Protective and defensive role of anthocyanins under plant abiotic and biotic stresses: An emerging application in sustainable agriculture

Plant stress is defined as a sudden change in environmental conditions that adversely affect the plant growth via disruption in various physio-morphological and metabolic processes (Rivero et al., 2021). Drought, salinity, temperature, pesticides, nutrient deficiency, floods, and pathogen attack, etc. are major types of plant stresses that exacerbate crop productivity globally (Cohen and Leach, 2019). Due to continued climate change and human activities, environmental stresses are expected to worsen in the future. During stress conditions, the plant undergoes various metabolic and physiological changes such as alteration in shoot/root biomass, reduction in photosynthesis and nutrient uptake, inhibition of flowering and seed formation which ultimately diminishes the plant growth and productivity (Fig. 1). The degree of this damage is determined by the severity, frequency, and time period of the exposure (Pandey et al., 2017). Due to immobile organisms, plants are unable to move away from adverse environmental conditions. Therefore, they have developed sophisticated signalling and protective systems to cope up with stress conditions. Reactive oxygen species (ROS) molecules primarily function as signal transduction molecules that reach the nucleus through redox reactions and control several pathways during plant acclimatization to stress (Das and Roychoudhury, 2014). When the level of ROS is too high, however, it becomes exceedingly hard to manage redox homeostasis of the cell because it causes instability of cellular membranes, nucleic acid, and protein, resulting in decreased photosynthetic efficiency and, eventually plant mortality (Kruk et al., 2019). To address this issue, plants have evolved a variety of ROS-scavenging techniques, including the induction of anthocyanin molecules, that exert protective effect under stress conditions (Bartwal et al., 2013, Naing and Kim, 2021, Shah and Smith, 2020, Sytar et al., 2018, Shi et al., 2022, Marone et al., 2022). Anthocyanin accumulation is one such strategy that may confer tolerance in plants under stress situations. Anthocyanins are water-soluble compounds belonging to the flavonoid group which imparts red, purple, orange, blue and brown color to many flowers, fruits and vegetables (Kovinich et al., 2014, Gu et al., 2018). Till today, more than 30 types of monomeric anthocyanidins have been reported. Among them, only six anthocyanidins are typically found in plants: cyanidin, malvidin, pelargonidin, peonidin, petunidin, and delphinidin (Khoo et al., 2017, Gitelson et al., 2001). Anthocyanins are synthesized via the phenylpropanoid pathway (PPP) which is the most conserved pathway in the plant kingdom (Eichenberger et al., 2018). After synthesis, anthocyanins are transported to vacuole through different pathways for storage purposes (Kaur et al., 2021). Upon stress conditions, anthocyanin molecules get accumulated in different plant tissues and provide significantly antioxidant activity to induce morpho-physiological and metabolic adaptations in plants, thereby recognized as nature’s “Swiss army knife” (Fig. 1) (Gould, 2004, Naing et al., 2017) (Fig. 1). Pigmented leaves and crops also play a crucial role in stress adaptation through increasing scavenging of ROS molecules (Zhang et al., 2019, Zhang et al., 2019, Zhang et al., 2019). Moreover, different accumulation and concentration of anthocyanins also take part in the plant biodiversity adaptation and the pigmented plants with higher anthocyanin content are not only better tolerant under different stress conditions but are also good for human nutrition (Kaur et al., 2022, Sytar et al., 2018, Chen et al., 2022, Sharma et al., 2022b, Sharma et al., 2022a, Kumari et al., 2020, Garg et al., 2022). For example, a significant anthocyanin accumulation was observed under salt stress in different colored wheat genotypes (Mbarki et al., 2018). In another case, when grapevine cells were subjected to Pi-deficiency, anthocyanin accumulation was detected, which was responsible for providing tolerance under stress (Yin et al., 2012). Under stress conditions, similar observations of induction of anthocyanins biosynthesis genes have been reported in various crops which confer tolerance to various plants (Kaur et al., 2022, Perin et al., 2019, Zhang et al., 2012, Kim et al., 2017). Given their antioxidant characteristics, anthocyanins protect plants against adverse environments through scavenging stress-induced ROS molecules and are thus regarded as one of the suitable approaches that can contribute to stress tolerance in plants (Sharma et al., 2019, Naing and Kim, 2021). Although the contribution of anthocyanins related to nutrition and pharmaceuticals are well-known, but significance of anthocyanins in stress response is still little understood. Therefore, the aim of the present review is to focus on factors affecting anthocyanin induction and their protective function against various stressors in multiple plant species. We have tried to cover natural anthocyanin variation in response to stresses as well as genetic transformation to introduce anthocyanin accumulation in non-accumulators and tried to comprehensively understand the stress mitigation mechanism of anthocyanins. To accomplish this, various databases, including PubMed, Google scholar, Nature communications, and ScienceDirect were used for literature searches till October, 2022. The relevant articles were identified using the following search items: anthocyanins and stress in plants, anthocyanin and drought tolerance. Anthocyanins/flavonoids/cold/salt/heavy metal/biotic stress, anthocyanin and micronutrient deficiency, anthocyanin/antioxidants and ROS signalling, anthocyanins and genetic engineering/ transgenic/over expression etc. In addition, the reference list of the included studies and their citations were also searched.

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