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- Herausgeber: John Wiley & Sons
- Kategorie: Wissenschaft und neue Technologien
- Sprache: Englisch
MANAGING PLANT STRESS USING SALICYLIC ACID Enables readers to understand the ability of salicylic acid in reducing the effects of abiotic stresses in different crop species Salicylic acid is an important plant hormone which acts as a multifunctional molecule and regulates key physiological and biochemical processes in plants. This book highlights the tremendous potential of treating plants with salicylic acid, either prior to or during stress. It focuses on the specific challenges and opportunities related to exogenous application or priming technology, such as the mode of application, new methodologies, and the potential impacts of salicylic acid on the environment. Sample topics covered in the book include: * The latest research on the ability of salicylic acid in reducing the effects of abiotic stresses in different crop species * The mechanism of action of salicylic acid at the biochemical and molecular level * Salicylic acid and its crosstalk with other plant hormones under stressful environments * Regulation of abiotic stress by salicylic acid at the gene level * The role of salicylic acid on the postharvest physiology of plants This book will be of significant interest to researchers, academics, and scientists working in the field of salicylic acid mediated responses in plants under challenging environments and with abiotic stress tolerance.
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Veröffentlichungsjahr: 2022
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Table of Contents
Title Page
Copyright Page
List of Contributors
Preface
1 Salicylic Acid: A Regulator of Plant Growth and Development
Introduction
Salicylic Acid and Plant Growth
Salicylic Acid and Photosynthesis
Salicylic Acid and Respiration
Salicylic Acid and Abiotic and Biotic Stress
Conclusions
References
2 Salicylic Acid‐Mediated Regulation of Plant Biology: An Omics Approach
Introduction
Programmed Cell Death (PCD)
SA as a Plant Growth Regulator
SA and Stresses
Resistance of Plants to Diseases
Proteome Analysis
Transcriptome Analysis
Genome Analysis
Conclusion
References
Note
3 Regulation of Plant Primary Metabolism by Salicylic Acid Under Abiotic Stress
Introduction
Primary Metabolism Regulation
SA Mediation and Abiotic Stress Tolerance Regulation
Conclusion
References
Appendix 3.A
Appendix 3.B
4 Regulation of Plant Secondary Metabolism by Salicylic Acid Under Abiotic Stress
Introduction
Biosynthesis Pathway of Different Secondary Metabolites in Plants
The Biosynthesis of SMs in Plants Exposed to Abiotic Stresses
Elicitation of Secondary Metabolites by Exogenously Applied Salicylic Acid Under Abiotic Stress
Conclusion and Future Prospects
References
5 How Does Salicylic Acid Regulate Mineral Nutrition in Plants Under Abiotic Stress? An Update
Introduction
Salicylic Acid and Regulation of Mineral Nutrition
Conclusion and Future Prospects
References
6 Seed Germination to Fruit Maturation Under Stressful Environment: Roles of Salicylic Acid
Introduction
Plant Face to Environmental Stress
Salicylic Acid as Plant Growth Regulator
Function of Salicylic Acid in Plant During Stressful Factors
Conclusion
References
Appendix 6.A
Appendix 6.B
7 Role of Salicylic Acid on Postharvest Physiology of Plants
Introduction
Biosynthesis and Metabolism of Salicylic Acid
Preharvest Factors Affecting Postharvest Quality
Physiological Roles of Salicylic Acid on Plants
Conclusion
References
8 Salicylic Acid‐Mediated Physiological and Molecular Mechanism in Plants Under Metal(loid) Stress
Introduction
Sources of Metalloids and Their Availability to the Plants
Effects of Metalloids in Plants and Tolerance of Plants to Stress
Phytohormones Used for Ameliorating the Metalloid Stress
Physiological Roles of Salicylic Acid in Plants Under Metal/Metalloid Stress
Omics‐Based Strategies for SA‐Induced Metalloid Tolerance in Plants
Conclusion
References
9 Salicylic Acid‐Mediated Physiological and Molecular Mechanisms in Plants Under Heat Stress
Introduction
Salicylic Acid History and Biosynthesis
Heat‐Stress Threshold
Physiological Mechanisms Mediated by SA Under HS
Molecular Mechanisms Mediated by Salicylic Acid Under Heat Stress
Conclusion and Future Prospects
References
10 Salicylic Acid‐Mediated Physiological and Antioxidant Enzyme Activity Mechanisms in Plants Under Chilling Stress
Introduction
Effect of Chilling and Salicylic Acid (SA) on Growth Factors
Conclusion
References
11 Salicylic Acid‐Mediated Physiological and Molecular Mechanisms in Plants Under Abiotic Stress
Introduction
Stress Hormones
Background of Salicylic Acid
Biosynthesis of Salicylic Acid in Plants
Salicylic Acid in Plants Under Metals/Metalloids
Epigenetics Functions in Salicylic Acid
Conclusion and Future Prospects
Conflict of Interest
Acknowledgements
References
12 Salicylic‐Acid Mediated Physiological and Molecular Mechanisms in Plants Under Drought Stress
Introduction
Adversities of Drought Stress
Role of SA in Photosynthesis and Sugars Biosynthesis under Drought Stress
Changes in ROS and Enzymatic Antioxidants in Plants Subjected to Drought Stress and Role of SA
Role of SA on Polyphenols Biosynthesis
Role of SA on Osmolytes Accumulation
Role of SA on Mineral Nutrients Uptake and Accumulation
Interaction of SA with Polyamines and Other Phytohormones in Plants Subjected to Drought Stress
Role of SA on Induction of Gene Expression Under Drought
Conclusions
References
13 ROS Regulation by Salicylic Acid Under Abiotic Stress
Introduction
Abiotic Stress in Plants and Activation of ROS Signaling
SA and ROS Interaction
SA Mediated Regulation of ROS Level (ROS Homeostasis)
SA‐Mediated ROS Detoxification (Activation of Antioxidant Defense System)
Conclusion and Future Perspective
References
14 Regulation of Photosynthesis by Salicylic Acid Under Optimal and Suboptimal Conditions
Introduction
Salicylic Acid (SA) and Regulation of Photosynthesis
Salicylic Acid as a Stress Factor for Plants
Effect of Lower Levels of Salicylic Acid on Plants
Effect of Elevated Levels of Salicylic Acid on Plants
Conclusion
References
15 Regulation of Abiotic Stress by Salicylic Acid at Gene Level: An Update
Introduction
Biosynthesis of Salicylic Acid
Metabolism of Salicylic Acid
Salicylic Acid Receptors in Plants
Salicylic Acid Transport in Plants
Implication of Salicylic Acid in Plants’ Abiotic Stress Tolerance
Conclusion
Acknowledgements
Author Contributions
References
16 Salicylic Acid and its Crosstalk with Other Plant Hormones Under Stressful Environments
Introduction
Function of SA Under Biotic Stress Conditions (Pathogen Defense Pathway)
Function of SA Under Abiotic Stress Conditions
Crosstalk of Salicylic Acid with Other Plant Hormones Under Stressful Environments
Conclusion and Perceptions
References
17 Post‐translational Modifications Mediated by the Phytohormone Salicylic Acid in Plants Growing Under Environmental Challenges
Introduction
SA Biosynthetic and Signaling Pathways
SA‐Induced Responses During Abiotic Stress
The Dual Role of SA in Redox Homeostasis During Abiotic Stress
Proteome Modifications Mediated by SA
How to Identify New PTMs Induced by SA
Perspectives
Acknowledgments
References
Index
End User License Agreement
List of Tables
Chapter 3
Table 3.B.1 Protective effects of SA in vegetal development under abiotic c...
Chapter 4
Table 4.1 Examples of changes in phytochemicals of plants under abiotic str...
Table 4.2 Phytochemicals alteration examples of plants by exogenously appli...
Chapter 5
Table 5.1 Salicylic acid‐induced improvement of different plant traits and ...
Chapter 6
Table 6.B.1 Roles of salicylic acid and beneficial impacts on crop cultures...
Chapter 7
Table 7.1 Influence of salicylic acid (SA) on the physiology of perishable ...
Chapter 8
Table 8.1 Effects of metals/metalloids on plants.
Table 8.2 SA‐induced effects in plants under metal/metalloid stress.
Table 8.3 Omics‐based strategies in SA‐regulated metalloid tolerance in pla...
Chapter 9
Table 9.1 HS threshold levels for some crops.
Table 9.2 Effect of SA on plant growth and physiological attributes under H...
Chapter 12
Table 12.1 SA‐mediated physiological responses under drought stress.
Table 12.2 Genes regulated by SA treatment under drought stress.
Chapter 14
Table 14.1 Effect of SA on the photosynthesis and photosynthetic pigments u...
List of Illustrations
Chapter 1
Figure 1.1 Schematization of the role of salicylic acid in plants.
Chapter 2
Figure 2.1 The hypothesized molecular mechanism of SA application on
Cucumis
...
Chapter 3
Figure 3.A.1 Control of physiological mechanisms implicating SA under abioti...
Figure 3.A.2 Different salicylic acid biosynthesis pathways and responses to...
Chapter 4
Figure 4.1 Effect of different abiotic stress and exogenous application of s...
Chapter 5
Figure 5.1 Influence of abiotic stress on plant and alleviation of oxidative...
Chapter 6
Figure 6.A.1 Plant face to environmental stress during life cycle.
Figure 6.A.2 Some functions of salicylic acid during the main stages of plan...
Chapter 7
Figure 7.1 Pathway of biosynthesis of salicylic acid (Beckers and Spoel 2006...
Figure 7.2 Preharvest factors affecting the postharvest quality of perishabl...
Figure 7.3 Brief overview of processes by which salicylic acid (SA) delays t...
Figure 7.4 Induction of chilling resistance in plants by salicylic acid. The...
Figure 7.5 Antioxidant systems in plants. Under normal conditions, cellular ...
Figure 7.6 Salicylic acid (SA) reduces the production of catalase (CAT), and...
Chapter 8
Figure 8.1 Uptake of heavy metals and metalloids by plants.
Figure 8.2 Illustration of the role of various phytohormones in ameliorating...
Figure 8.3 SA‐regulated Omics‐based approaches to identify stress‐responsive...
Chapter 10
Figure 10.1 The effects of chilling and salicylic acid (SA) on (a) shoot MDA...
Figure 10.2 The effects of chilling and salicylic acid on the activity of ca...
Figure 10.3 The effects of chilling and salicylic acid on H
2
O
2
content in (a...
Chapter 11
Figure 11.1 A model shows the biosynthesis of salicylic acid (SA) pathway st...
Chapter 13
Figure 13.1 Summarized form of synchronized working of antioxidative system ...
Figure 13.2 Depicting complexities of ROS signaling inside an abiotically st...
Figure 13.3 Role of SA to mitigate ROS in plants.
Chapter 14
Figure 14.1 Elicit role of SA on photosynthesis under optimal and suboptimal...
Chapter 15
Figure 15.1 Mechanisms affecting SA biosynthesis in the shikimate pathway in...
Figure 15.2 The phenylalanine ammonia‐lyase (PAL) pathway for SA biosynthesi...
Figure 15.3 Glycosylation of SA in
Arabidopsis
: the
Arabidopsis
genome encod...
Figure 15.4 Simplified schematic representation of drought stress defense in...
Figure 15.5 Overview of the role of SA in rice, bean, and tomato in toleranc...
Figure 15.6 Simplified schematic representation of cold stress defense in
Ar
...
Chapter 16
Figure 16.1 Salicylic acid biosynthesis pathway.
Chapter 17
Figure 17.1
A proposed model of SA
‐
driven PTMs in plant cells. (a)
Methyl SA...
Figure 17.2
Methodologies to analyze protein post‐translational modification
...
Guide
Title Page
Copyright Page
List of Contributors
Preface
Table of Contents
Begin Reading
Index
WILEY END USER LICENSE AGREEMENT
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Managing Plant Stress Using Salicylic Acid
Managing Plant Stress Using Salicylic AcidPhysiological and Molecular Aspects
Edited by
Anket Sharma
Zhejiang A&F University, Hangzhou, China
Renu Bhardwaj
Guru Nanak Dev University, Punjab, India
Vinod Kumar
Government Degree College, Ramban, J&K, India
Bingsong Zheng
Zhejiang Agriculture and Forestry University, Hangzhou, China
Durgesh Kumar Tripathi
Amity University, Uttar Pradesh, Noida, India
This edition first published 2023© 2023 John Wiley & Sons Ltd
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The right of Anket Sharma, Renu Bhardwaj, Vinod Kumar, Bingsong Zheng, and Durgesh K. Tripathi to be identified as the authors of the editorial material in this work has been asserted in accordance with law.
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List of Contributors
Muhammad AamerResearch Centre on Ecological Sciences, Jiangxi Agricultural University, Nanchang, China
Mahsa AbbasiDepartment of Horticultural Science, Gorgan University of Agricultural Sciences and Natural Resources, Gorgan, Iran
Aniqa AfzalDepartment of Plant Pathology, University of Agriculture, Faisalabad, Pakistan
Hassan Ahmed Ibraheem AhmedTasmanian Institute of Agriculture, University of Tasmania, Hobart, Australia
Hanan Sayed AliBotany and Microbiology Department, Faculty of Science, Damietta University, Damietta, Egypt
Imran AshrafDepartment of Agronomy, University of Agriculture Faisalabad, Faisalabad, Pakistan
Leila BensidhoumLaboratoire de Maitrise des Energies Renouvelables, Faculté des Sciences de la Nature et de la Vie, Université de Béjaia, Bejaia, Algérie
Renu BhardwajDepartment of Botanical and Environmental Sciences, Guru Nanak Dev University, Amritsar, Punjab, India
Tamanna BhardwajDepartment of Botanical and Environmental Sciences, Guru Nanak Dev University, Amritsar, Punjab, India
Maria BatoolCollege of Plant Science and Technology, Huazhong Agricultural University, Wuhan, China
Faiçal BriniBiotechnology and Plant Improvement Laboratory, Center of Biotechnology of Sfax, (CBS)‐University of Sfax, Sfax, Tunisia
Muhammad Umer ChatthaDepartment of Agronomy, University of Agriculture, Faisalabad, Pakistan
Sardar Alam CheemaDepartment of Agronomy, University of Agriculture Faisalabad, Faisalabad, Pakistan
Nasser DelangizDepartment of Plant Biotechnology and Breeding, Faculty of Agriculture, University of Tabriz, Tabriz, Iran
Shalini DhimanDepartment of Botanical and Environmental Sciences, Guru Nanak Dev University, Amritsar, India
Heba Talat EbeedBotany and Microbiology Department, Faculty of Science, Damietta University, Damietta, Egypt
Nahla A. El‐SherifBotany Department, Faculty of Science, Ain Shams University, Cairo, Egypt; Biology Department, Taibah University, Madinah, Saudi Arabia
Ivette García‐SotoCentro de Ciencias Genómicas, Universidad Nacional Autónoma de México, Cuernavaca, México
Mouna GhorbelBiotechnology and Plant Improvement Laboratory, Center of Biotechnology of Sfax, (CBS)‐University of Sfax, Sfax, Tunisiaand University of Hai’l, Hail city, Kingdom of Saudi Arabia
Huang GuoqinResearch Centre on Ecological Sciences, Jiangxi Agricultural University, Nanchang, China
Fasih Ullah HaiderCollege of Resource and EnvironmentalSciences, Gansu Agricultural University, Lanzhou, ChinaandDepartment of Agronomy, University of Agriculture Faisalabad, Faisalabad, Pakistan
Tang HaiyingResearch Centre on Ecological Sciences, Jiangxi Agricultural University, Nanchang, China
Muhammad Umair HassanResearch Centre on Ecological Sciences, Jiangxi Agricultural University, Nanchang, ChinaandDepartment of Agronomy, University of Agriculture, Faisalabad, Pakistan
Dhriti KapoorDepartment of Botany, School of Bioengineering and Biosciences, Lovely Profesional University, Phagwara, India
Rupinder KaurDepartment of Biotechnology, DAV College, Amritsar, Punjab, India
Kanika KhannaDepartment of Botanical and Environmental Sciences, Guru Nanak Dev University, Amritsar, Punjab, India
Sukhmeen Kaur KohliDepartment of Botanical and Environmental Sciences, Guru Nanak Dev University, Amritsar, India
Jaspreet KourDepartment of Botanical and Environmental Sciences, Guru Nanak Dev University, Amritsar, Punjab, India
Behnam Asgari LajayerDepartment of Soil Science, Faculty of Agriculture, University of Tabriz, Tabriz, Iran
Dong Jin LeeDepartment of Crop Science & Biotechnology, Dankook University, Cheonan‐si, Republic of Korea
Israel Maruri‐LópezCentro de Ciencias Genómicas, Universidad Nacional Autónoma de México, Cuernavaca, Morelos, México
Sina Siavash MoghaddamDepartment of Plant Production and Genetics, Faculty of Agriculture, Urmia University, Urmia, Iran
Ebrahim MoghisehNuclear Agriculture Research School, Nuclear Science and Technology Research Institute, Karaj, Iran
Slimane MokraniLaboratoire de Maitrise des Energies Renouvelables, Faculté des Sciences de la Nature et de la Vie, Bejaia, AlgérieandLaboratory of Research on Biological Systems and Geomantics, Faculty of Nature and Life Sciences, Mascara, Algeria
El‐hafid NabtiLaboratoire de Maitrise des Energies Renouvelables, Faculté des Sciences de la Nature et de la Vie, Université de Bejaia, Bejaia, Algérie
Faisal NadeemDepartment of Agronomy, The University of Haripur, Haripur, Pakistan
Latifeh PourakbarDepartment of Biology, Faculty of Science, Urmia University, Urmia, Iran
Muthusamy RamakrishnanState Key Laboratory of Subtropical Silviculture, Zhejiang A&F University, Hangzhou, Zhejiang, China
Adnan RasheedKey Laboratory of Crops Physiology, Ecology and Genetic Breeding, Ministry of Education/College of Agronomy, Jiangxi Agricultural University, Nanchang, China
Abdul RehmanDepartment of Crop Science & Biotechnology, Dankook University, Cheonan‐si, Republic of Korea
Mario SerranoCentro de Ciencias Genómicas, Universidad Nacional Autónoma de México, Cuernavaca, Morelos, México
Babar ShahzadTasmanian Institute of Agriculture, University of Tasmania, Hobart, Australia
Ashutosh SharmaFaculty of Agricultural Sciences, DAV University, Jalandhar, Punjab, India
Neerja SharmaDepartment of Botanical and Environmental Sciences, Guru Nanak Dev University, Amritsar, Punjab, India
Neha SharmaDepartment of Botanical and Environmental Sciences, Guru Nanak Dev University, Amritsar, India
Pooja SharmaDepartment of Botanical and Environmental Sciences, Guru Nanak Dev University, Amritsar, Punjab, IndiaandDepartment of Microbiology, DAV University, Jalandhar, Punjab, India
Vasudha SharmaDepartment of Soil, Water, and Climate, University of Minnesota, Saint Paul, USA
Vivek SharmaAgricultural and Biological Engineering Department, University of Florida, Gainesville, USA
Arun Dev SinghDepartment of Botanical and Environmental Sciences, Guru Nanak Dev University, Amritsar, Punjab, India
Geetika SrihindiDepartment of Botany, Punjabi University, Patiala, Punjab, India
Aman UllahDepartment of Plant Sciences, College of Agricultural and Marine Sciences Sultan Qaboos University, Al‐Khoud, Oman
Mingbing ZhouState Key Laboratory of Subtropical Silviculture, Zhejiang A&F University, Hangzhou, China
Preface
Plants, during their life, encounter various challenges, and one of the main threats for their normal development is abiotic stress. The major abiotic stresses possessing threat to plants are water scarcity, salinity, extreme temperatures, heavy metals, and pesticides. All these abiotic factors can cause phytotoxicity, either directly or indirectly, resulting in interruption to plant’s growth and development which eventually decrease plant yield. Researchers all over the world have already specified abiotic stresses as the main danger for agrarian systems. However, plant’s internal defense system tries to counterattack the negative impacts of abiotic stresses by regulating their biological processes. But, above a threshold level of a particular stress, even plant’s internal antioxidative defense system is unable to entirely protect plants from the deleterious effects of abiotic stress. So, plants need some external stimulus or support to boost the level of defense system to survive under challenging environments. Exogenous application of plant hormones is one of the best and eco‐friendly approaches to trigger the defense system of stressed plants.
In the recent past, a lot of investigations have been focused on studying the mechanisms of plant hormone‐mediated regulation of plant growth and development under abiotic stress conditions. Salicylic acid is an important plant hormone which acts a multifunctional molecule and regulates key physiological and biochemical processes in plants. This hormone also provides resistance to plants against abiotic stresses by regulating key cell signaling pathways. Exogenous application of SA helps in convalescing the growth and development of stressed plants by reducing the oxidative stress accompanied by enhanced performance of antioxidative defense machinery. At present, scientists all over the world are very keen to study the deep mechanisms of SA‐modulated abiotic stress responses by using various advanced molecular techniques. These advancements in research approach can be beneficial in exploring some important genetic pathways which can be applied to develop abiotic stress‐tolerant plant varieties. So, recently, many studies have been carried out to find the deep molecular mechanisms explaining SA‐mediated regulation of plant growth under abiotic stress. So, our purpose is to compile all the latest developments described in the arena of SA‐mediated regulation of abiotic stress.
The first chapter explains the general roles of salicylic acid in plant biology. Chapter 2 discusses the role of salicylic acid in plants during stressful conditions in relation to omics approaches. Chapter 3 focuses on describing the possible role of salicylic acid in regulation of primary metabolisms like respiration and photosynthesis in plants growing under challenging conditions. The next chapter discusses salicylic acid‐mediated secondary metabolism in plants under abiotic stress. Further chapters explore the role of salicylic acid in stressed plants by important aspects like mineral nutrition, seed germination to fruit maturation. Chapter 7 brings the updated knowledge about the role of salicylic acid in the postharvest technology. The next chapters focus on exploring salicylic acid‐mediated physiological and molecular mechanism in plants under stresses like metal(loid), heat, chilling, and drought. Chapter 14 describes in detail the regulation of photosynthesis by salicylic acid under optimal and suboptimal conditions. Further chapters focus on describing the roles of salicylic acid in mediating stress conditions in plants at genetic levels including the phytohormonal cross talk and post‐transnational modifications.
This book is a collection of recent developments in the field of salicylic acid biology in relation to challenging environment conditions. To the academic and industry sectors, the book provides useful hints for the development of eco‐friendly stress‐mediating approaches as well as helps to understand the future importance and involvement of salicylic acid in safe food production. Therefore, we believe that this book will be a vital source of information for scientists and academics working in the field of abiotic stress tolerance.
Dr. Anket SharmaDr. Renu BhardwajDr. Vinod KumarProf. Bingsong ZhengDr. Durgesh Kumar Tripathi
1Salicylic Acid: A Regulator of Plant Growth and Development
Neha Sharma1, Vivek Sharma2, Vasudha Sharma3, and Renu Bhardwaj1
1 Department of Botanical and Environmental Sciences, Guru Nanak Dev University, Amritsar, India
2 Agricultural and Biological Engineering Department, University of Florida, Gainesville, FL, USA
3 Department of Soil, Water, and Climate, University of Minnesota, Saint Paul, MN, USA
Introduction
In plants, the phytohormones act as endogenous signals, both spatially and temporally, regulating a number of physiological functions. The cross talk between various phytohormones helps the plant to withstand biotic and abiotic stresses. This cross talk of plant hormones has evolved into a complex network within the plants, thus helping the plants having a balanced reaction to developmental and environmental stimuli (Sharma et al. 2018, 2019a; Koo et al. 2020). Salicylic acid (SA) or ortho‐hydroxybenzoic acid is a member of the group of plant phenolics with a seven‐carbon (C) skeleton. A study of reproductive structures and leaves of 34 plant species confirmed that SA is ubiquitously distributed in plant kingdom (Raskin et al. 1990). The name SA is from Salix (Latin word) as it was found to be an active constituent of willow tree bark (Salix sp.) which was used extensively to cure fever and aches (Khan et al. 2015).
The biosynthesis of SA in plants involves the isochorismate synthase (ICS) pathway and phenylalanine ammonia‐lyase (PAL) pathway (Janda et al. 2014). The ICS pathway was first discovered in Pseudomonas species and the PmsCEAB gene cluster was found to play the key role in the synthesis of SA. The conversion of chorismate to isochorismate (IC) is catalyzed by PmsC gene and then isochorismate pyruvatelyase encoded by the PmsB gene converts IC to SA making SA synthesis from chorismate a two‐step process (Mercado‐Blanco et al. 2001; Lefevere et al. 2020). In the PAL pathway, the key enzyme is chorismate mutase (CM) which catalyzes the process of converting CM to prephenate. Prephenate gets converted to phenylalanine (Phe), which in turn is converted to trans‐cinnamic acid (tCA) by the enzyme PAL. The next step involves the catalyzing of the conversion of tCA to benzoic acid (BA) by abnormal inflorescence meristem1 (AIM1), which is a multifunctional protein (MFP) family member (Rylott et al. 2006; Arent et al. 2010). The last step in the PAL pathway is the conversion of BA to SA which is presumed to be catalyzed by benzoic acid hydroxylase (Lefevere et al. 2020).
The ICS as well as PAL pathways to synthesize SA start from chorismate, and the importance of both ICS and PAL varies in different species of plants, as not all enzymes which catalyze various reactions in these pathways have been found in all plants. The ICS pathway plays an important role in SA biosynthesis in Arabidopsis, and PAL has been found to be more important in rice, while in soybeans, both pathways contribute equally (Silverman et al. 1995; Duan et al. 2014).
In plants, SA plays a significant part in the growth, development, and in the protection from biotic and abiotic stresses (Khan et al. 2015; Sharma et al. 2019b, 2020; Prakash et al. 2021) (Figure 1.1). The role of SA in defense mechanisms of plant was established during the last 30 years and before that it was recognized as an unimportant secondary plant metabolite. Since 1979, when White (1979) reported the role of SA in tobacco plants’ disease resistance, numerous findings showed the role of SA as an important regulatory substance in plants (Chen et al. 2009). Studies have shown that in plants, SA plays a vital part in disease resistance, DNA damage/repair, seed germination, fruit yield, and thermogenesis (Dempsey and Klessig 2017). Increased levels of SA are seen in the presence of an infection, and if supplied exogenously, SA strengthens the plant defense system (Lefevere et al. 2020). In this review, we have focused on the role of SA in plants as a regulator of growth and development and providing resistance against various stresses.
Figure 1.1 Schematization of the role of salicylic acid in plants.
Source: Based on Khan et al. 2015; Sharma et al. 2019b, Sharma et al. 2020; Prakash et al. 2021.
Salicylic Acid and Plant Growth
SA plays an important role in plant growth, along with other phytohormones, and its effects on growth, when applied exogenously, is affected by the species of the plant and its stage of development as well as its concentration (Vicente and Plasencia 2011). It has been reported that more than 1 mM of SA is considered a high concentration and has negative effects (Koo et al. 2020). Barley and maize seeds did not show any germination when imbibed in >3 mM of SA (Guan and Scandalios 1995; Xie et al. 2007). On the contrary, when maize seeds were imbibed in ~0.3 mM– ~0.9 mM of SA, an increased germination speed and enhanced shoot length were recorded (Sallam and Ibrahim 2015). SA (aqueous solution), when applied to soybean shoots in the form of spray, increased the shoot and root growth significantly. Although, photosynthetic rate was not found to have any significant effect with this treatment (Gutiérrez‐Coronado et al. 1998). In soybean, wheat, maize, and chamomile, SA has been found to stimulate growth. An increased growth of ~20 and 45% in the shoots and roots, respectively, was observed in soybean plants when treated with 10 nM, 100 μM, and up to 10 mM of SA. In wheat seedlings, development of larger ears and enhancement of cell division was observed in the shoot apical meristems, with a treatment of 50 μM SA. In chamomile plants, 50 μM SA stimulated the growth and an opposite effect was observed at a concentration of 250 μM SA (Gutiérrez‐Coronado et al. 1998; Shakirova et al. 2003; Gunes et al. 2007; Kováčik et al. 2009). In apple, strawberry, and mango plants, fruit setting was enhanced with SA treatment (Shaaban et al. 2011; Kazemi 2013; Ngullie et al. 2014).
The relationship between SA, reactive oxygen species (ROS), and mitogen‐activated protein kinase (MAPK) cascades has been found to be very important in regulating plant growth. Zhang and Klessig (1997) found that Arabidopsis MPK6 is an orthologue of tobacco SA‐induced protein kinase (SIPK), and has been suggested to have an important role in growth and development (Bush and Krysan 2007; Wang et al. 2007, 2008). It has been reported that in regulation of cell growth, MAPK cascades act as mediators between phytohormones, SA, and ROS signaling (Foreman et al. 2003; Potocký et al. 2007).
SA also plays an important role in regulating flowering. Lee and Skoog (1965) indicated its flower‐inducing effects for the first time. 4 μM SA was reported to promote flower bud formation from callus of tobacco. Then, Cleland and Ajami (1974) reported the isolation and identification of SA in aphid honeydew as the substantial factor for flower induction in short‐day plant Xanthium strumarum. Later, SA’s role in Impatiens balsamina, Oncidium (orchid species), Pisita stratiotes L., and Arabidopsis thaliana as a stimulatory factor on flowering was demonstrated. In thermogenic plants, the inflorescences were found to have high levels of endogenous SA (Raskin et al. 1990), while in non‐thermogenic plants, SA levels were found to increase twofold in tobacco and fivefold in Arabidopsis leaves at the time of initiation or transition toward flowering (Yalpani et al. 1993; Abreu and Munné‐Bosch 2009). Similarly, Arabidopsis plants, which were SA deficient (NahG, sid1/eds5, and sid2) exhibited a phenotype having late flowering (Martínez et al. 2004). In sunflower, it was discovered that the transcription factor HAHB10 (belonging to HD‐Zip II family) plays a role in responding to biotic stress and inducing flowering and it was observed that treatment with SA induces the HAHB10 expression (Dezar et al. 2011).
However, the possibility of endogenous regulation by SA alone in case of flowering was weakened as there was not much difference in the levels of SA in aphid honeydew from flowering as well as vegetative plant parts. Thus, it was established that SA promotes and regulates flowering in conjugation with other plant growth regulators (Raskin 1992).
Salicylic Acid and Photosynthesis
In plants, photosynthesis is considered to be a very delicate physiological process. Heat stress can harm the photosynthetic apparatus, and plants have developed protective mechanisms like dissipating the excess excitation energy, utilizing heat shock proteins and plant growth regulators. SA has been reported to play a significant part during abiotic stresses to plants (Wang et al. 2010; Kohli et al. 2017, 2018). SA application enhanced the photosynthetic capacity in barley and spring wheat plants which were exposed to salt and drought stress (El‐Tayeb 2005; Arfan et al. 2007). In tobacco and Arabidopsis, when SA was applied exogenously, it improved the heat tolerance (Dat et al. 1998a, 1998b; Lopez‐Delgado et al. 1998; Senaratna et al. 2003; Clarke et al. 2004). In heat‐stressed young grape leaves, the thiobarbituric acid reactive substances were decreased upon spraying with SA (0.1 mM solution) which indicates that SA helped in tolerance of intrinsic heat stress (Wang and Li 2006).
In a study conducted by Fariduddin et al. (2003), foliar application of SA (aqueous solution) to 60 days old Brassioca juncea plants showed higher net photosynthetic rate and increased pod count and seed yield as compared with the control. Studies have suggested that SA plays a significant role in photosynthesis as it affects the structure of chloroplast and leaf (Uzunova and Popova 2000), contents of carotenoid and chlorophyll, and closure of stomata. It has also been reported to affect the carbonic anhydrase as well as RuBisCO (ribulose‐1,5‐bisphosphate carboxylase/oxygenase) enzymes (Vicente and Plasencia 2011). Haroun et al. (1998) reported that SA application promotes photosynthetic pigments in the leaves of lupine (Lupinus termis Forssk.). Further, a study conducted on Egyptian Lupine confirmed that foliar application of SA (25, 50, and 75 ppm) resulted in a rise in chlorophyll a,b and carotenoids in leaves (Gomaa et al. 2015).
The closing of stomata in plants is controlled by different phytohormones, and is considered to undertake a vital role in photosynthesis (Acharya and Assmann 2009). Studies have shown a significant role of SA in the functioning of guard cells, which is in turn essential for the closure of stomata (Melotto et al. 2006). Mateo et al. (2004) reported that stomatal gas exchange was reduced fourfold within two hours after treating Arabidopsis with 0.4 mM SA.
Salicylic Acid and Respiration
SA plays a significant role in regulating the alternative oxidase pathway (AOX) via induction of its gene expression (Kapulnik et al. 1992; Rhoads and McIntosh 1992). AOX, for maintaining the growth rate homeostasis, leads a controlled synthesis of ATP as non‐proton‐driven carrier (Moore et al. 2002). AOX is further reported to decrease the production of ROS in mitochondria. Thus, SA targeting AOX plays a significant role for regulation of plant growth. Studies have also emphasized on the role of SA in controlling the oxidative phosphorylation and electron transport in plant mitochondria (Xie and Chen 1999; Norman et al. 2004).
Adding 2–20 μM SA in a cell suspension culture of tobacco resulted in increased cyanide‐resistant O2 uptake. Calorimetric measurements showed an increase of 60% in the rate of evolution of heat from the cells (Kapulnik et al. 1992). SA has been reported to exert a dual action on mitochondrial respiration, depending on concentration, in plant and animal mitochondria. At a lower concentration (0.1–1.0 mM), on the oxidative phosphorylation, it exerts mainly an uncoupling reaction, helping in increasing the substrate oxidation rate. At a high concentration (more than 1 mM), SA significantly suppressed State 3 respiration (Howell et al. 2004; Nulton‐Persson et al. 2004; de Souza et al. 2011). In a study conducted in the mitochondria of sugarbeet (Beta vulgaris L.), taproots, and lupine (Lupinus luteus L.) cotyledons, to understand the effect of SA on generation of membrane potential (Δψ) at malate oxidation, it was observed that the Δψ dissipation was a result of the uncoupling and inhibitory action of SA on respiration. It was concluded that in a plant cell under stress, increased concentration of SA has a direct and strong effect on the basic mitochondrial functions, viz., ATP synthesis, oxidation of respiratory substrates, ROS production, and membrane potential generation and the degree of the extent of these effects not only depends on the amount of time the organelle was exposed to SA but also on its concentration and the mitochondrial structure and function in different plants and animals (Shugaev et al. 2014).
Salicylic Acid and Abiotic and Biotic Stress
Agricultural productivity all over the world is affected by abiotic and biotic stresses to the plants (Handa et al. 2018, 2019; Wang et al. 2019; Arif et al. 2019; Roychoudhury and Tripathi 2019; Kapoor et al. 2019, 2020; Deshmukh et al. 2020; Roychoudhury and Tripathi 2020; Dhiman et al. 2021). Plant growth regulators are well known to be involved in the plant‐responses to stress (Yadav et al. 2018; Singh et al. 2020). SA has been found to induce stress resistance in plants having biotic stress (Kumar 2014) and also to increase tolerance to abiotic stresses such as heat, salinity, drought, and metal (Khan and Khan 2013; Fayez and Bazaid 2014; Khan et al. 2014; Zhang et al. 2015; Prakash et al. 2021). An application of SA exogenously to Oryza sativa, Zea mays, and Phaseolus vulgaris plants exposed to lead, cadmium, and copper metal stresses were found to have improved photosynthetic traits as well as enhanced growth. It also modulated the antioxidant defense system of the plants (Krantev et al. 2008; Zengin 2014; Arif et al. 2016). The exogenous application of SA increased endogenous SA content by enhancing OsWRKY45 gene expression, which in turn lowered the H2O2 content and thus prevented membrane damage in Oryza sativa plants exposed to cadmium metal stress (Chao et al. 2010). Similarly, Belkadhi et al. (2015) also reported that SA regulated the H2O2 accumulation and thus resulted in an enhanced tolerance to Cd in Linum usitatissimum. Chlorosis caused by Fe deficiency has also been reported to be inhibited by SA (Kong et al. 2014).
In crops like Vicia faba, Brassica juncea, Medicago sativa, and Vicia radiata, SA has been found to strengthen the stress mechanism against salinity stress (Azooz 2009; Nazar et al. 2011, 2015; Palma et al. 2013; Khan et al. 2014). In case of Triticum aestivum, exogenous application of SA (0.5 mM) resulted in an improvement of salt stress tolerance via increase in the enzyme activity of ascorbate‐GSH pathway and enhancement in the level of GPX1, GPX2, DHAR, GR, GST1, GST2, MDHAR, and GS antioxidant genes (Li et al. 2013). Similarly, in case of Ozone stress to plants, SA promotes molecular and physiological changes as it works as a signal molecule (Tamaoki 2008). It has also been suggested that SA plays a role in the signaling network which integrates phytohormones like JA, ethylene in ozone‐stressed plants (Rao et al. 2002). The modulation of antioxidative enzymes, detoxification of superoxide radicals, prevention of oxidative damage, and protection of metabolic enzymes and membranes have been reported following exogenous SA application in plants which are UV‐B radiation stressed (Mohammed and Tarpley 2009). A decrease in chromosome aberration level caused by UV‐B in the meristematic root tip cells of Crepis capillaris was reported with the application of SA (Rančelienė and Vyšniauskienė 2012). Li et al. (2014) reported significant improvement in photosynthetic function, when SA was applied exogenously to plants exposed to UV‐B.
There are studies on the significant part of SA in plants having temperature stress. Modulation of antioxidant enzymes in temperature‐stressed (2 °C) Z. mays has been reported by Janda et al. (1999). Under chilling stress, SA was found to protect ultrastructures in the seedlings of Musa acuminata (Kang et al. 2007). In Hordeum vulgare genotypes, exogenous application of SA enhanced antioxidant enzyme and ice nucleation activity and thus provided cold tolerance (Mutlu et al. 2013). Similarly, in lemon fruit, SA increased total phenolics synthesis and also improved tolerance to chilling temperature (Siboza et al. 2014). In case of drought stress, a number of studies have reported the stress mitigation role of SA application. Habibi (2012) reported an increase in the rate of net CO2 assimilation and plant dry mass when supplemented with 500 μM SA to drought‐stressed H. vulgare plants. A foliar application (1.0 μM of SA) was found to enhance the antioxidative defense system in Z. mays plants (draught‐tolerant vs. drought‐sensitive cultivar) exposed to drought stress (Saruhan et al. 2012). It has been reported that oHCA was associated with the biosynthesis of SA and thus played an important part in providing drought tolerance to O. sativa plants (Pál et al. 2014).
SA has also been found as a key element in providing the plants the resistance to various microbial pathogens like fungi, bacteria, viruses, and oomycetes (Kunkel and Brooks 2002; Vlot et al. 2009). Glazebrook (2005) established a positive correlation between resistance responses of plants against hemibiotrophic and biotrophic pathogens and the levels of endogenous SA. In case of Citrus sinensis plants, a 45% reduction in disease caused by Xanthomonas axonopodis was reported (Wang and Liu 2012). The resistance of tomato and Arabidopsis plants was enhanced against Botrytis cinerea with the application of SA (Ferrari et al. 2003; Li and Zou 2017). Similarly, treating tomato plants with SA has also been reported to provide resistance against Fusarium oxysporum and Alternaria alternata (Esmailzadeh et al. 2008; Jendoubi et al. 2015). A retarded nymph development was observed in case of Oebalus pugnax, which is a piercing and sucking insect, when SA was applied exogenously to O. sativa plants (Stella de Freitas et al. 2019).
The exogenous treatment of plants subjected to pathogen stress with SA results in enhanced expression of pathogenesis‐related genes (PR1, PR2, and PR5) and this overexpression of genes increases tolerance to not only pathogens but also various other abiotic stresses (Hong and Hwang 2005; Sarowar et al. 2005; Ali et al. 2018). The mechanism of SA‐induced tolerance to plants against stress can include various steps like osmolyte accumulation (glycinebetaine, proline, soluble sugars, and amines), thus maintaining osmotic homeostatis, enhancing scavenging of ROS, production of stress proteins, regulation of transpiration, mineral uptake regulation, inhibition of ethylene synthesis, and increase in the production of secondary plant metabolites (Horváth et al. 2007; Khan et al. 2015; Emamverdian et al. 2020).
Conclusions
Plants’ growth and regulation requires a regular cross talk between various plant growth regulators, among which SA has been found to play a vital role. Seed germination, plant physiological functions, flowering, and tolerance against abiotic and biotic stresses are the various aspects of plant growth and regulation, where the role of SA has been found and elaborated in various research studies. Since its discovery as a healing substance obtained from the bark of willow tree, SA has been proved to be valuable not only for humans but also for plants. There are certain reports which suggest that application of SA over a certain amount brings negative effects. Thus, studies involving exogenous application of SA in an optimal amount for positive plant growth and negative growth of unwanted plants in crop production can open new aspects. Further, manipulating levels of endogenous SA in transgenic plants can open new areas in crop biotechnology and thus help in growing crops with positive growth and increased stress tolerance.
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