Plant secondary metabolites (SMs) play important roles in plant survival and in creating ecological connections between other species. In addition to providing a variety of valuable natural products, secondary metabolites help protect plants against pathogenic attacks and environmental stresses. Given their sessile nature, plants must protect themselves from such situations through accumulation of these bioactive compounds. Indeed, secondary metabolites act as herbivore deterrents, barriers against pathogen invasion, and mitigators of oxidative stress. The accumulation of SMs are highly dependent on environmental factors such as light, temperature, soil water, soil fertility, and salinity. For most plants, a change in an individual environmental factor can alter the content of secondary metabolites even if other factors remain constant. In this review, we focus on how individual environmental factors affect the accumulation of secondary metabolites in plants during both biotic and abiotic stress conditions. Furthermore, we discuss the application of abiotic and biotic elicitors in culture systems as well as their stimulating effects on the accumulation of secondary metabolites. Specifically, we discuss the shikimate pathway and the aromatic amino acids produced in this pathway, which are the precursors of a range of secondary metabolites including terpenoids, alkaloids, and sulfur- and nitrogen-containing compounds. We also detail how the biosynthesis of important metabolites is altered by several genes related to secondary metabolite biosynthesis pathways. Genes responsible for secondary metabolite biosynthesis in various plant species during stress conditions are regulated by transcriptional factors such as WRKY, MYB, AP2/ERF, bZIP, bHLH, and NAC, which are also discussed here.

Plant secondary metabolites, especially flavonoids, are major metabolites widely found in plants that play several key roles in plant defence and signalling in response to stress conditions. The most studied among these flavonoids are kaempferol and quercetin due to their anti-oxidative potential and their key roles in the defence system, making them more critical for plant adaptation in stress environments. Kaempferol and quercetin in plants have great therapeutic potential for human health. Despite being well-studied, some of their functional aspects regarding plants and human health need further evaluation. This review summarizes the emerging potential of kaempferol and quercetin in terms of antimicrobial activity, bioavailability and bioactivity in the human body as well as in the regulation of plant defence in response to stresses and as a signalling molecule in terms of hormonal modulation under stress conditions. We also evaluated the safe use of both metabolites in the pharmaceutical industry.

Melatonin is a multifunctional signaling molecule that is ubiquitously distributed in different parts of a plant and responsible for stimulating several physio-chemical responses to adverse environmental conditions. In this review, we show that, although plants are able to biosynthesize melatonin, the exogenous application of melatonin to various crops can improve plant growth and development in response to various abiotic and biotic stresses (e.g., drought, unfavorable temperatures, high salinity, heavy metal contamination, acid rain, and combined stresses) by regulating antioxidant machinery of plants. Current knowledge suggests that exogenously applied melatonin can enhance the stress tolerance of plants by regulating both the enzymatic and non-enzymatic antioxidant defense systems. Enzymic antioxidants upregulated by exogenous melatonin include superoxide dismutase, catalase, glutathione peroxidase, and enzymes involved in the ascorbate–glutathione cycle (ascorbate peroxidase, monodehydroascorbate reductase, dehydroascorbate reductase, and glutathione reductase), whereas levels of non-enzymatic antioxidants such as ascorbate, reduced glutathione, carotenoids, tocopherols, and phenolics are also higher under stress conditions. The enhanced antioxidant system consequently exhibits lower lipid peroxidation and greater plasma membrane integrity when under stress. However, these responses vary greatly from crop to crop and depend on the intensity and type of stress, and most studies to date have been conducted under controlled conditions. This means that a wider range of crop field trials and detailed transcriptomic analysis are required to reveal the gene regulatory networks involved in the between melatonin, antioxidants, and abiotic stress.

The tolerance of plant growth-promoting endophytes (PGPEs) against various concentrations of cadmium (Cd) and nickel (Ni) was investigated. Two glutathione-producing bacterial strains (Enterobacter ludwigii SAK5 and Exiguobacterium indicum SA22) were screened for Cd and Ni accumulation and tolerance in contaminated media, which showed resistance up to 1.0 mM. Both strains were further evaluated by inoculating specific plants with the bacteria for five days prior to heavy metal treatment (0.5 and 1.0 mM). The enhancement of biomass and growth attributes such as the root length, shoot length, root fresh weight, shoot fresh weight, and chlorophyll content were compared between treated inoculated plants and treated non-inoculated plants. Both strains significantly increased the accumulation of Cd and Ni in inoculated plants. The accumulation of both heavy metals was higher in the roots than in the shoots, however; Ni accumulation was greater than Cd. Heavy metal stress-responsive genes such as OsGST, OsMTP1, and OsPCS1 were significantly upregulated in treated non-inoculated plants compared with treated inoculated plants, suggesting that both strains reduced heavy metal stress. Similarly, abscisic acid (ABA) was increased with increased heavy metal concentration; however, it was reduced in inoculated plants compared with non-inoculated plants. Salicylic acid (SA) was found to exert synergistic effects with ABA. The application of suitable endophytic bacteria can protect against heavy metal hyperaccumulation by enhancing detoxification mechanisms.

As a result of climate change, crops often experience high-temperature stress that can drastically hinder plant growth and development. In soybean, an economically important crop that is highly sensitive to heat stress, the use of plant growth-promoting rhizobacteria (PGPRs) represents a promising strategy for countering the negative effects of heat stress. Accordingly, a novel strain of Bacillus tequilensis (i.e. SSB07) that grows actively at high temperatures was isolated, identified, and characterized in the present study. SSB07 improved the growth of Chinese cabbage seedlings and was shown to produce the gibberellins GA₁, GA₃, GA₅, GA₈, GA₁₉, GA₂₄, and GA₅₃, as well as indole-3-acetic acid and abscisic acid. The application of B. tequilensis SSB07 was also found to increase the shoot length and biomass, leaf development, and photosynthetic pigment contents of soybean plants, and under heat stress, SSB07 inoculation significantly increased the endogenous jasmonic acid and salicylic acid contents of the phyllosphere and significantly down-regulated the production of stress-responsive ABA. Thus, B. tequilensis SSB07 shows promise for countering the negative effects of climate change on crop growth and development.

Drought and ultraviolet radiation (UV radiation) are the coexisting environmental factors that negatively affect plant growth and development via oxidative damage. Flavonoids are reactive, scavenging oxygen species (ROS) and UV radiation-absorbing compounds generated under stress conditions. We investigated the biosynthesis of kaempferol and quercetin in wild and flavanone 3-hydroxylase (F3H) overexpresser rice plants when drought and UV radiation stress were imposed individually and together. Phenotypic variation indicated that both kinds of stress highly reduced rice plant growth parameters in wild plants as compared to transgenic plants. When combined, the stressors adversely affected rice plant growth parameters more than when they were imposed individually. Overaccumulation of kaempferol and quercetin in transgenic plants demonstrated that both flavonoids were crucial for enhanced tolerance to such stresses. Oxidative activity assays showed that kaempferol and quercetin overaccumulation with strong non-enzymatic antioxidant activity mitigated the accumulation of ROS under drought and UV radiation stress. Lower contents of salicylic acid (SA) in transgenic plants indicated that flavonoid accumulation reduced stress, which led to the accumulation of low levels of SA. Transcriptional regulation of the dehydrin (DHN) and ultraviolet-B resistance 8 (UVR8) genes showed significant increases in transgenic plants compared to wild plants under stress. Taken together, these results confirm the usefulness of kaempferol and quercetin in enhancing tolerance to both drought and UV radiation stress.