Glycine betaine mediated changes in growth, photosynthetic efficiency, antioxidant system, yield and quality of mustard
Introduction
Glycine betaine (GB) is a well-known quaternary ammonium compound (zwitterion) naturally found in haemophilic archaebacterial, bacteria, marine invertebrates, plants and animals (Chen and Murata, 2002, 2008, 2011; Figueroa-Soto and Valenzuela-Soto, 2018) and was isolated first from Beta vulgaris (Scheibler et al. 1869). GB is an organic metabolite and its extreme water-solubility and low viscosity make it a key osmoregulatory (Fariduddin et al. 2013). Besides acting as an osmoprotectant, it stabilizes quaternary structure of proteins and enzymes, improves the antioxidant defense system, photosystem-II, reduces membrane permeability and H2O2 mediated signaling (Ma et al. 2007; Chen and Murata, 2011; Oukarroum et al. 2012; Tian et al. 2017; Annunziata et al. 2019; Ali et al. 2020; Huang et al. 2020). GB generally accumulates in the cytosol, plastids, chloroplast and mitochondria of many plant species and further its accumulation increases in response to stressful conditions (Ashraf and Foolad, 2007; Park et al. 2007; Ahmed et al. 2019). In plants, GB biosynthesis occurs via a two-step oxidation pathway of choline. The first step is catalyzed by choline monooxygenase (CMO) via the oxidation of choline to an intermediate betaine aldehyde. While the second oxidation step is mediated by a NAD(P)+ dependent betaine aldehyde dehydrogenase (BADH) enzyme (Xu et al. 2018; Ali et al. 2020). Amaranthaceae, Asteraceae, Malvaceae, Portulacaceae and Poaceae are among the naturally GB accumulating families (Ashraf and Foolad, 2007; Chen and Murata, 2011). However, GB accumulation varies considerably from species to species, and in tolerant genotypes generally it accumulates at higher level than the sensitive ones. Plants utilize exogenously applied GB and translocate it to almost all plant organs. This exogenous application enhances the level of GB in plants that are unable to synthesize it. Exogenously applied or transgenes generated GB can regulate the gene expression and trigger responses to abiotic stress conditions (Ashraf and Foolad, 2007; Chen and Murata, 2011; Giri, 2011). Previous reports showed that as an osmoprotectant, GB has the potential to improve the growth and physiology of plants under normal and stressful conditions. For example, exogenously applied GB promoted crop growth and improved plant water status, accumulation of proline, CO2 fixation, water use efficiency and stomatal conductance under normal and stressed conditions (Ather et al. 2015; Ahmad et al. 2020).
Indian mustard is the second important edible oilseed crop of India and supplies about 7% of worlds edible oil. However, its production is still insufficient to fulfil the daily needs of the people (Hayat et al. 2012). Several efforts have been made to enhance its productivity by utilizing the ‘treatment-response strategies”. Keeping this in view, the present experiment was designed to explore the optimal concentration of leaf-applied GB so that the full genetic potential of the crop could be exploited.
Section snippets
Plant material
The healthy seeds of Brassica juncea L. cv. Pusa Agrani were brought from the Indian Agricultural Research Institute, New Delhi. Prior to sowing these seeds were surface sterilized by mercuric chloride (0.01%) solution for about five minutes with continuous shaking accompanied by thorough rinsing with double distilled water (DDW) three to four times to remove the mercuric chloride from the seed coats.
Treatment preparation
The stock solution of GB was prepared by dissolving the desired amount of GB in 5 ml of DDW in
Effect of GB on growth attributes
The data (Fig. 1A-G) showed that root length (RL), shoot length (SL), root fresh mass (RFM), shoot fresh mass (SFM), root dry mass (RDM), shoot dry mass (SDM) and APL were found to be increased by the foliar application of GB. The spray application of 30 mM GB gave the maximum values but its effect was statistically similar with that of 20 mM GB. The application of 20 mM GB increased RL by 12.2%, SL by 11.79%, RFM by 12.5%, RDM by 11.26%, SFM by 13.39%, SDM by 13.40% per plant and APL by 14.73%
Discussion
GB regulates diverse metabolic processes in plants under normal and myriad conditions (Ather at al., 2015). The present experiment was performed to improve the understanding of effect of GB on the performance of Indian mustard by undertaking growth, physio-biochemical, histochemical, microscopical, yield and quality analyses (Fig 1-8). The data (Fig. 1A-B, G) revealed that the GB-treated plants showed a beneficial effect on RL, SL and APL. Such improvement in these characters could be ascribed
Conclusion
The present study revealed that the foliar spray of GB enhanced the overall performance of the oilseed crop. Foliar spray of 20 mM GB proved beneficial for most of the parameters studied and this was further supported by the histochemical and confocal microscopic studies. The effects of 30 mM GB on most parameters were found to be at par with that of 20 mM GB. However, proline and GB contents were found to be increased with the foliar spray of GB in a concentration dependent manner. The foliar
Author(s) contribution
SI performed the experiment, analyzed the data and wrote the article.
ZAP and SHS helped in histochemical and microscopic studies as also in the preparation of histogram.
FM designed and supervised the experiment as well as guided the preparation of manuscript.
Declaration of Competing Interest
The authors declare that no conflict of interest exists.
Acknowledgements
SI, ZAP and SHS are thankful to UGC New Delhi, India for providing the Non-Net Fellowship. The authors are also grateful to the University Sophisticated Instrumentation Facility (USIF) AMU, Aligarh for carrying out the confocal and SEM analyses.
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