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外源腐植酸对玉米拔节期干旱胁迫影响的缓解效应

Mitigative effect of exogenous humic acid on drought stress in maize at the jointing stage

  • 摘要: 为探究外源腐植酸对玉米应对干旱胁迫的生理调控机制及其阶段性响应特征,该研究通过桶栽试验,于玉米六叶期喷施不同质量浓度(0、0.50、0.75、1.00、1.25、1.50 g/L)以矿源黄腐酸为主要成分的腐植酸水溶肥(humic acid water-soluble fertilizer,HA-WSF)。试验在拔节期进行12 d干旱处理后复水8 d,系统测定叶片持水能力、叶绿素相对含量、抗氧化酶活性、氧化损伤产物及渗透调节物质等生理指标。结果表明:喷施腐植酸可通过提高叶片相对含水量与叶绿素相对含量,增强超氧化物歧化酶和过氧化氢酶活性,减少丙二醛与过氧化氢积累,并促进脯氨酸、可溶性糖及可溶性蛋白合成等多重生理途径,缓解拔节期干旱胁迫对玉米的伤害。腐植酸的调控效应呈现阶段性与浓度依赖:旱后6 d,以0.75 ~ 1.00 g/L浓度通过物理保水与激活超氧化物歧化酶和过氧化氢酶主导防御;旱后12 d,最适浓度提升至1.25 g/L,调控核心转向以脯氨酸和可溶性糖积累为核心的渗透调节。综合来看,叶面喷施0.75~1.25 g/L为最佳浓度范围。该研究系统揭示了腐植酸通过水分平衡-抗氧化防御-渗透调节多级联动机制,动态增强玉米干旱适应与复水恢复能力的生理整合模式,为理解腐植酸调控作物逆境适应的动态机制提供了一种视角。

     

    Abstract: Drought stress during the jointing stage critically constrains maize productivity, particularly under the context of increasing frequency of prolonged drought events. Humic acid water-soluble fertilizer (HA-WSF) has emerged as a promising biostimulant for enhancing crop stress tolerance, yet its dynamic regulatory mechanisms remain inadequately understood, especially regarding stage-specific responses and recovery processes. To investigate the physiological regulation and stage-specific response characteristics of exogenous humic acid in alleviating drought stress in maize, a pot experiment was conducted using the maize variety Zhengdan 958. HA-WSF at concentrations of 0, 0.50, 0.75, 1.00, 1.25, and 1.50 g/L was foliar-applied at the six-leaf stage. Plants were subsequently subjected to a 12-day moderate drought treatment at the jointing stage, followed by an 8-day rewatering period to simulate post-rainfall recovery. Physiological parameters including leaf relative water content (RWC), water loss rate (WLR), SPAD value, antioxidant enzyme activities (superoxide dismutase, SOD; catalase, CAT; peroxidase, POD), oxidative damage markers (malondialdehyde, MDA; hydrogen peroxide, H2O2), osmotic regulators (proline, Pro; soluble sugar, SS), and soluble protein (SP) were systematically measured at four time points (0, 6, 12, and 20 days after drought initiation).Results revealed that HA-WSF application alleviated drought-induced damage through integrated physiological pathways, with effects exhibiting pronounced stage- and concentration-dependent patterns. At 6 days of drought, a concentration of 0.75 g/L primarily maintained leaf hydration by reducing water loss and activating SOD and CAT, constituting an early physical–antioxidant defense. At 12 days of drought, the optimal concentration shifted to 1.00–1.25 g/L, with regulatory focus transitioning to osmotic adjustment driven by coordinated accumulation of Pro and SS, alongside sustained SOD and CAT activities. Partial least squares path modeling further demonstrated that the antioxidant system served as the core mediator in reducing oxidative damage, exhibiting a direct negative effect on oxidative damage (path coefficient = –1.053, P < 0.001) and explaining 72.2% of its variance, whereas osmotic adjustment showed no direct effect. The total effect of HA-WSF concentration on oxidative damage was –0.302, with the indirect effect via the antioxidant system accounting for 192% of the total effect, underscoring the predominant role of antioxidant defense. Notably, POD exhibited distinct functional divergence from SOD and CAT, showing significant positive correlations with MDA and H2O2, suggesting its involvement in structural defense rather than direct ROS scavenging under prolonged stress. Following rewatering, optimal recovery of SPAD value and SP content was achieved at 1.00 g/L HA-WSF, indicating accelerated photosynthetic restoration and highlighting the carryover benefits of pre-stress biostimulant application. Collectively, the optimal foliar application concentration range for enhancing drought tolerance was identified as 0.75–1.00 g/L, with lower concentrations sufficient for early protection and higher concentrations required for sustained osmotic regulation under prolonged stress. These findings elucidate a multi-level synergistic mechanism by which HA-WSF dynamically enhances drought adaptation and post-drought recovery in maize through integrated regulation of water balance, antioxidant defense, and osmotic adjustment. The results further emphasize the temporal shift in regulatory priorities from early-stage water conservation to later-stage osmotic maintenance, providing theoretical and technical references for precision application of humic acid in drought-prone maize production systems under increasingly variable rainfall patterns.

     

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