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Файл:Ординатура / Хирургия / Библиотека им академика М.И. Перельмана / Книга_5217_Библиотеки_им_академика_М_И_Перельмана.pdf
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- •Also of interest
- •Contents
- •Part I: Introduction
- •1.1.2.3 Sustainability and future perspectives
- •1.2 Alkaloids, flavonoids, terpenoids, and other active compounds
- •1.2.1 Alkaloids
- •1.2.2 Flavonoids
- •1.2.3 Terpenoids
- •1.2.4 Other active compounds
- •1.3 Chemical structures and pharmacological effects
- •1.3.1 Chemical structures and effects of alkaloids
- •1.3.2 Chemical structures and effects of flavonoids
- •1.3.3 Chemical structures and effects of terpenoids
- •1.3.4 Structures and effects of other compounds
- •1.4.2 Flavonoids
- •1.4.3 Terpenoids
- •1.4.4 Other active compounds
- •1.5 Chemical structures and pharmacological effects
- •1.5.1 Chemical structures and effects of alkaloids
- •1.1 Introduction to medicinal and aromatic plants
- •1.1.1 Historical background
- •1.1.1.1 Historical background
- •1.1.2 Traditional and modern uses
- •1.1.2.1 Traditional uses
- •1.1.2.2 Modern uses
- •1.5.2 Chemical structures and effects of flavonoids
- •1.5.3 Chemical structures and effects of terpenoids
- •1.5.4 Structures and effects of other compounds
- •1.6 Aromatic plants in everyday life
- •1.6.1 The importance of essential oils and aromatherapy
- •1.6.2 Applications in the cosmetics and food industry
- •1.6.3 Food industry
- •1.7.1 Protection of endangered species
- •1.7.2 Sustainable harvesting methods
- •1.8.1 Protection of endangered species
- •1.8.1.1 Threats to endangered species
- •1.8.2 Conservation strategies
- •1.8.2.1 Protection of natural habitats (in situ conservation)
- •1.8.3 Participation of local communities
- •1.8.3.1 Education and awareness
- •1.8.3.2 International collaborations
- •1.8.3.3 Sustainable harvesting and trade
- •1.8.4 Sustainable harvesting methods
- •1.8.4.1 The importance of sustainable harvesting
- •1.8.4.2 Sustainable harvesting principles
- •1.8.4.3 Sustainable harvesting techniques
- •1.8.4.4 Monitoring and evaluating the harvesting process
- •1.8.4.5 The economic dimension of sustainable harvesting
- •1.8.4.6 International approaches and legal regulations
- •1.8.4.6.1 International approaches
- •1.8.4.6.2 Legal regulations
- •1.8.4.6.3 Protection of local communities and traditional knowledge
- •1.8.5 Many countries are protecting biodiversity
- •1.8.5.1 Global conservation efforts
- •1.8.5.2 Protected areas and conservation in natural habitats
- •1.8.5.3 Ex situ conservation and gene banks
- •1.9 Challenges and future prospects
- •1.9.1 Impacts of climate change
- •1.9.2 Genetic and biotechnological approaches
- •1.9.2.1 Protection of genetic diversity and breeding studies
- •1.9.2.2 Genomic and transcriptomic approaches
- •1.9.2.3 Culture tissue techniques
- •1.9.2.4 CRISPR/Cas9 technology
- •1.9.2.5 Metabolic engineering and synthetic biology
- •1.9.2.6 Bioinformatics and data analysis
- •1.10 Case studies and regional practices
- •1.10.1 Successful projects in specific regions
- •1.10.1.1 India: Ayurveda and biodiversity conservation projects
- •1.10.1.2 Brazil: sustainable collection projects in the Amazon forest
- •1.10.1.3 Turkey: protection and production of endemic plants
- •1.10.1.4 Africa: integration of local knowledge with modern practices
- •1.10.2.1 Documentation and protection of traditional knowledge
- •1.10.2.2 Scientific validation and application
- •1.10.2.3 Education and awareness
- •1.10.2.4 Patents and intellectual property rights
- •1.10.2.5 Public and private sector collaboration
- •1.11 Conclusions
- •References
- •2.1 Introduction
- •2.3.1 Plant selection
- •2.3.1.1 Random plant selection
- •2.3.1.2 Plant selection based on ethnopharmacology and traditional uses
- •2.3.1.3 Plant selection by HTS technologies
- •2.3.1.4 Plant selection through virtual screening
- •2.3.1.5 Phytochemical databases
- •2.3.2.1 Comminution and homogenization
- •2.3.3 Extraction
- •2.3.3.1 Conventional extraction techniques
- •2.3.3.2 Maceration
- •2.3.3.3 Infusion
- •2.3.3.4 Decoction
- •2.3.3.5 Percolation
- •2.3.3.13 Pressurized liquid extraction
- •2.3.3.14 Enzyme-assisted extraction
- •2.3.3.15 Solid-phase microextraction
- •2.3.3.6 Hydrodistillation and steam distillation
- •2.3.3.7 Soxhlet extraction
- •2.3.3.8 Advanced extraction techniques
- •2.3.3.9 Ultrasound-assisted extraction
- •2.3.3.10 Pulsed-electric field extraction
- •2.3.3.11 Microwave-assisted extraction
- •2.3.3.12 Supercritical extraction
- •2.3.3.16 Bioassay-guided fractionation of plant extracts
- •2.3.4 Isolation and purification
- •2.3.4.3 Gas chromatography (GC)
- •2.3.4.4 Column chromatography (CC)
- •2.3.4.5 Ion exchange chromatography (IEC)
- •2.3.5 Elucidation of the chemical structure
- •2.3.5.1 Nuclear magnetic resonance (NMR)
- •2.3.5.2 Mass spectrometry (MS) and high-resolution mass spectrometry (HRMS)
- •2.3.5.4 UV-visible spectroscopy
- •2.3.6 Evaluation of therapeutic efficacy with bioassays
- •2.3.7 Preclinical and clinical researches
- •2.3.8 Structural modifications and developing new analogues
- •2.4 The use of omics technologies in drug discovery and development
- •2.4.1 Genomics
- •2.4.2 Metabolomics
- •2.4.3 Proteomics
- •2.5 Future scope
- •2.6 Conclusion
- •References
- •3.1 Introduction
- •3.2 Bioactive compounds
- •3.2.1 Alkaloids
- •3.2.2 Terpenoids (terpenes)
- •3.2.3 Phenolics
- •3.3 Industrial importance of biological active compounds
- •3.4 Industrial use of MAPs
- •3.5 Essential oils
- •3.6 MAPs in the dye industry
- •3.6.1 Use of MAPs in the perfumery
- •3.6.2 Use of MAPs in cosmetics
- •3.6.3 Use of MAPs in plastic production
- •3.6.4 Other industrial applications
- •3.6.5 MAPs in energy production
- •3.6.6 MAPs in agricultural applications
- •3.7 Salt stress
- •3.7.1 Nutrient
- •3.7.2 Productivity
- •3.7.3 Photosynthesis
- •3.8 Drought stress
- •3.9 Heavy metals
- •3.10 Heat stress
- •3.11 Soil pH
- •3.12 Light intensity
- •3.13 Pest and disease management
- •3.14 Conclusion and future perspective
- •References
- •4.1 Introduction
- •4.2 Toxic compounds and their effects
- •4.2.1 Alkaloids
- •4.2.2 Glycosides
- •4.2.3 Essential oils
- •4.2.4 Saponins
- •4.2.5 Coumarins
- •4.3 Poisonous medicinal plants
- •4.3.1 Digitalis purpurea (foxglove)
- •4.3.2 Atropa belladonna (deadly nightshade)
- •4.3.3 Aconitum napellus (monkshood, aconite)
- •4.3.4 Conium maculatum (hemlock)
- •4.3.5 Nerium oleander (oleander)
- •4.3.6 Datura stramonium (jimsonweed)
- •4.3.7 Ricinus communis (castor bean)
- •4.3.8 Taxus baccata (English yew)
- •4.3.9 Hyoscyamus niger (black henbane)
- •4.3.10 Cicuta virosa (water hemlock)
- •4.3.11 Veratrum viride (false hellebore)
- •4.3.12 Helleborus niger (Christmas rose)
- •4.3.13 Mandragora officinarum (mandrake)
- •4.3.14 Ageratina altissima (white snakeroot)
- •4.3.15 Bryonia alba (white bryony)
- •4.3.16 Colchicum autumnale (autumn crocus)
- •4.3.17 Chelidonium majus Linn. – Papaveraceae
- •4.4 Aromatic plants and poisons
- •4.4.1 Artemisia absinthium (wormwood)
- •4.4.2 Sassafras albidum (sassafras)
- •4.4.3 Lavandula angustifolia (lavender)
- •4.4.4 Rosmarinus officinalis (rosemary)
- •4.4.5 Mentha pulegium (pennyroyal)
- •4.4.6 Eucalyptus globulus (eucalyptus)
- •4.4.7 Myristica fragrans (nutmeg)
- •4.4.8 Thuja occidentalis (white cedar)
- •4.4.9 Illicium verum (star anise)
- •4.4.10 Syzygium aromaticum (clove)
- •4.4.11 Juniperus sabina (savin juniper)
- •4.4.12 Pimpinella anisum (anise)
- •4.4.13 Lavandula stoechas (French lavender)
- •4.4.14 Artemisia vulgaris (mugwort)
- •4.4.15 Melaleuca alternifolia (tea tree)
- •4.4.16 Pelargonium graveolens (rose geranium)
- •4.5 Safe use and precautions
- •4.5.1 Safety guidelines and precautions
- •4.6 Conclusions
- •References
- •5.1 Introduction
- •5.2 Effect of drought or water deficiency on the morphology of medicinal plants
- •5.4 Effect of drought or water deficiency on secondary metabolites of medicinal plants
- •5.5 Different approaches to mitigate the negative effects of drought stress on plants
- •5.6 Case studies
- •5.7 Conclusions
- •References
- •6.1 Introduction
- •6.2 Importance of medicinal and aromatic plants
- •6.3 Salinity effect on medicinal plants
- •6.3.1 Effects on growth and development
- •6.3.2 Impact on photosynthesis and water relations
- •6.3.3 Ionic stress and nutrient imbalance
- •6.3.4 Oxidative stress and antioxidant response
- •6.3.5 Impact on secondary metabolite production
- •6.4 Molecular responses to salinity stress
- •6.5.1 Amino acids
- •6.5.2 Proteins
- •6.5.3 Carbohydrates
- •6.5.4 Lipids
- •6.6 Study of alkaloids through proteomic and other approaches
- •6.7 Phenolic compounds during stress
- •6.8 Strategies for improving salt tolerance in MAPs
- •6.8.1 Exogenous application of plant growth regulators
- •6.8.2 Use of beneficial microorganisms
- •6.8.3 Genetic approaches
- •6.8.4 CRISPR/Cas9 gene editing
- •6.8.5 Agronomic practices
- •6.8.6 Use of mulches
- •6.8.7 Application of organic amendments
- •6.8.8 Silicon supplementation
- •6.8.9 Application of polyamines
- •6.8.10 Nanofertilizers and nanoparticles
- •6.8.11 Application of melatonin
- •6.9.1 Water relations and osmotic adjustment
- •6.9.2 Ion homeostasis and nutrient balance
- •6.10 Molecular mechanisms of salt tolerance
- •6.11.1 Genetic engineering strategies
- •6.11.2 Identification of salt-tolerant genes
- •6.11.3 Use of plant growth regulators
- •6.12 Conclusion and key points
- •References
- •7.1 Introduction
- •7.2 Heavy metals and their effects on the environment
- •7.4 Processes of heavy metal uptake by roots
- •7.5 Transport and accumulation in various plant tissues
- •7.8 Plant defense mechanisms against heavy metals
- •7.9 Molecular and genetic responses to heavy metal contamination
- •7.11 Selection of heavy metal-resistant plants
- •7.12 Case studies
- •7.13 Conclusions
- •References
- •8.1 Introduction
- •8.2 Metabolic and hormonal responses to abiotic stress
- •8.3 Water stress
- •8.3.1 Drought stress
- •8.3.2 Waterlogging stress
- •8.4 Temperature stress
- •8.4.1 High temperature (heat shock)
- •8.4.2 Low-temperature stress
- •8.5 Light stress
- •8.6 Salt stress
- •8.7 Nutrient stress
- •8.8 Heavy metal stress
- •8.9 Molecular docking calculation for stress
- •8.10 Conclusion
- •References
- •Part III: Pharmaceutical use of medicinal plants
- •9.1 Introduction
- •9.2 General properties of medicinal and aromatic plants used in burn treatment
- •9.2.1 Phytochemical content and mechanisms of action
- •9.2.2 Antimicrobial effects
- •9.2.3 Wound-healing effects
- •9.2.4 Analgesic effects
- •9.2.5 Advantages and disadvantages of herbal treatments
- •9.2.5.1 Advantages
- •9.2.5.2 Disadvantages
- •9.3 Medicinal and aromatic plants used in burn treatment
- •9.3.1 Aloe vera
- •9.3.1.1 Clinical effects
- •9.3.2 Calendula officinalis (Calendula)
- •9.3.3 Centella asiatica (gotu kola)
- •9.4 Molecular basis of plant action mechanisms
- •9.4.1 Cellular mechanisms in wound healing
- •9.4.2 Innovative research methods in herbal treatments
- •9.4.2.1 Omic technologies: genomic, proteomic, and metabolomic approaches
- •9.5 Formulation and application methods of herbal products
- •9.5.1 Pharmaceutical formulations
- •9.5.2 Dosage and application methods
- •9.5.3 Nanotechnological approaches
- •9.5.4 Factors affecting chemical stability
- •9.5.4.1 Stability enhancement methods
- •9.5.4.2 Importance of storage conditions
- •9.5.4.3 Stability tests and quality control
- •9.6 Clinical research and evidence-based practices
- •9.6.1 Clinical studies
- •9.6.2.1 Meta-analyses and literature reviews
- •9.7 Safety and side effects
- •9.7.1 Toxicological risks
- •9.7.2 Side effects and contraindications
- •9.8 Integration of traditional knowledge and modern science
- •9.8.1 Ethnobotany and traditional knowledge
- •9.8.2 Cultural and regional diversity
- •9.9 Future research areas and innovation
- •9.9.1 Pharmacogenetics and personalized medicine
- •9.9.2 Biodegradable and smart materials
- •9.9.3 Combined use of herbal treatments
- •9.10 Conclusion
- •References
- •10.1 Introduction
- •10.2 COPD
- •10.3 Asthma
- •10.4 Pneumonia
- •10.5 Lung cancer
- •References
- •11.1 Introduction
- •11.2 Oxidative stress
- •11.2.1 Reactive oxygen species
- •11.2.2 Sources and generation of free radicals
- •11.3.1 Lipid peroxidation
- •11.3.2 Protein oxidation
- •11.3.3 DNA oxidation
- •11.4 Defense of the organism against ROS
- •11.4.1 Free radicals and antioxidants
- •11.4.2 Antioxidants action mechanism
- •11.5 Methods for determination of antioxidative activity
- •11.5.1 Methods based on hydrogen atom transfer
- •11.5.2 Methods based on electron transfer
- •11.5.3 Other methods for determination of antioxidant potential
- •11.6 Medicinal and aromatic plants as natural antioxidants
- •11.7 MAPs with antioxidant activity
- •11.8 Conclusion
- •References
- •12.1 Introduction
- •12.2 Definition, historical documents, and distribution related to the study of the usage of MAPs
- •12.3 Antibacterial activity of MAPs
- •12.4 Extracts and essential oils from MAPs as antibacterial agents
- •12.5 Compounds of essential oils with antibacterial properties and their activity against a variety of bacterial strains
- •12.6.2 Terpenoids from MAPs as antibacterial agents
- •12.6.3 Alkaloids from MAPs as antibacterial agents
- •12.7.2 Clinopodium nepeta (L). Kuntze
- •12.7.3 Lavandula officinalis
- •12.7.4 Helichrysum italicum
- •12.7.5 Mentha piperita
- •12.8 Conclusion
- •References
- •13.1 Introduction
- •13.2 Medicinal and aromatic plant-derived extracts
- •13.2.1 Extraction techniques of MAPs
- •13.2.2 Influence of extraction operational parameters
- •13.3 MAPs in skin care products
- •13.3.1 MAPs as photoprotective agents against UV light and skin damage
- •13.3.2 Regenerative and wound-healing properties of MAP-derived agents
- •13.3.3 MAPs as skin anti-aging and whitening agents
- •13.4 MAPs in hair cosmetics
- •13.4.1 MAPs in hair products
- •13.4.2 MAPs in hair growth products
- •13.5 MAPs in oral hygiene products
- •13.5.1 Formulations for toothpaste and mouthwash
- •13.5.2 MAPs in prevention of dental caries
- •13.6 MAPs enhanced by sustainable materials in cosmetics
- •13.6.1 Nanotechnology in cosmetic formulations
- •13.6.2 Innovative nanocarrier materials
- •13.7 Conclusion
- •Abbreviations
- •References
- •14.1 Introduction

Chapter 6 Impact of salinity stress on medicinal and aromatic plant biotechnology 233
Table 6.1: Impact of salinity on the growth and development of medicinal plants.
Parameter Observation Example species
Seed germination Reduced germination rate under high salinity levels Catharanthus roseus
Chlorophyll
Content and
biomass
Allocation Shift toward root biomass; reduction in shoot biomass Moringa oleifera
Flowering Delayed or inhibited flowering due to ionic and osmotic stress Trachyspermum
Decreased chlorophyll concentration and photosynthetic
efficiency
St. John’s wort
ammi
of reproductive success and medicinal compound production, reduced flowering
under salinity stress can severely affect yield, particularly in species that derive medicinal properties from their flowers [41].
6.3.2 Impact on photosynthesis and water relations
Salinity stress disrupts the photosynthetic machinery of MAPs, reducing both the chlorophyll content and photosynthetic efficiency. For example, salinity-stressed St. John’s
wort plants displayed an 18.9% reduction in chlorophyll levels compared to unstressed controls [42]. Furthermore, salinity affects critical parameters related to photosynthetic performance. Reductions in F
(maximum quantum yield of PSII), Fv/F
v/Fm
(maximum primary yield of PSII photochemistry), and PI (performance index) under
salinity stress highlight significant losses in photosynthetic capacity, leading to reduced biomass and growth [43].
Salt stress disrupts plant water uptake by altering soil solution dynamics and creating artificial drought conditions, despite water availability [44, 45]. To counteract
this osmotic challenge, plants synthesize osmoprotectants like amino acids and carbohydrates, which help maintain cellular equilibrium in saline environments [46]. This
adaptive response enables plants to mitigate some of the harmful effects of salt exposure, showcasing their resilience to environmental stressors.
6.3.3 Ionic stress and nutrient imbalance
Salt stress in plants leads to an overaccumulation of sodium and chloride ions, disrupting key cellular processes in Figure 6.1. Studies on moringa have shown increased Na
and Cl- levels under saline conditions, accompanied by reduced potassium uptake [47,
48]. This ionic imbalance interferes with the absorption of essential nutrients like potas-
o
+

Figure 6.1: Effect of soil salinity on photosynthesis and water retention.
234 Fatemeh Ahmadi
sium, calcium, and magnesium [49]. Consequently, plants experience impaired growth,
metabolic disturbances, and decreased productivity due to the combined effects of ion
toxicity and nutrient deficiencies [50].
6.3.4 Oxidative stress and antioxidant response
Salt stress triggers excessive production of reactive oxygen species (ROS) in plants,
threatening cellular components. Plants respond by activating antioxidant defenses, including enzymes like superoxide dismutase and catalase, as well as nonenzymatic compounds such as ascorbic acid and flavonoids [51, 52]. These mechanisms help neutralize
ROS, protecting cellular structures and enhancing plant resilience under saline conditions. Studies on medicinal plants have shown increased antioxidant activity correlating
with salt exposure levels, demonstrating plants’ adaptive strategies against oxidative
stress [53, 54].
6.3.5 Impact on secondary metabolite production
Salt stress paradoxically affects medicinal plants, often stunting growth, while boosting the production of valuable secondary metabolites [55]. Under saline conditions,

Chapter 6 Impact of salinity stress on medicinal and aromatic plant biotechnology 235
these plants frequently increase their synthesis of compounds like tannins, saponins,
and phenols [56]. This response involves changes in key biosynthetic pathways that
are sensitive to environmental stress [57]. Research has shown that salt exposure can
enhance essential oil production in various aromatic plants, potentially serving as a
survival mechanism [58]. This phenomenon has significant implications for the medicinal plant industry, suggesting that controlled salinity could be used to stimulate the
production of high-value bioactive compounds [59]. By integrating stress management
into cultivation practices, growers may enhance both the therapeutic efficacy and economic value of medicinal plants. This approach represents a promising strategy for
optimizing medicinal plant production in challenging environments [60].
6.4 Molecular responses to salinity stress
Medicinal plants adapt to salt stress through complex molecular mechanisms, involving changes in gene expression, protein activity, and metabolite production. Proteomic studies have revealed stress-induced alterations in proteins crucial for tolerance,
photosynthesis, and secondary metabolite synthesis [61]. Salt exposure triggers the upregulation of genes linked to osmolyte production, ion transport, and antioxidant defenses. This molecular plasticity not only enhances salt tolerance but also promotes
the accumulation of valuable secondary metabolites. Understanding these intricate
molecular responses provides insights into how medicinal plants adapt to saline conditions, while potentially increasing their therapeutic and economic value [62].
6.5 Salt stress and primary metabolites
in medicinal plants
6.5.1 Amino acids
Salt stress triggers significant shifts in plant amino acid metabolism. Many amino
acids, including alanine, arginine, and glycine, increase under saline conditions, with
proline showing the most dramatic rise. Nonprotein amino acids and amides also accumulate. Proline’s buildup, observed in various medicinal plants, results from reduced proline oxidase activity and serves as a key osmoprotectant [60]. This helps
maintain cellular balance and structure under salt stress. The overall increase in free
amino acids, seen in plants like Catharanthus roseus, aids in osmotic adjustment and
provides resources for energy and biosynthesis during stress. These metabolic
changes highlight plants’ adaptive strategies for surviving in saline environments,
demonstrating the crucial role of amino acids in stress resilience [57].

236 Fatemeh Ahmadi
6.5.2 Proteins
The elevation of free amino acids in plants experiencing salt stress is partially explained by the breakdown of proteins. For example, protein degradation has been observed in Catharanthus roseus exposed to salinity [63]. In chamomile and sweet marjoram, salt stress reduced the levels of soluble proteins, likely due to protein
aggregation within the cells [64]. In Achillea fragratissima, a decline in crude protein
synthesis was noted at 4,000 ppm salinity [65]. However, some studies indicate an increase in protein synthesis under higher salinity levels, suggesting that plants may
store nitrogen in proteins for use during recovery from stress [66].
6.5.3 Carbohydrates
Salinity disrupts carbohydrate metabolism in plants, typically leading to imbalances
due to reduced photosynthesis and nutrient availability. For instance, fennel plants
exhibited a decrease in carbohydrate content under saline conditions [67]. In contrast,
plants such as Salvia officinalis and Satureja hortensis showed an increase in carbohydrates as salinity levels rose, highlighting species-specific responses to salt stress
[68, 69].
6.5.4 Lipids
Salt stress significantly impacts the lipid profile of plants, influencing fatty acid and
oil synthesis. For example, in Ricinus communis, salinity reduced oil yield in roots but
increased oil content in shoots [70]. In Coriandrum sativum, salinity stress led to a notable decrease in total fatty acid content, with reductions in key fatty acids such as αlinolenic and linoleic acids as NaCl concentrations increased [71–75].
6.6 Study of alkaloids through proteomic and other approaches
Proteomic studies have enhanced our knowledge of secondary metabolite production
in medicinal plants, revealing complex mechanisms behind the synthesis of therapeutically valuable compounds [76, 77]. Researchers are using advanced techniques like
cell cultures and metabolic engineering to increase the yield of these naturally limited
substances. Studies on plants such as Catharanthus roseus have shown how various
factors, including phytohormones and salt stress, affect protein expression and alkaloid production [78, 79]. This research has identified key enzymes and proteins in-

Chapter 6 Impact of salinity stress on medicinal and aromatic plant biotechnology 237
volved in metabolite synthesis, providing insights into how plants respond to stress
by altering their biochemical pathways. These findings open new possibilities for enhancing the production of medicinal compounds for pharmaceutical and commercial
use [80]. In Chelidonium majus, two-dimensional gel electrophoresis revealed 21 proteins associated with stress signaling, nucleic acid binding, and defense responses,
shedding light on the molecular mechanisms underpinning salinity tolerance [81].
Studies on Papaver somniferum identified codeinone reductase as a central enzyme in
morphine biosynthesis, emphasizing its critical role in stress-induced secondary metabolite production [82].
These advancements in proteomics provide invaluable insights into the complex
molecular pathways that regulate secondary metabolite production, paving the way
for optimized cultivation practices and biotechnological innovations to maximize the
medicinal and economic potential of these compounds [83, 84].
6.7 Phenolic compounds during stress
Phenolic compounds have emerged as important indicators of salt stress in plants.
These diverse molecules, numbering around 9,000, play crucial roles in plant defense,
particularly in neutralizing ROS generated during stress conditions [85]. Salt stress disrupts photosynthetic processes, leading to increased ROS production and oxidative
damage. In response, plants synthesize various phenolic compounds, including phenolic
acids, flavonoids, and proanthocyanidins, which act as antioxidants [85]. Research on
crop plants consistently shows elevated phenolic content under saline conditions. For
example, spearmint and Achillea fragratissima exhibit higher levels of phenolic acids
and tannins when exposed to salt stress. Matricaria chamomilla demonstrates increased
production of specific phenolic acids like protocatechuic, chlorogenic, and caffeic acids.
Similarly, Nigella sativa and Mentha pulegium show a positive correlation between phenolic accumulation and salinity levels [86]. In Nigella, cultivated in saline soils, compounds such as quercetin, apigenin, and trans-cinnamic acid are found in higher concentrations, illustrating the plant’s adaptive response to salt stress through enhanced
phenolic synthesis.
6.8 Strategies for improving salt tolerance in MAPs
The growing issue of soil salinity poses a major threat to the cultivation of MAPs, which
are valued for their therapeutic and economic significance. To address this, various strategies have been developed, ranging from traditional agronomic methods to advanced biotechnological approaches, aimed at mitigating the impact of salinity on MAPs [86]

238 Fatemeh Ahmadi
Table 6.2: Strategies to improve salt tolerance in medicinal plants.
Strategy Mechanism Example
plants
Use of plant growth
regulators
Application of
Mycorrhizae
Genetic engineering Overexpression of salt-tolerance genes like SOS and PCS Artemisia annua
Exogenous
osmoprotectants
Enhances stress tolerance through hormonal regulation and
antioxidant activity
Improves nutrient uptake and water relations under saline
conditions
Protects cellular structures and maintains osmotic balance Chamomile and
Basil and
chamomile
Marjoram
sage
(Table 6.2). This section explores these strategies, focusing on their mechanisms and effectiveness in enhancing salt tolerance [87].
6.8.1 Exogenous application of plant growth regulators
Plant growth regulators (PGRs) have shown promise in enhancing the salt tolerance of
MAPs by influencing key physiological processes [88]. Among these, salicylic acid (SA) has
demonstrated particular efficacy in alleviating salt stress effects. Research on St. John’s
wort, exposed to saline conditions, revealed that SA treatment significantly boosted
growth parameters and photosynthetic performance [89]. SA application led to marked
improvements in chlorophyll levels, photosystem efficiency, and electron transport. Additionally, SA treatment reduced stress hormone levels, while increasing antioxidant enzyme activity, thereby improving the plant’s ability to manage oxidative stress under saline conditions. These findings underscore the potential of PGRs, especially SA, as a
practical approach to enhancing salt tolerance in medicinal plants, offering a promising
strategy for maintaining crop productivity in salt-affected areas [90].
Gibberellic acid (GA3): Gibberellic acid has shown considerable potential in alleviating the detrimental effects of salinity on MAPs. For instance, in basil (Ocimum basili-
cum), GA3 application under saline conditions significantly improved growth metrics
such as plant height, leaf area, and biomass (both fresh and dry weight). Moreover,
GA3 enhanced the production of essential oils and their primary constituents, including linalool and methyl chavicol. These benefits are attributed to GA3’s role in preserving membrane stability, enhancing antioxidant enzyme activity, and regulating
osmolyte accumulation, which collectively contribute to improved stress resilience
[91, 92].
Brassinosteroids (BRs): Brassinosteroids have also demonstrated promising effects on salt tolerance in MAPs. For example, foliar application of 24-epibrassinolide
in chamomile (Matricaria chamomilla) under saline conditions enhanced growth,

Chapter 6 Impact of salinity stress on medicinal and aromatic plant biotechnology 239
photosynthetic efficiency, and essential oil content. BRs improve salt tolerance by activating the plant’s antioxidant defense mechanisms, enhancing water relations, and
promoting the accumulation of compatible solutes. These combined effects enable
plants to maintain physiological stability and adapt to saline environments more effectively [93].
The application of PGRs, including SA, GA3, and BRs, underscores their potential
as valuable tools for mitigating salt stress in MAPs, ultimately supporting improved
growth, productivity, and secondary metabolite synthesis in challenging environments.
6.8.2 Use of beneficial microorganisms
Utilizing beneficial microorganisms, such as arbuscular mycorrhizal fungi (AMF) and
plant growth-promoting rhizobacteria (PGPR), offers an eco-friendly approach to enhance salt tolerance in MAPs [94]. These microorganisms improve plant resilience to
salinity stress by enhancing nutrient uptake, increasing water absorption, balancing
hormones, and activating stress-related biochemical pathways. AMF enhances root–soil
interactions, facilitating phosphorus uptake, while PGPR promotes root growth, produces beneficial phytohormones, and bolsters antioxidant defenses 95]. For instance, inoculating Origanum majorana with Glomus mosseae helps mitigate salinity effects by improving growth and essential oil production. Similarly, in basil (Ocimum basilicum),
PGPR strains like Pseudomonas putida and Bacillus lentus enhance growth and essential
oil yield under salt stress by improving antioxidant enzyme activity and nutrient absorption [96]. Overall, these beneficial microbes play a crucial role in supporting plant
health and productivity in saline environments, aligning with sustainable agricultural
practices.
6.8.3 Genetic approaches
Genetic strategies hold great potential for improving salt tolerance in MAPs by combining traditional breeding with advanced biotechnological methods. Traditional
breeding allows for the selection and crossing of salt-tolerant varieties, while modern
techniques such as genetic engineering and marker-assisted selection enable precise
targeting of genes linked to salinity resistance [97]. These approaches can lead to the
development of MAPs that are more resilient to saline conditions, enhancing their medicinal and economic value. Research has identified specific genes that contribute to
salt tolerance, such as the Na
introduced into Artemisia annua, resulting in improved growth and artemisinin production under salt stress [98]. Overall, these genetic advancements provide promising
+/H+
antiporter gene (NHX1), which has been successfully

240 Fatemeh Ahmadi
pathways for enhancing the resilience of MAPs against salinity, supporting sustainable agricultural practices.
RNA Interference (RNAi) technology: RNAi has been used to suppress genes that
hinder salt tolerance [99]. For example, silencing the SmMYB39 gene in Salvia miltior-
rhiza enhanced the plant’s tolerance to salt stress. This was achieved by increasing
the production of phenolic acids and tanshinones, which are important medicinal
compounds [100].
6.8.4 CRISPR/Cas9 gene editing
The CRISPR/Cas9 genome editing system offers precise and efficient modifications,
making it a promising approach for enhancing salinity tolerance in MAPs. While its
application in MAPs is still developing, this tool shows great promise for creating salttolerant plants with improved medicinal properties [101].
6.8.5 Agronomic practices
Agronomic interventions are essential for managing the effects of salinity on MAPs by
improving soil conditions and ensuring better plant growth in saline environments [102].
Irrigation management: Efficient irrigation techniques can control soil salinity
levels. Drip irrigation, for example, has been shown to maintain lower salinity in the
root zone compared to traditional methods like furrow irrigation. Studies on Rosmar-
inus officinalis (rosemary) revealed that drip irrigation not only improved plant
growth but also increased the yield and quality of essential oils under saline conditions [103].
6.8.6 Use of mulches
Mulching is an effective agronomic strategy to mitigate salinity stress. Organic mulches
help reduce water loss through evaporation and prevent salt buildup in the root zone.
For example, straw mulch applied to Salvia officinalis (sage) significantly enhanced
growth and essential oil yield under saline conditions. This practice also improved soil
moisture retention, minimized temperature fluctuations, and boosted soil biological activity [104, 105].

Chapter 6 Impact of salinity stress on medicinal and aromatic plant biotechnology 241
6.8.7 Application of organic amendments
Organic amendments, such as compost and biochar, have proven effective in improving soil health and mitigating salinity-induced stress in MAPs. These amendments enhance soil structure, promote water retention, and alleviate the negative effects of
salt stress [106]. For instance, in Foeniculum vulgare (fennel), vermicompost application significantly enhanced plant-growth-boosted essential oil yield, and improved antioxidant activity under saline conditions. These improvements were attributed to enhanced soil properties, increased nutrient availability, and stimulation of beneficial
microbial activity [107, 108].
6.8.8 Silicon supplementation
Silicon (Si) supplementation has emerged as an effective method for enhancing plant
tolerance to salinity [109]. While not essential for growth, Si plays a vital role in mitigating abiotic stresses, including salt stress. In basil, Si treatment has been shown to
alleviate the negative impacts of salinity, resulting in improved growth, enhanced
photosynthesis, and increased essential oil production [110]. These benefits are linked
to heightened antioxidant enzyme activity, better water absorption, reduced sodium
accumulation, and improved potassium balance. Similarly, in peppermint (Mentha pi-
perita), Si application leads to better growth and essential oil yield, while reinforcing
antioxidant defenses and maintaining membrane integrity under saline conditions.
Overall, Si supplementation proves beneficial for promoting resilience in plants facing
salinity stress [111].
6.8.9 Application of polyamines
Polyamines, including putrescine, spermidine, and spermine, are crucial for regulating plant growth and stress responses, especially under saline conditions [112]. Their
application has been recognized for enhancing salt tolerance in MAPs by stabilizing
cell membranes, reducing oxidative damage from ROS, and balancing ion levels. Additionally, polyamines promote the expression of stress-related genes, improve nutrient uptake, and assist in osmotic adjustment, thereby strengthening plants’ ability
to cope with salinity [113]. For example, in chamomile, putrescine application improved growth, flower yield, and essential oil content under salt stress by increasing
antioxidant enzyme activity and enhancing photosynthetic efficiency [114]. In sage,
spermidine application helped mitigate salinity effects by improving water retention and promoting compatible solute accumulation, which supported growth and
essential oil production. Overall, polyamines present a promising strategy for sus-

242 Fatemeh Ahmadi
tainable MAP cultivation in saline environments due to their multifunctional roles
in enhancing plant resilience [115].
6.8.10 Nanofertilizers and nanoparticles
Nanofertilizers and nanoparticles are innovative tools for enhancing nutrient uptake
efficiency and improving plant resilience to salinity. In Mentha piperita (peppermint),
zinc oxide nanoparticles (ZnO NPs) significantly enhanced growth, essential oil yield,
and antioxidant activity under saline conditions. These improvements were linked to
increased accumulation of compatible solutes, better photosynthesis, and more stable
nutrient levels [116]. Similarly, in basil, iron oxide nanoparticles (Fe
salt stress by improving growth, increasing essential oil production, and boosting antioxidant enzyme activity. These nanoparticles also supported secondary metabolite
synthesis and helped maintain membrane stability in plants subjected to salinity [117].
NPs) alleviated
₃O₄
6.8.11 Application of melatonin
Melatonin is recognized as an effective enhancer of salt tolerance in various plants, including medicinal and aromatic species. Acting both as a growth regulator and antioxidant, melatonin helps mitigate salinity stress, which has garnered significant research
interest. In lemon balm (Melissa officinalis), applying melatonin has been shown to improve growth, essential oil yield, and antioxidant activity under saline conditions [118].
This enhancement is linked to melatonin’s ability to promote the accumulation of compatible solutes like proline, which aids in osmotic regulation, stabilizes chloroplast
structures, and maintains ionic balance by reducing sodium uptake and increasing potassium retention. These mechanisms collectively improve the plant’s resilience to salinity [84].
Melatonin’s potential to enhance salt tolerance, combined with its ability to boost
secondary metabolite production, makes it a promising solution for the sustainable
cultivation of MAPs in saline environments, offering both therapeutic and economic
benefits [119]. Similarly, in Lavandula angustifolia (lavender), melatonin treatment alleviated the adverse effects of salt stress on growth and essential oil production. The
protective mechanism involved heightened antioxidant enzyme activity, improved
water relations, and stabilization of membrane integrity [120, 121].
The strategies for improving salt tolerance in MAPs are diverse, ranging from
basic agronomic practices to advanced biotechnological techniques. Exogenous applications of PGRs, beneficial microorganisms, silicon, polyamines, and melatonin have
shown great potential for enhancing salt tolerance in MAPs by modulating physiologi-
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