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Chapter 5 Impact of drought stress on the medicinal and aromatic plants’ biochemistry 223
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nanoparticles and
2
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Fatemeh Ahmadi
Chapter 6 Impact of salinity stress on medicinal and aromatic plant biotechnology
Abstract: Salinity stress is a critical environmental challenge affecting the growth, de-
velopment, and metabolic processes of medicinal and aromatic plants (MAPs). These plants are highly valued for their secondary metabolites, which have extensive appli cations in medicine, cosmetics, and the food industry. Salinity disrupts key physiologi­cal functions, including photosynthesis, water relations, and nutrient balance, while inducing oxidative stress and altering metabolite production. Despite these chal lenges, MAPs exhibit remarkable physiological, biochemical, and molecular adaptive mechanisms, such as osmotic adjustment, ion homeostasis, antioxidant defenses, and regulation of stress-responsive genes. This chapter provides a comprehensive analysis of the effects of salinity on MAPs, highlighting changes in growth metrics, photosyn thesis, and secondary metabolite production. It delves into molecular mechanisms that enable salt tolerance, such as the SOS pathway, ion transporters, and transcrip tion factors. Furthermore, the chapter explores strategies to enhance salt resilience in MAPs, including the use of plant growth regulators, beneficial microorganisms, ge netic engineering, and agronomic practices like mulching and silicon supplementa­tion. The integration of advanced biotechnological tools, such as CRISPR/Cas9 and omics approaches, is also discussed to optimize salt tolerance and metabolite produc tion. By leveraging these insights, sustainable solutions for MAP cultivation in saline environments can be achieved, ensuring the continued economic and therapeutic sig nificance of these plants.
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Keywords: antioxidant defenses, CRISPR/Cas9, genetic engineering, plant growth regu­lators, salt tolerance mechanisms

6.1 Introduction

High soil salt content presents a formidable ecological hurdle, severely impacting crop development and output [ centrations in the plant’s environment, interfering with vital physiological processes. Originally confined to dry regions, this problem has now expanded globally, driven by both natural events and human activities [
Corresponding author: Fatemeh Ahmadi, School of Agriculture and Environment, University of
Western Australia, Crawley, WA 6009, Australia, e-mail: fatemeh.ahmadi@uwa.edu.au
1]. This phenomenon results from excessive salt con-
2, 3]. Researchers gauge soil salinity by
230 Fatemeh Ahmadi
measuring the soil solution’s electrical conductivity (EC) [4]. The USDA Salinity Labo­ratory considers soil to be saline when its saturated paste extract (EC
) exceeds 4 dS/
e
m, though many plants react negatively to lower levels, with field conditions poten­tially exacerbating these effects [5]. Salt-induced stress manifests through osmotic and ionic mechanisms. The former restricts water uptake, creating drought-like con­ditions even in wet soils [6]. The latter involves cellular damage from ion overload, particularly Na
+
and Cl–, disrupting plant metabolic functions. Salinity types include dryland and irrigation-induced, based on salt accumulation methods. Dryland salin­ity occurs in nonirrigated areas due to natural phenomena like groundwater move­ments [7]. Irrigation salinity results from repeated use of salt-rich water, leading to salt buildup [8]. Furthermore, salinity is classified as primary or secondary based on its source. Primary salinity evolves naturally through geological processes, while secondary salinity results from human interventions such as land clearing and poor irrigation management [9, 10].
Salt-affected soils are classified based on their electrical conductivity, sodium con­tent, and pH levels, as three primary types: saline, sodic, and saline-sodic soils [11, 12]. These soil conditions significantly challenge plant survival by creating complex physi­ological stress mechanisms. Plants respond to salinity through sophisticated adaptive strategies at molecular and cellular levels [13]. They develop multiple defense mecha­nisms to counteract salt stress, including accumulating protective osmolytes, regulat­ing ion homeostasis, enhancing antioxidant defenses, modifying gene expression, and producing stress-responsive hormones like abscisic acid (ABA) [14, 15]. The primary survival strategies involve maintaining water balance, preventing toxic ion accumula­tion, and protecting cellular structures from oxidative damage. These adaptive re­sponses enable plants to survive and potentially thrive in challenging saline environ­ments [16]. Understanding these intricate plant responses provides crucial insights for developing salt-tolerant crop varieties, ultimately supporting agricultural productivity in regions with challenging soil conditions [17, 18].

6.2 Importance of medicinal and aromatic plants

For centuries, medicinal and aromatic plants (MAPs) have been closely linked to human health and cultural heritage. These plants, celebrated for their diverse bioac­tive compounds, have served as the cornerstone of traditional medicine and continue to hold significant importance in modern healthcare, pharmaceutical advancements, and a wide range of industrial applications [19]. The significance of MAPs goes beyond their medicinal properties, encompassing economic, ecological, and cultural impor­tance. The use of plants for healing predates written history, with early civilizations, including those in Egypt, China, India, and Greece, developing detailed herbal medi­cine systems. For instance, the Egyptian Ebers Papyrus, a document from 1550 BCE,
Chapter 6 Impact of salinity stress on medicinal and aromatic plant biotechnology 231
describes over 850 plant-based remedies. Similarly, traditional Chinese medicine and Ayurveda have harnessed MAPs for thousands of years, creating complex pharmaco­peia and therapeutic approaches [20].
MAPs form a vital component of global biodiversity, with more than 50,000 me­dicinal species among the estimated 422,000 flowering plants. This biodiversity is es­sential for ecological stability and represents a vast resource for discovering new medicines. However, growing demand for MAPs has resulted in overharvesting, ne­cessitating robust conservation measures [21]. Conservation strategies include in situ preservation of natural habitats, ex situ conservation in botanical gardens and gene banks, adopting sustainable harvesting techniques, and promoting cultivation to alle­viate pressure on wild populations [22].
The therapeutic value of MAPs arises from their phytochemical composition, which includes alkaloids (e.g., morphine from Papaver somniferum), glycosides (e.g., digoxin from Digitalis lanata), terpenoids (e.g., artemisinin from Artemisia annua), phenolics (e.g., curcumin from Curcuma longa), and flavonoids (e.g., quercetin from various sources) [23]. These compounds often exhibit synergistic effects, enhancing their medicinal efficacy. For example, garlic (Allium sativum) derives its therapeutic properties from organosulfur compounds like allicin, known for its antibacterial, anti­fungal, and antiviral properties [24].
MAPs are extensively used in managing diverse health conditions. Examples include Ginkgo biloba and garlic for cardiovascular health, paclitaxel from Pacific yew and vin­cristine from Madagascar periwinkle for cancer treatment, bitter melon and fenugreek for diabetes, Echinacea species for respiratory ailments, and St. John’s wort for mental health concerns. Additionally, MAPs are a cornerstone of drug discovery, contributing directly or indirectly to about 25% of modern medicines. The process involves studying traditional applications, isolating bioactive components, determining their chemical structures, and conducting preclinical and clinical trials. Notable drugs derived from MAPs include aspirin (from Salix species), morphine, quinine (from Cinchona species), and artemisinin [25].
Aromatic plants, valued for their ability to produce essential oils, find extensive applications across various fields, including medicine, perfumery, cosmetics, and aro­matherapy [26]. These essential oils are renowned for their wide-ranging biological activities, such as the potent antimicrobial properties of tea tree oil (Melaleuca alterni-
folia), the anti-inflammatory effects associated with lavender oil (Lavandula angustifo­lia), and the calming, anxiolytic benefits evidenced in research on lavender oil inhala-
tion [27]. The global essential oil market, which was valued at USD 7.03 billion in 2020, is projected to experience consistent growth due to increasing demand across indus­tries. Similarly, the herbal medicine market, initially valued at USD 83 billion in 2019, is expected to witness exponential growth, with predictions estimating its value at USD 550 billion by 2030 [28]. These markets underscore the economic importance of MAPs, which support agriculture, industrial processing, international trade, and em­ployment, especially in developing nations [29, 30].
232 Fatemeh Ahmadi
Despite their immense potential, MAPs face challenges such as maintaining con­sistent quality, standardizing cultivation and processing, navigating diverse regula­tory frameworks, addressing conservation needs alongside growing demand, and safeguarding intellectual property rights related to traditional knowledge [31]. Future directions for research include refining phytochemical analysis with advanced tech­nologies, employing omics approaches like genomics and metabolomics, utilizing bio­technological methods to produce plant-derived compounds, validating traditional uses through rigorous clinical trials, and developing sustainable cultivation and pro­duction practices [32].
The contributions of MAPs to healthcare, the economy, and cultural heritage remain indispensable. Their role in modern drug discovery, the growing demand for natural products, and their potential to address global health issues reinforce their significance [33]. As research progresses, integrating traditional knowledge with modern science will be crucial to ensuring their sustainable use and conservation for future generations [34]. The multidisciplinary importance of MAPs – from traditional medicine and global trade to biodiversity and cultural preservation – requires a holistic approach to their study and sustainable utilization. Advancing our understanding through collaborative efforts promises to unlock even greater potential from these extraordinary plants [35].

6.3 Salinity effect on medicinal plants

Salinity stress severely impacts MAPs, affecting their growth, photosynthesis, and me­tabolite production [36, 37]. These plants develop complex adaptive mechanisms to survive in saline environments. Studying these responses offers valuable insights for enhancing plant resilience and developing salt-tolerant cultivation methods, showcas­ing the remarkable adaptability of plants under environmental stress [38].
6.3.1 Effects on growth and development
Salinity stress significantly impairs plant growth, particularly in medicinal species, by disrupting essential physiological processes [39]. High salt levels interfere with chloro­phyll production and photosynthetic efficiency, leading to stunted development that is summarized in Table 6.1. Research on plants like moringa demonstrates how salt exposure can dramatically reduce growth metrics and biomass allocation, underscor­ing the profound impact of salt stress on plant health [37].
Salinity stress also affects plants at the germination stage. Higher salt concentra­tions often delay or entirely inhibit seed germination, which can significantly reduce the establishment of crops and overall yield [40]. Additionally, salt stress interferes with the flowering process in several MAPs. As flowering is often a key determinant