Humic Acid Modulates Ionic Homeostasis, Osmolytes Content, and Antioxidant Defense to Improve Salt Tolerance in Rice

Abstract

The sensitivity of rice plants to salinity is a major challenge for rice growth and productivity in the salt-affected lands. Priming rice seeds in biostimulants with stress-alleviating potential is an effective strategy to improve salinity tolerance in rice. However, the mechanisms of action of these compounds are not fully understood. Herein, the impact of priming rice seeds (cv. Giza 179) with 100 mg/L of humic acid on growth and its underlaying physiological processes under increased magnitudes of salinity (EC = 0.55, 3.40, 6.77, 8.00 mS/cm) during the critical reproductive stage was investigated. Our results indicated that salinity significantly reduced Giza 179 growth indices, which were associated with the accumulation of toxic levels of Na⁺ in shoots and roots, a reduction in the K⁺ and K⁺/Na⁺ ratio in shoots and roots, induced buildup of malondialdehyde, electrolyte leakage, and an accumulation of total soluble sugars, sucrose, proline, and enzymic and non-enzymic antioxidants. Humic acid application significantly increased growth of the Giza 179 plants under non-saline conditions. It also substantially enhanced growth of the salinity-stressed Giza 179 plants even at 8.00 mS/cm. Such humic acid ameliorating effects were associated with maintaining ionic homeostasis, appropriate osmolytes content, and an efficient antioxidant defense system. Our results highlight the potential role of humic acid in enhancing salt tolerance in Giza 179.

Keywords: antioxidants; humic acid priming; osmolytes; rice; salinity.

Figure 1

Effect of seed priming with humic acid (100 mg/L) on oxidative stress parameters (A) MDA content and (B) EL percentage of Giza 179 plants grown under increased levels of seawater. Values are the means ± SE of three replicates. Different bar letters indicate significant differences at p < 0.05.

Figure 2

Effect of seed priming with humic acid (100 mg/L) on osmolyte content (A) total soluble sugars, (B) sucrose, (C) total carbohydrates, (D) soluble proteins, and (E) proline content of Giza 179 plants grown under increased levels of seawater. Values are the means ± SE of three replicates. Different bar letters indicate significant differences at p < 0.05.

Figure 3

Effect of seed priming with humic acid (100 mg/L) on antioxidant activities, (A) total flavonoids, (B) total phenols, (C) catalase activity (CAT), and (D) peroxidase activity (POX) of Giza 179 plants grown under increased levels of seawater. Values are the means ± SE of three replicates. Different bar letters indicate significant differences at p < 0.05.

Figure 3

Effect of seed priming with humic acid (100 mg/L) on antioxidant activities, (A) total flavonoids, (B) total phenols, (C) catalase activity (CAT), and (D) peroxidase activity (POX) of Giza 179 plants grown under increased levels of seawater. Values are the means ± SE of three replicates. Different bar letters indicate significant differences at p < 0.05.

Figure 4

Principal component analysis (PCA) of growth, physiological, biochemical, and mineral traits in the investigated humic acid (100 mg/L) treated and non-treated Giza 179 plants grown under increased levels of seawater. (A) PCA score plot and (B) PCA biplot of 21 measured traits.

Figure 5

Heatmap of Pearson correlation analysis of all investigated traits in humic acid (100 mg/L) treated and non-treated Giza 179 plants grown under increased levels of seawater. Red and blue colors represent positive and negative correlations, respectively, according to the color scale.