with grass. Journal of Ecology 98, 156-167.
| Crossref | Google Scholar |
Crews TE, Peoples MB (2004) Legume versus fertilizer sources of nitrogen: ecological tradeoffs and human needs. Agriculture, Ecosystems & Environment 102(3), 279-297.
| Crossref | Google Scholar |
De Gerónimo E, Aparicio VC (2022) Changes in soil pH and addition of inorganic phosphate affect glyphosate adsorption in agricultural soil. European Journal of Soil Science 73(1), 13188.
| Crossref | Google Scholar |
Donzelli D, De Michele C, Scholes RJ (2013) Competition between trees and grasses for both soil water and mineral nitrogen in dry savannas. Journal of Theoretical Biology 332, 181-190.
| Crossref | Google Scholar | PubMed |
Ferguson BJ, Indrasumunar A, Hayashi S, Lin MH, Lin YH, Reid DE, Gresshoff PM (2010) Molecular analysis of legume nodule development and autoregulation. Journal of Integrative Plant Biology 52, 61-76.
| Crossref | Google Scholar | PubMed |
Fitter AH (1977) Influence of mycorrhizal infection on competition for phosphorus and potassium by two grasses. New Phytologist 79, 119-125.
| Crossref | Google Scholar |
Fox J, Weisberg S (2019) ‘An R companion to applied regression.’ 3rd edn. (Sage: Thousand Oaks, CA, USA). Available at https://www.john-fox.ca/Companion/
Fynn RWS, O’Connor TG (2005) Determinants of community organization of a South African mesic grassland. Journal of Vegetation in Science 16, 93-102.
| Crossref | Google Scholar |
Han M, Zhang H, Liu M, Tang J, Guo X, Ren W, Zhao Y, Yang Q, Guo B, Han Q, Feng Y, Feng Z, Wu H, Yang X, Kong D (2024) Increased dependence on nitrogen-fixation of a native legume in competition with an invasive plant. Plant Diversity 46, 510-518.
| Crossref | Google Scholar |
Høgh-Jensen H, Schjoerring JK (1997) Interactions between white clover and ryegrass under contrasting nitrogen availability: N2 fixation, N fertilizer recovery, N transfer, and water use efficiency. Plant and Soil 197, 187-199.
| Crossref | Google Scholar |
Høgh-Jensen H, Schjoerring JK (2010) Interactions between nitrogen, phosphorus and potassium determine growth and N2-fixation in white clover and ryegrass leys. Nutrient Cycling in Agroecosystems 87, 327-338.
| Crossref | Google Scholar |
Horst WJ, Kamh M, Jibrin JM, Chude VO (2001) Agronomic measures for increasing P availability to crops. Plant and Soil 237, 211-223.
| Crossref | Google Scholar |
Hortal S, Lozano YM, Bastida F, Armas C, Moreno JL, Garcia MC, Pugnaire FI (2017) Plant–plant competition outcomes are modulated by plant effects on the soil bacterial community. Scientific Reports 7, 17756.
| Crossref | Google Scholar | PubMed |
Iqbal S, Akhtar J, Naz T, Riaz U, Hussain S, Mazhar Z, Iqbal MM (2020) Root morphological adjustments of crops to improve nutrient use efficiency in limited environments. Communications in Soil Science and Plant Analysis 51, 2452-2465.
| Crossref | Google Scholar |
Jhariya MK, Banerjee A, Yadav DK, Raj A (2018) Leguminous trees an innovative tool for soil sustainability. In ‘Legumes for soil health and sustainable management’. (Eds R Meena, A Das, G Yadav, R Lal) pp. 315–345. (Springer: Singapore) 10.1007/978-981-13-0253-4_10
Ledgard SF (2001) Nitrogen cycling in low input legume-based agriculture, with emphasis on legume/grass pastures. Plant and Soil 228, 43-59.
| Crossref | Google Scholar |
Li X, Zeng R, Liao H (2016) Improving crop nutrient efficiency through root architecture modifications. Journal of Integrative Plant Biology 58, 193-202.
| Crossref | Google Scholar | PubMed |
Liebman M, Dyck E (1993) Crop rotation and intercropping strategies for weed management. Ecological Applications 3(1), 92-122.
| Crossref | Google Scholar |
Magadlela A, Kleinert A, Dreyer LL, Valentine AJ (2014) Low-phosphorus conditions affect the nitrogen nutrition and associated carbon costs of two legume tree species from a Mediterranean-type ecosystem. Australian Journal of Botany 62, 1-9.
| Crossref | Google Scholar |
Magadlela A, Vardien W, Kleinert A, Steenkamp ET, Valentine AJ (2016) Variable P supply affects N metabolism in a legume tree, Virgilia divaricata, from nutrient-poor Mediterranean-type ecosystems. Functional Plant Biology 43, 287-297.
| Crossref | Google Scholar | PubMed |
Magadlela A, Makhaye N, Pérez-Fernández M (2021) Symbionts in Mucuna pruriens stimulate plant performance through nitrogen fixation and improved phosphorus acquisition. Journal of Plant Ecology 14, 310-322.
| Crossref | Google Scholar |
Matiwane SE, Aremu AO, Valentine AJ, Magadlela A (2019) Nutritional status of KwaZulu-Natal soils affects microbe symbiosis, nitrogen utilization and growth of Vigna radiata (L.) R. Walczak. South African Journal of Botany 126, 115-120.
| Crossref | Google Scholar |
Matson PA, Parton WJ, Power AG, Swift MJ (1997) Agricultural intensification and ecosystem properties. Science 277(5325), 504-509.
| Crossref | Google Scholar | PubMed |
Mendoza R, García I, Depalma D, Fernández López C (2016) Competition and growth of a grass–legume mixture fertilised with nitrogen and phosphorus: effect on nutrient acquisition, root morphology and symbiosis with soil microorganisms. Crop & Pasture Science 67, 629-640.
| Crossref | Google Scholar |
Mortimer PE, Pérez-Fernández MA, Valentine AJ (2008) The role of arbuscular mycorrhizal colonization in the carbon and nutrient economy of the tripartite symbiosis with nodulated Phaseolus vulgaris. Soil Biology and Biochemistry 40, 1019-1027.
| Crossref | Google Scholar |
Ndabankulu K, Tsvuura Z, Magadlela A (2022) Alien invasive Leucaena leucocephala successfully acquires nutrients by investing in below-ground biomass compared to native Vachellia nilotica in nutrient-amended soils in South Africa. AoB Plants 14, plac026.
| Crossref | Google Scholar |
Ndzwanana Z, Tsvuura Z, Valentine AJ, Pérez-Fernández MA, Magadlela A (2019) Differential patterns of nitrogen nutrition and growth cost of the indigenous Vachellia sieberiana and the introduced Chromolaena odorata in the savanna environment. AoB Plants 11, plz008.
| Crossref | Google Scholar |
Nielsen KL, Bouma TJ, Lynch JP, Eissenstat DM (1998) Effects of phosphorus availability and vesicular–arbuscular mycorrhizas on the carbon budget of common bean (Phaseolus vulgaris). New Phytologist 139, 647-656.
| Crossref | Google Scholar |
Pirhofer-Walzl K, Rasmussen J, Høgh-Jensen H, Eriksen J, Soegaard K, Rasmussen J (2012) Nitrogen transfer from forage legumes to nine neighbouring plants in a multi-species grassland. Plant Soil 350, 71-84.
| Crossref | Google Scholar |
R Core Team (2025) ‘R: a Language and Environment for Statistical Computing.’ (R Foundation for Statistical Computing: Vienna, Austria) Available at https://www.R-project.org/
Rajendren K, Mohan E (2014) Tree species with potential of nitrogen fixation in agroforestry system adopted by farmers in semi-arid region of southern India. Biological Research Bulletin 3, 1-4.
| Google Scholar |
Rather LJ, Shahid-ul-Islam, Mohammad F (2015) Acacia nilotica (L.): a review of its traditional uses, phytochemistry, and pharmacology. Sustainable Chemistry and Pharmacy 2, 12-30.
| Crossref | Google Scholar |
Sanchez PA, Shepherd KD, Soule MJ, Place FM, Buresh RJ, Izac AMN, Uzo Mokwunye AU, Kwesiga FR, Ndiritu CG, Woomer PL (1997) Soil fertility replenishment in Africa: an investment in natural resource capital. In ‘Replenishing soil fertility in Africa’. SSSA Special Publication No. 51. (Eds RJ Buresh, PA Sanchez, F Calhoun) pp. 1–46. (SSSA: Madison, WI, USA)
Sithole N, Tsvuura Z, Kirkman K, Magadlela A (2021) Nitrogen source preference and growth carbon costs of Leucaena leucocephala (Lam.) de Wit saplings in South African grassland soils. Plants 10(11), 2242.
| Crossref | Google Scholar | PubMed |
Stomph T, Dordas C, Baranger A, de Rijk J, Dong B, Evers J, Gu C, Li L, Simon J, Jensen ES, Wang Q, Wang Y, Wang Z, Xu H, Zhang C, Zhang L, Zhang WP, Bedoussac L, van der Werf W (2020) Chapter One – Designing intercrops for high yield, yield stability and efficient use of resources: are there principles? Advances in Agronomy 160, 1-50.
| Crossref | Google Scholar |
Sulieman S, Ha CV, Schulze J, Tran LP (2013) Growth and nodulation of symbiotic Medicago truncatula at different levels of phosphorus availability. Journal of Experimental Botany 64, 2701-2712.
| Crossref | Google Scholar | PubMed |
Temperton VM, Mwangi PN, Scherer-Lorenzen M, Schmid B, Buchmann N (2007) Positive interactions between nitrogen-fixing legumes and four different neighbouring species in a biodiversity experiment. Oecologia 151, 190-205.
| Crossref | Google Scholar | PubMed |
Thomas D, Sumberg JE (1995) A review of the evaluation and use of tropical forage legumes in sub-Saharan Africa. Agriculture, Ecosystems and Environment 54, 151-163.
| Crossref | Google Scholar |
Vardien W, Mesjasz-Przybylowicz J, Przybylowicz WJ, Wang Y, Steenkamp ET, Valentine AJ (2014) Nodules from Fynbos legume Virgilia divaricata have high functional plasticity under variable P supply levels. Journal of Plant Physiology 171, 1732-1739.
| Crossref | Google Scholar | PubMed |
Veneklaas EJ, Lambers H, Bragg J, Finnegan PM, Lovelock CE, Plaxton WC, Price CA, Scheible WR, Shane MW, White PJ, Raven JA (2012) Opportunities for improving phosphorus-use efficiency in crop plants. New Phytologist 195, 306-320.
| Crossref | Google Scholar | PubMed |