Humic Acid Alleviates Aluminum Toxicity in Citrus grandis (L.) Osbeck: Insight from Growth, Gas Exchange, and Related...

Most Citrus spp. trees in China are cultivated in acidic soils with low soil organic matter and high Al³⁺. The mechanisms of humic acid (HA) to alleviate Al³⁺ stress in plants remain unclear. 'Sour pummelo' (Citrus grandis (L.) Osbeck) seedlings were exposed to 0.5 (HA0.5), 0.1 (HA0.1), or 0 (HA0) mM sodium humate and 1.2 (Al1.2) or 0 (Al0) mM AlCl₃·6H₂O for 128 days. Thereafter, the research examined biomass; Al and mineral nutrients; leaf photosynthetic performance; and leaf and root nonstructural carbohydrates, reactive oxygen species metabolism, and related physiological parameters. Al1.2 significantly reduced whole plant dry weight (DW), root DW, leaf CO₂ assimilation (A_CO2), and chlorophyll a + b concentration by 61%, 45%, 61%, and 35%, respectively, at HA0, but only 48%, 17%, 44%, and 11%, respectively, at HA0.5. Further analysis suggested that the addition of HA endowed Citrus with Al resilience by the following several aspects: (a) lessened tissue (leaf, stem, and root) concentrations of Al and enhanced capacity to maintain macronutrient (S, K, Mg, Ca, N, and P) homeostasis at Al1.2; (b) improved capacity to combat oxidative stress at Al1.2; and (c) enhanced A_CO2 and growth at Al1.2. Further analysis indicated that HA-mediated alleviation of growth decline caused by Al1.2 involved (a) reduced ability to absorb Al and less root-to-shoot Al transport and (b) increased ability to maintain macronutrient homeostasis and to combat oxidative stress; and that HA-mediated alleviation of leaf chlorophyll and A_CO2 decline and photosynthetic electron transport chain impairment involved less leaf Al concentration and improved leaf macronutrient homeostasis. To conclude, the addition of HA lowered roots' ability to absorb Al and tissue Al concentration and subsequently mitigated Al-toxic impairment to root growth and function, thereby enhancing the ability of plants to maintain macronutrient homeostasis, and hence alleviating Al1.2-stimulated oxidative damage and inhibition of A_CO2 and growth.