Hydrogen Sulfide Interacts with 5-Aminolevulinic Acid to Enhance the Antioxidant Capacity of Pepper (Capsicum annuum L.) Seedlings under Chilling Stress

5-Aminolevulinic acid (ALA) is the precursor of tetrapyrrole synthesis, and hydrogen sulfide (H2S) is a gas signal molecule. Studies have shown that exogenous ALA and H2S can alleviate abiotic stress. This study evaluated the roles of ALA and H2S and their interactions in regulating antioxidant activity in pepper seedlings under chilling stress. Chilling stress significantly inhibited the growth of pepper seedlings and increased the amounts of hydrogen peroxide (H2O2), superoxide anion (O2(center dot-)), and malondialdehyde (MDA). ALA and/or H2S increased the activities of superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT). Moreover, ALA and/or H2S enhanced the ascorbate (AsA)-glutathione (GSH) cycle by increasing the contents of AsA and GSH, the ratio of AsA to dehydroascorbic acid and GSH to glutathione disulfide increased, and the activities of ascorbate peroxidase (APX), glutathione reductase (GR), dehydroascorbate reductase (DHAR) and monodehydroascorbate reductase (MDHAR) also increased. At the transcriptional level, ALA and/or H2S upregulated the expressions of CaSOD, CaPOD, CaCAT, CaAPX, CaGR, CaDHAR, and CaMDHAR in seedlings under chilling stress. ALA and/or H2S also reduced the contents of H-2 O-2,O-2(center dot-) and MDA, eventually mitigating the inhibitory effects of chilling stress on pepper seedling growth. The combination of ALA and H2S had a better effect than ALA or H2S alone. Moreover, ALA and H2S interact to regulate the oxidative stress response of pepper seedlings under chilling stress.