Exploring the Biochemical Profiles of Medicinal Plants Cultivated Under Stressful Environmental Conditions

Manoj Kumar Sharma*

Department of Botany, Janta Vedic College, Baraut (Baghpat) India.

Corresponding Author E-mail: mbhardwaj1501@gmail.com

DOI : http://dx.doi.org/10.12944/CARJ.12.1.07

Article Publishing History

Received: 29 Nov 2023
Accepted: 27 Jan 2024
Published Online: 31 Jan 2024

Review Details

Plagiarism Check: Yes
Reviewed by: Dr. Abdulrahman R.Mahmood
Second Review by: Dr. Mohini Kuchekar
Final Approval by: Dr. Torit Baran Bagchi

Article Metrics

Views     PDF Download PDF Downloads: 366

Google Scholar

Abstract:

The therapeutic potential of medicinal plants, attributed to diverse bioactive chemicals, is impacted by dynamic metabolic changes under environmental stressors such as salt, drought, and heavy metal exposure. This study aimed to elucidate the biochemical alterations in stressed medicinal plants and their implications for therapeutic qualities. Two groups of plants were compared: one cultivated in optimal conditions, serving as a control, and the other exposed to stressors mimicking real-world scenarios. Biochemical parameters, including total phenolic content, flavonoids, antioxidant activity, and specific secondary metabolites, were assessed using established analytical methods. In a case study involving Aloe vera under elevated soil salinity, a significant decrease in total phenolic content and antioxidant activity was observed. Surprisingly, aloin, an anti-inflammatory bioactive compound, exhibited notable upregulation. Stress-induced biochemical variations in medicinal plants differed significantly from those in ideal conditions, demonstrating adaptive responses, including modulation of secondary metabolite synthesis. While certain stresses led to an overall decrease in secondary metabolites, specific instances showed an overexpression of beneficial chemicals. Exploring drought stress in Lavandula angustifolia (lavender), Vitis vinifera, and Artemisia tridentata, revealed a marked reduction in flavonoids and specific secondary metabolites. The study aimed to establish connections between metabolic alterations and stress-adaptive reactions. The findings contribute to understanding how environmental stress influences the therapeutic efficacy of medicinal plants and shed light on the molecular pathways governing stress-induced changes. In the investigation of heavy metal exposure, Hypericum perforatum, and Coriandrum sativum subjected to cadmium-contaminated soil exhibited an overall decline in secondary metabolites, including reduced hypericin content. Contrarily, hyperforin, linked to antidepressant properties, experienced a significant increase. These results have implications for pharmaceutical applications, sustainable farming practices, and the conservation of medicinal plant biodiversity amidst changing environmental dynamics. The study underscores the intricate relationship between environmental stressors and the biochemical makeup of medicinal plants, emphasizing their resilience and capacity to produce valuable bioactive molecules in challenging conditions. The comparative biochemical analysis of stressed medicinal plants involves assessing various parameters to comprehend how stress influences plant metabolism and bioactive compound production.

Keywords:

Antioxidant activity; Abiotic stress; Bioactive compounds; Biochemical profiles; Medicinal plants; Metabolic pathways; Plant cultivation; Phytochemicals; Stressful environments; Secondary metabolites

Download this article as: 

Copy the following to cite this article:

Sharma M. K. Exploring the Biochemical Profiles of Medicinal Plants Cultivated Under Stressful Environmental Conditions. Curr Agri Res 2024; 12(1). doi : http://dx.doi.org/10.12944/CARJ.12.1.07

Copy the following to cite this URL:

Sharma M. K. Exploring the Biochemical Profiles of Medicinal Plants Cultivated Under Stressful Environmental Conditions. Curr Agri Res 2024; 12(1). Available from: https://bit.ly/48Zaozw


Introduction

The necessity for sustainable and creative approaches to healthcare has increased recently due to the growing global population and environmental issues. The potential therapeutic applications of medicinal plants, which are rich sources of bioactive chemicals, have attracted a lot of attention. However, several environmental factors, particularly stress levels, can affect the quantity and quality of these bioactive chemicals in medicinal plants. Stress factors like as drought, salinity, and temperature fluctuations are prevalent globally and significantly affect the growth and development of plants. Extended periods of stress allow plants to adjust and develop more resilience, which boosts their resistance. Optimising the growth of medicinal plants and realising their full therapeutic potential requires an understanding of their physiological reactions to stress.1

The purpose of this review is to perform a thorough comparative biochemical investigation of a few selected medicinal plants that are cultivated under stressful environments. We aim to clarify the adaptive processes used by these plants and investigate how these responses may promote the accumulation of bioactive chemicals with potential therapeutic value by examining the biochemical changes generated by stress.2

The traditional medical applications and economic significance of the medicinal plants considered for this study have guided the selection process. Our objective is to evaluate the metabolic responses of these plants and imitate real-world situations by exposing them to controlled stress settings.3 In addition to advancing our knowledge of the alterations brought about by stress in these therapeutic plants, this comparative study will offer insightful information about how these species can be sustainably cultivated under a variety of environmental circumstances.

Key Objectives

Identification of Stress-Responsive Biochemical Pathways

Explore the biochemical pathways that are activated or modulated in response to stress conditions in medicinal plants.

Quantification of Bioactive Compounds

Measure the concentrations of key bioactive compounds, such as secondary metabolites and antioxidants, in stressed and non-stressed plants.

Assessment of Medicinal Potential

Evaluate the potential enhancement of medicinal properties in plants subjected to stress conditions by comparing the bioactive compound profiles with traditionally grown counterparts.

Elucidation of Stress Adaptation Mechanisms

Investigate the molecular and biochemical mechanisms employed by medicinal plants to adapt to stress, providing insights into their resilience and survival strategies.

Practical Implications for Cultivation

Provide practical recommendations for the cultivation of medicinal plants under stress conditions, aiming to optimize the production of bioactive compounds for medicinal purposes.

Through the pursuit of these goals, the project hopes to further our understanding of the complex interactions between medicinal plants and their surroundings and provide useful information to the domains of plant biochemistry, traditional medicine, and sustainable agriculture. The results could lead to the creation of methods to increase these plants’ yield and therapeutic value, which would ultimately be advantageous for the pharmaceutical sector and the world’s health care system.4 Plant scientists can use the provided data to identify and study these occurrences to gain a better understanding of the mechanics behind plant stress responses. We can anticipate the effects of recently discovered compounds and the abuse of chemicals in agriculture by comprehending this interaction, which is a natural characteristic of the planet.5

A review paper will go into great detail about the complex molecular mechanisms that medicinal plants use to adjust to and manage stress.

Stress-Responsive Biochemical Pathways

Stress reactions are essential to plants’ ability to survive in harsh conditions. To detect and react to a wide range of stressors, including biotic (such as infections) and abiotic (such as drought and salinity) challenges, plants have developed complex signalling pathways.6 These signalling pathways frequently entail an intricate web of molecular actions that impact metabolic functions, gene expression, and physiological adjustments.

Abscisic Acid (ABA) Pathway:

ABA is a key phytohormone that plays a central role in response to abiotic stresses, such as drought and high salinity.7

ABA signalling involves ABA receptors, which activate downstream components, leading to changes in gene expression and physiological responses like stomatal closure to reduce water loss.8

Jasmonic Acid (JA) and Ethylene Pathways:

These pathways are often activated in response to biotic stresses, including herbivore attack and pathogen infection.9

JA and ethylene signalling pathways crosstalk and regulate the expression of defence-related genes, leading to the synthesis of compounds like jasmonic acid and ethylene that deter herbivores and inhibit pathogen growth.10

Salicylic Acid (SA) Pathway:

SA is another important phytohormone involved in plant defence against pathogens, particularly biotrophic and hemi-biotrophic microbes.

SA signalling induces the expression of pathogenesis-related (PR) genes and activates systemic acquired resistance (SAR), enhancing the plant’s ability to resist subsequent infections.11,12

Mitogen-Activated Protein Kinase (MAPK) Cascades:

MAPK cascades are central components in stress signalling pathways, integrating signals from various upstream receptors.13

Activated MAPKs phosphorylate and activate transcription factors, leading to the expression of stress-responsive genes.14

Calcium-Mediated Signaling:

Calcium ions (Ca2+) serve as second messengers in response to various stresses, triggering downstream signalling events.15

Changes in cytosolic calcium levels activate calcium-dependent protein kinases (CDPKs) and other calcium-binding proteins, modulating gene expression and other stress responses.16

Reactive Oxygen Species (ROS) Signaling:

ROS, such as superoxide radicals and hydrogen peroxide, are produced in response to stress and act as signalling molecules.17

ROS activate antioxidant defence systems and signal transduction pathways, contributing to stress acclimation.18

Phosphatidylinositol Signaling Pathway:

Phosphatidylinositol signalling is involved in stress responses, including responses to drought and salinity.19

Phospholipase C (PLC) is activated, leading to the production of inositol trisphosphate (IP3) and diacylglycerol (DAG), which modulate ion channels and gene expression.20

These signalling pathways often intersect and crosstalk, forming a complex regulatory network that allows plants to fine-tune their responses to different stress conditions. Understanding these pathways is crucial for developing strategies to enhance stress tolerance in crops and other plants of economic or ecological importance.

Reactive Oxygen Species (ROS) and Antioxidant Defence:

Reactive Oxygen Species (ROS) play a dual role in plant biology, serving as both signalling molecules and potential sources of cellular damage. While ROS are generated in response to various environmental stresses, their accumulation can lead to oxidative stress, causing cellular damage to lipids, proteins, and nucleic acids.21 To counteract the detrimental effects of ROS, plants have evolved sophisticated antioxidant defence mechanisms.

Antioxidant defence systems include enzymatic and non-enzymatic components that work together to detoxify ROS and maintain cellular redox balance. Enzymes such as superoxide dismutase (SOD), catalase (CAT), and peroxidases play crucial roles in scavenging ROS, converting them into less harmful compounds. Non-enzymatic antioxidants, including ascorbate, glutathione, and tocopherols, also contribute to ROS detoxification.22

The delicate balance between ROS production and antioxidant defence is essential for plant health and stress tolerance. Dysregulation of this balance can result in oxidative damage, compromising cellular functions and ultimately affecting plant growth and development.

Phytohormones and Stress Responses

Phytohormones, also known as plant hormones, are signalling molecules that regulate various physiological processes in plants. These hormones play a crucial role in plant growth, development, and responses to environmental stimuli, including stress.19, 22, 23 Here’s a brief overview of some key phytohormones and their involvement in stress responses:

Abscisic Acid (ABA)

ABA is a key phytohormone involved in the plant’s response to various stress conditions, particularly drought and high salinity.

It regulates stomatal closure to reduce water loss during drought, promotes root growth to access deeper soil layers for water uptake, and influences gene expression related to stress tolerance.7, 8, 24, 25

Jasmonic Acid (JA) and Methyl Jasmonate (MeJA):

These hormones are associated with the plant’s response to biotic stress, such as herbivore attack and pathogen infection.

They induce the expression of defence-related genes, leading to the production of secondary metabolites, toxins, and proteins that deter herbivores and inhibit pathogen growth.9, 10

Salicylic Acid (SA)

SA is another hormone involved in plant defence responses, particularly against biotrophic pathogens (those that extract nutrients from living plant cells).

It induces the expression of defence genes and activates systemic acquired resistance (SAR), long-lasting and broad-spectrum resistance to pathogens.11, 12, 26

Ethylene (ET)

Ethylene is a gaseous hormone that regulates various aspects of plant growth and development, as well as responses to both biotic and abiotic stress.

It is involved in the response to mechanical stress, pathogen attack, and senescence. In flooded conditions, ethylene can promote aerenchyma formation to enhance oxygen transport to submerged roots.8, 27

Cytokinins

Cytokinins play a role in cell division and differentiation and are involved in mitigating stress responses, especially drought stress.

They promote the synthesis of antioxidants and enhance the plant’s ability to withstand oxidative stress.28, 29

Gibberellins (GAs):

GAs are primarily associated with promoting stem elongation and seed germination, but they also play a role in stress responses.

They may be involved in responses to environmental stresses such as cold and nutrient deficiencies.30, 31

Brassinosteroids (BRs):

BRs influence cell elongation, division, and differentiation and contribute to stress tolerance, including resistance to drought and salinity.

Plants integrate these hormonal signals to orchestrate adaptive responses to various environmental stresses. The crosstalk between different phytohormones allows plants to fine-tune their responses and optimize survival strategies in challenging conditions.32, 33

Secondary Metabolite Biosynthesis

Secondary metabolites are organic compounds that are not directly involved in the normal growth, development, or reproduction of an organism, including plants. However, they often play crucial roles in plant adaptation to environmental stresses. These compounds are synthesized through complex biochemical pathways and contribute to the plant’s defence mechanisms against herbivores, pathogens, and various abiotic stresses. Here’s a brief overview of secondary metabolite biosynthesis during plant growth stress:

Phenolic Compounds

Phenolic compounds, including flavonoids and phenolic acids, are common secondary metabolites. They act as antioxidants, scavenging reactive oxygen species (ROS) produced under stress conditions.

These compounds also contribute to the plant’s defence against pathogens and herbivores by inhibiting microbial growth and deterring herbivore feeding.34

Terpenoids

Terpenoids, such as terpenes and essential oils, are synthesized from isoprenoid precursors. They play roles in plant defence against herbivores and pathogens.

Some terpenoids act as volatile organic compounds (VOCs) that can attract predators of herbivores or serve as signals to neighbouring plants about impending stress.35

Alkaloids

Alkaloids are nitrogen-containing compounds with diverse structures and functions. Examples include alkaloids like nicotine, caffeine, and morphine.

Alkaloids often have toxic effects on herbivores and may deter feeding. They can also have antimicrobial properties and contribute to the plant’s defence against pathogens.36

Glucosinolates

Glucosinolates are sulphur-containing compounds found mainly in the Brassicaceae family. They play a role in defence against herbivores and pathogens.

When plant tissues are damaged, glucosinolates are hydrolysed to produce toxic compounds that deter herbivores and inhibit microbial growth.36, 37

Polyacetylenes

Polyacetylenes are compounds containing multiple carbon-carbon triple bonds. They have been identified in some plants as stress-responsive metabolites.

These compounds are involved in defence against herbivores and pathogens and may contribute to the plant’s resistance to abiotic stresses.36, 38

Tannins

Tannins are polyphenolic compounds that can bind and precipitate proteins. They are often involved in plant defence against herbivores by interfering with digestion and nutrient absorption.

Tannins can also have antimicrobial properties and contribute to the plant’s defence against pathogens.36, 39

The biosynthesis of secondary metabolites is tightly regulated and often induced in response to stress signals. Environmental stresses, such as herbivore attack, pathogen infection, UV radiation, and nutrient deficiency, can trigger the activation of specific pathways leading to the production of these metabolites. The accumulation of secondary metabolites is part of the plant’s adaptive response to stress and contributes to its overall resilience and survival.

Osmo-protectants and Water-Use Efficiency

Osmo-protectants and water-use efficiency play pivotal roles in mitigating the impact of plant stress. Osmo-protectants are compounds that help plants maintain cellular structure and function under adverse conditions, such as drought or salinity. They act as protective agents, safeguarding cells from damage caused by water loss and osmotic stress.40

Water-use efficiency, on the other hand, refers to the plant’s ability to optimize water consumption while sustaining growth and development. Plants employ various strategies to enhance water-use efficiency, including the regulation of stomatal openings to minimize water loss through transpiration.41

In the context of plant stress, understanding the interplay between osmoprotectants and water-use efficiency is crucial for developing strategies to enhance plant resilience. Research in this area focuses on identifying key osmoprotectants and mechanisms that contribute to improved water-use efficiency, ultimately aiding in the development of stress-tolerant crops and sustainable agricultural practices.42

Heat Shock Proteins (HSPs) and Chaperones

Heat Shock Proteins (HSPs) and chaperones are essential components of a plant’s stress response, particularly during exposure to elevated temperatures or other environmental stresses. HSPs are a diverse group of molecular chaperones that assist in protein folding, assembly, and transport under normal conditions. However, their expression significantly increases in response to stress, serving as a protective mechanism for the plant.43

During stress, HSPs play a crucial role in preventing protein denaturation and aggregation, ensuring the maintenance of cellular protein homeostasis. They assist in refolding damaged proteins and, in some cases, facilitate the degradation of irreversibly damaged proteins.43

Chaperones, working in concert with HSPs, are molecular assistants that aid in the proper folding of newly synthesized proteins and the refolding of misfolded or denatured proteins during stress. Together, HSPs and chaperones contribute to the stabilization of cellular proteins, maintaining cellular functions and preventing stress-induced damage.43

Research in this field aims to unravel the specific roles of different HSPs and chaperones, their regulation, and their interactions with other cellular components. Understanding these mechanisms is essential for developing strategies to enhance plant stress tolerance, improve crop yields, and ensure food security in the face of changing environmental conditions.

Genetic and Epigenetic Regulation

Genetic and epigenetic regulation are fundamental processes that plants employ to cope with environmental stressors. Genetic regulation involves the activation or suppression of specific genes in response to stress, while epigenetic regulation involves modifications to the structure of DNA or associated proteins without altering the underlying genetic code.

Genetic Regulation

During plant stress, the expression of certain stress-responsive genes is modulated to produce proteins that help the plant adapt. These genes may be involved in the synthesis of protective compounds, the activation of signalling pathways, or the regulation of stress-related processes. Genetic regulation is crucial for the plant’s ability to mount a rapid and targeted response to stress conditions.44

Epigenetic Regulation

Epigenetic changes, such as DNA methylation and histone modifications, play a key role in modulating gene expression without altering the underlying DNA sequence. These changes can be heritable and influence how genes are activated or repressed in response to stress. Epigenetic modifications serve as a memory system for the plant, allowing it to “remember” previous encounters with stress and adjust its response accordingly.44

Research in this area explores the specific genes and pathways involved in the genetic response to stress, as well as the epigenetic modifications that contribute to stress memory and adaptation. Understanding the interplay between genetic and epigenetic regulation provides insights into the molecular mechanisms underlying plant stress responses and can inform strategies for developing stress-tolerant crops through targeted genetic or epigenetic engineering.

Cross-Talk Between Pathways

The cross-talk between pathways emerges as a pivotal and intricate aspect of plant stress responses, orchestrating a sophisticated network of interactions to optimize the adaptation of plants to challenging environmental conditions. Scientific evidence substantiates the existence of cross-talk between various signalling pathways, providing a nuanced understanding of the interconnected molecular processes involved in plant stress responses.23, 45

Studies have elucidated the dynamic interplay between hormonal signalling pathways, such as ABA, JA, and SA, revealing both synergistic and antagonistic interactions that finely tune the plant’s defence mechanisms. Additionally, cross-talk between these hormonal pathways and other signalling components, such as ROS and MAPK cascades, adds layers of complexity to the regulatory network governing stress responses.

The evidence gleaned from molecular biology, genomics, and proteomics approaches underscores the significance of cross-talk in shaping the multifaceted plant stress response. By unravelling these intricate interactions, researchers can pinpoint key nodes and regulatory points, offering potential targets for engineering crops with enhanced stress tolerance.

As the field continues to advance, a more comprehensive understanding of cross-talk between pathways during plant stress will not only deepen our knowledge of fundamental plant biology but also pave the way for innovative strategies in crop improvement. Harnessing this knowledge holds the promise of developing resilient crop varieties capable of withstanding diverse environmental stresses, contributing to global food security in the face of a changing climate.13, 14, 21, 23, 24

Impact on Bioactive Compound Production

The impact of environmental stress on bioactive compound production in plants is a subject of significant scientific interest, with evidence highlighting the dynamic modulation of secondary metabolites in response to stress conditions. Bioactive compounds, such as antioxidants, phenolics, and terpenoids, are not only crucial for the plant’s defence mechanisms but also contribute to the nutritional and medicinal value of crops.

Antioxidant Production

Scientific studies demonstrate that environmental stress, such as drought, salinity, and high temperatures, often leads to an increased production of antioxidants in plants. Antioxidants, including flavonoids and polyphenols, serve as protective molecules by scavenging reactive oxygen species (ROS) generated during stress. This response helps mitigate oxidative damage and maintain cellular integrity.7, 8, 9, 10, 17, 22, 45

Phytochemical Accumulation

Various stresses can induce the accumulation of specific phytochemicals with potential health benefits. For instance, exposure to UV radiation has been shown to stimulate the production of flavonoids in some plants, which not only protect against UV-induced damage but also contribute to the plant’s adaptability to environmental challenges.2, 34, 46

Hormonal Regulation of Metabolites

Hormonal signalling pathways, particularly abscisic acid (ABA) and jasmonic acid (JA), play pivotal roles in orchestrating the plant’s response to stress and influencing the production of bioactive compounds. Scientific evidence suggests that the crosstalk between these hormonal pathways and the regulation of secondary metabolism contributes to the synthesis of compounds with protective and signalling functions.7, 9, 10, 24, 48

Altered Terpenoid Profiles

Certain stress conditions, such as herbivory or pathogen attack, can lead to alterations in the production of terpenoids. Terpenoids, including essential oils, contribute to plant defence against herbivores and have diverse ecological roles. Scientific studies have explored how stress-induced changes in terpenoid profiles can influence plant interactions with other organisms.1, 3, 36, 37, 38

Understanding the impact of stress on bioactive compound production is not only relevant for elucidating plant adaptation mechanisms but also holds implications for human health, agriculture, and the pharmaceutical industry. By deciphering the molecular mechanisms involved, researchers can potentially manipulate these pathways to enhance the production of bioactive compounds in crops, resulting in plants with improved nutritional and medicinal properties.

Technological Advances in Studying Stress Responses

Technological advances have significantly transformed our ability to study stress responses in plants, providing researchers with powerful tools to unravel the intricate molecular mechanisms underlying plant adaptation to environmental challenges. Here’s a brief overview of some key technological advances and their scientific contributions:

Omics Technologies

Genomics

High-throughput sequencing technologies have enabled a comprehensive analysis of plant genomes, facilitating the identification of stress-responsive genes and genomic variations associated with stress tolerance.14, 16, 43, 44, 49

Transcriptomics

RNA sequencing (RNA-seq) allows researchers to study global changes in gene expression during stress, providing insights into the regulatory networks governing plant responses.14, 16, 43, 44, 49

Proteomics

Mass spectrometry-based proteomics enables the identification and quantification of proteins involved in stress responses, revealing dynamic changes in the plant proteome.14, 16, 43, 44, 49

Metabolomics

Advanced analytical techniques, such as mass spectrometry and nuclear magnetic resonance spectroscopy, have enhanced the profiling of plant metabolites. This has led to a deeper understanding of stress-induced changes in the metabolome, including the production of bioactive compounds.38, 50, 51, 52

Advanced Imaging Techniques

Fluorescence Microscopy

Live-cell imaging using fluorescent markers allows real-time visualization of cellular processes, including stress-related events such as reactive oxygen species (ROS) dynamics and ion fluxes.21, 53, 54

Confocal and Super-resolution Microscopy

These techniques provide high-resolution imaging, enabling detailed studies of subcellular structures and protein localization during stress.21, 53, 54

CRISPR/Cas9 Genome Editing

CRISPR/Cas9 Technology

This revolutionary gene-editing tool enables precise modification of plant genomes, allowing researchers to investigate the functions of specific genes involved in stress responses. CRISPR/Cas9 has facilitated the development of stress-tolerant crop varieties.43, 55, 56

Remote Sensing and Phenotyping

Remote Sensing

Satellite and drone-based remote sensing technologies provide real-time monitoring of plant health and stress at the field level, offering insights into the spatial and temporal dynamics of stress responses.57

High-throughput Phenotyping

Automated phenotyping platforms allow the rapid and non-destructive assessment of plant traits, facilitating the study of stress-induced changes in morphology, physiology, and growth.57

Computational Biology

Bioinformatics

Advanced computational tools and algorithms are employed to analyse large-scale omics data, uncover patterns, and identify key regulatory elements in stress-responsive pathways.58, 59

These technological advances, supported by scientific evidence, have revolutionized our ability to dissect the molecular and physiological aspects of plant stress responses. The integration of these tools has led to a more comprehensive understanding of stress adaptation mechanisms and has opened new avenues for the development of stress-tolerant crops and sustainable agricultural practices.

Bioactive Compounds

Plants that grow under stress conditions often produce a variety of bioactive compounds as part of their adaptive responses. These compounds play a crucial role in the plant’s defence mechanisms and can have potential benefits for human health. Plants synthesize these bioactive compounds in response to various stressors, such as drought, high salinity, extreme temperatures, and pathogen attacks. The presence of these compounds in plant-based foods can have positive implications for human nutrition and health, making them potentially valuable in the development of functional foods and nutraceuticals.2 Here’s a brief overview of some key bioactive compounds found in plants under stress:

Introduction to Bioactive Compounds

Definition and Significance

Plants exposed to stress conditions, such as environmental challenges or external threats, often produce a diverse array of bioactive compounds as part of their adaptive strategies. These compounds serve various functions, including defence mechanisms against pests, pathogens, and environmental stressors. Bioactive compounds in stressed plants are of interest not only for their ecological roles but also for their potential applications in human health and nutrition. These compounds can include antioxidants, antimicrobials, and other molecules that contribute to the plant’s resilience. Studying and harnessing these bioactive compounds offers opportunities for developing stress-tolerant crops and discovering novel compounds with therapeutic or nutritional benefits for humans.2, 53, 60, 61, 62

Classes of Bioactive Compounds

Alkaloids

Alkaloids are a group of naturally occurring organic compounds that contain basic nitrogen atoms. Many plants produce alkaloids as secondary metabolites, and their production can be influenced by various environmental stress conditions.36 Here’s a brief introduction to alkaloids in plants under stress.

Role of Alkaloids in Stress Response

Research has demonstrated that alkaloid production in plants can increase in response to stressors such as herbivory, pathogen attacks, and environmental stress.63 Nicotiana spp studies involving tobacco plants, alkaloid levels, including nicotine, have been shown to increase in response to herbivore attacks, serving as a chemical defence mechanism.64

Antimicrobial Properties

Alkaloids often exhibit antimicrobial properties, and their synthesis can be induced in plants under pathogenic stress conditions.65 Quinine, an alkaloid found in the bark of Cinchona trees, has long been used for its antimalarial properties and is produced by the plant in response to stressors, including fungal infections.61, 62

Osmotic Stress Response

Some alkaloids, like betaines, function as osmoprotectants, helping plants cope with osmotic stress caused by factors such as drought or high salinity. Glycine betaine, an alkaloid-like compound, has been identified in various plant species and is associated with enhanced tolerance to abiotic stress conditions.66, 67, 68

Induction by Abiotic Stress

Studies indicate that alkaloid biosynthesis in plants can be influenced by abiotic stress factors, including temperature extremes and nutrient deficiencies. Alkaloid production in medicinal plants like opium poppy (Papaver somniferum) is known to be influenced by environmental factors, with stress conditions potentially enhancing alkaloid content.69, 70

Scientific evidence supports the idea that alkaloid production in plants is a dynamic and adaptive response to various stress conditions, contributing to the plant’s defence mechanisms. These alkaloids not only play a role in plant protection but also have implications for human uses, such as in medicine and agriculture.

Flavonoids

Flavonoids are a diverse group of secondary metabolites found in plants, and their production can be influenced by environmental stress conditions. Here’s a brief introduction to flavonoids in plants under stress.

Role of Flavonoids in Stress Response

Numerous studies have demonstrated that plants increase the synthesis of flavonoids in response to various stressors, including UV radiation, pathogen attacks, and drought. In a study on grapevine (Vitis vinifera), exposure to UV-B radiation led to a significant increase in the accumulation of flavonoids, particularly anthocyanins, which act as protective pigments against UV damage.2, 34, 46, 71

Antioxidant Properties

Flavonoids are known for their antioxidant activity, helping plants combat oxidative stress caused by environmental factors. This antioxidant capacity has been linked to the presence of hydroxyl groups in the flavonoid structure. Quercetin, a flavonoid commonly found in various plant species, has been shown to scavenge free radicals and protect plant cells from oxidative damage induced by stress.7, 8, 9, 10, 17, 18, 21, 22, 45

UV Protection and Photoprotection

Flavonoids, particularly flavonols, contribute to UV protection by absorbing UV-B and UV-A radiation. They act as photoprotective agents, preventing damage to cellular components. Studies on Arabidopsis thaliana have shown that flavonoid accumulation increases under UV-B exposure, providing photoprotection and reducing DNA damage.2, 34, 46, 71

Pathogen Defence

Flavonoids play a role in plant defence against pathogens by inhibiting the growth of bacteria and fungi. Their production is often induced by infection. Flavonoids such as catechins and isoflavonoids have been shown to have antimicrobial properties, contributing to the plant’s defence against various pathogens.2, 53, 60, 61, 62

Scientific evidence supports the notion that flavonoids in plants are not only products of stress responses but also active participants in plant adaptation and defence mechanisms. Their diverse functions, including antioxidant activity, UV protection, and pathogen defence, make flavonoids crucial components of the plant’s response to stress conditions.

Terpenoids Compounds

Terpenoids, also known as isoprenoids, constitute a diverse class of secondary metabolites found in plants. Their production can be influenced by various stress conditions, and they play crucial roles in plant adaptation and defence mechanisms. Here’s a brief introduction to terpenoids in plants under stress.

Role of Terpenoids in Stress Response

Research has demonstrated that plants often increase the synthesis of terpenoids in response to environmental stress, including herbivore attacks, pathogen infections, and abiotic stressors like drought and high temperatures. In a study on sagebrush (Artemisia tridentata), exposure to drought stress resulted in increased terpenoid production, potentially serving as a defence against herbivores and contributing to the plant’s ability to withstand water scarcity.1, 3, 35, 36, 37, 38

Herbivore Deterrence and Defence

Terpenoids play a key role in deterring herbivores and defending plants against insect attacks. They can act as repellents, antifeedants, or even toxins against herbivores. The emission of volatile terpenoids by pine trees (Pinus spp.) has been shown to deter herbivorous insects, and the composition of emitted terpenoids can change in response to herbivore presence.1, 3, 9, 10, 34, 35, 36, 37, 38

Antimicrobial Properties

Certain terpenoids exhibit antimicrobial properties, contributing to the plant’s defence against pathogens. They can inhibit the growth of bacteria and fungi. Essential oils, rich in terpenoids, have been demonstrated to have antimicrobial activity. For example, tea tree oil from Melaleuca alternifolia contains terpenoids like terpinen-4-ol with antimicrobial properties.2,9, 10, 36, 39, 53, 60, 61, 62

Abiotic Stress Tolerance

Some terpenoids act as osmoprotectants, helping plants tolerate abiotic stress conditions such as high salinity and extreme temperatures. In studies on lavender (Lavandula angustifolia), the synthesis of terpenoids, including essential oils, increased under conditions of water deficit, suggesting a role in drought tolerance.7, 8, 27, 36, 37, 38, 67

Scientific evidence supports the idea that terpenoids in plants are dynamic molecules that respond to stress conditions by participating in defence mechanisms against herbivores and pathogens, as well as contributing to abiotic stress tolerance. Their diverse functions make terpenoids essential players in the complex interplay between plants and their environment.

Phenolic compounds

Phenolic compounds are a diverse group of secondary metabolites found in plants, and their production can be influenced by various stress conditions. These compounds play significant roles in plant adaptation and defence mechanisms. Here’s a brief introduction to phenolic compounds in plants under stress, supported by scientific evidence:

Role of Phenolic Compounds in Stress Response

Research has consistently shown that plants increase the synthesis of phenolic compounds in response to environmental stress, such as drought, high salinity, and exposure to pathogens. Studies on grapevine (Vitis vinifera) have demonstrated an increase in phenolic compound levels, including flavonoids, in response to drought stress. These compounds contribute to the plant’s ability to cope with water scarcity.7, 8, 9, 10, 17, 22, 45

Antioxidant Properties

Phenolic compounds, especially flavonoids and polyphenols, are known for their antioxidant activity. They help plants combat oxidative stress induced by environmental factors. Quercetin, a flavonoid with antioxidant properties, has been identified in various plant species and has been shown to scavenge free radicals, protecting plant cells from oxidative damage caused by stress.7, 8, 9, 10, 17, 18, 21, 22, 45, 72

Pathogen Defence

Phenolic compounds contribute to plant defence against pathogens by inhibiting the growth of bacteria and fungi. Their production is often induced by infection. In response to pathogen attacks, plants such as wheat (Triticum aestivum) produce phenolic compounds like hydroxycinnamic acids as part of their defence mechanism against fungal pathogens.9, 10, 11, 12, 26, 73, 74

UV Protection

Phenolic compounds, particularly flavonoids, can act as sunscreens, protecting against UV radiation. Their production is often induced by exposure to UV light. Arabidopsis thaliana plants subjected to UV-B radiation show an increase in the accumulation of phenolic compounds, contributing to photoprotection and reducing damage caused by UV stress.2, 34, 46, 71, 75

Scientific evidence supports the idea that phenolic compounds in plants play crucial roles in response to stress conditions, contributing to antioxidant defence, protection against pathogens, and adaptation to environmental challenges. The diverse functions of phenolic compounds make them integral components of the plant’s stress response mechanisms.

Stress-Induced Changes in Bioactive Compounds

Stress-induced changes in bioactive compounds in plants are complex and varied, reflecting the plants’ adaptive responses to environmental challenges. These changes can include alterations in the levels of secondary metabolites such as phenolic compounds, alkaloids, terpenoids, and other bioactive substances.7, 8, 9, 10, 17, 22, 45

Phenolic Compounds

Studies have shown that various stress conditions, including drought, salinity, and pathogen attacks, can lead to increased production of phenolic compounds in plants. For example, in wheat plants, drought stress induces the accumulation of phenolic acids, contributing to the antioxidant defence and stress tolerance.7, 8, 9, 10, 17, 22, 45, 76

Alkaloids

Alkaloid levels often increase in response to stressors such as herbivory and pathogen attacks. For instance, in tobacco plants, alkaloid production, including nicotine, is enhanced in response to herbivore feeding, acting as a chemical defence mechanism.7, 8, 9, 10, 17, 22, 45, 64

Terpenoids

Terpenoid synthesis is frequently upregulated under stress conditions. For example, in sagebrush exposed to drought stress, there is an increase in terpenoid production, potentially contributing to the plant’s defence against herbivores and aiding in water stress tolerance.7, 8, 9, 10, 17, 22, 45, 77

Flavonoids

Flavonoid levels often rise in response to various stressors, such as UV radiation, drought, and pathogen attacks. In grapevines, exposure to UV-B radiation increases flavonoid, particularly anthocyanin, production, protecting against UV damage.7, 8, 9, 10, 17, 22, 45, 78

Oxidative Stress Responses

Environmental stressors can induce oxidative stress in plants. In response, plants increase the production of antioxidant bioactive compounds, such as polyphenols and flavonoids, to counteract the effects of reactive oxygen species (ROS) generated under stress conditions.7, 8, 9, 10, 17, 22, 45

Hormonal Regulation

Stress-induced changes in bioactive compounds are often mediated by plant hormones. For instance, abscisic acid (ABA) plays a crucial role in regulating the biosynthesis of stress-related compounds like flavonoids and terpenoids under drought conditions.7, 8, 9, 10, 17, 22, 45, 79

Stress-induced changes in bioactive compounds represent dynamic responses of plants to environmental challenges. These changes not only contribute to the plant’s survival and adaptation but also have implications for human uses, such as the development of stress-tolerant crops and the discovery of bioactive compounds with potential applications in medicine and nutrition.

Adaptive Strategies for Bioactive Compound Production

Plants employ adaptive strategies for the production of bioactive compounds under stress conditions, enhancing their survival and defence mechanisms. These strategies involve the upregulation of specific biochemical pathways leading to the synthesis of secondary metabolites.

Phenolic Compounds

Phenolic compounds, including flavonoids and polyphenols, act as antioxidants and defence molecules. Under stress conditions like drought, studies on grapevines (Vitis vinifera) show increased levels of phenolic compounds, contributing to the antioxidant defence and stress tolerance.7, 8, 9, 10, 17, 22, 34, 36, 39, 45

Alkaloids

Alkaloids serve as chemical defences against herbivores and pathogens. In response to herbivore attacks, such as in tobacco plants (Nicotiana spp.), alkaloid production, including nicotine, is induced, deterring herbivores and protecting the plant.61- 70

Terpenoids

Terpenoids act as deterrents against herbivores and have antimicrobial properties. Sagebrush (Artemisia tridentata) increases terpenoid production under drought stress, potentially aiding in defence against herbivores and enhancing water stress tolerance.7, 8, 9, 10, 17, 22, 36, 45, 46

Flavonoids

Flavonoids contribute to UV protection, antioxidant defence, and pathogen resistance. Under UV-B radiation, Arabidopsis thaliana plants exhibit an increase in flavonoid levels, providing photoprotection and reducing DNA damage.7, 8, 9, 10, 17, 18, 21, 22, 45, 46, 75

Osmoprotectants

Osmoprotectants like betaines help plants cope with osmotic stress. Glycine betaine, a betaine osmoprotectant, is synthesized in response to abiotic stressors, such as drought, contributing to stress tolerance in various plant species 7, 8, 27, 36, 37, 38, 42, 53, 66, 67, 68

Hormonal Regulation

Plant hormones, such as abscisic acid (ABA), mediate the biosynthesis of stress-related compounds. ABA regulates the production of flavonoids and terpenoids under drought conditions, facilitating adaptive responses in plants.7, 8, 9, 10, 17, 22, 23, 45, 79, 80

Antimicrobial Compounds

Some bioactive compounds, including certain alkaloids and terpenoids, have direct antimicrobial properties. Essential oils rich in terpenoids exhibit antimicrobial activity, contributing to the plant’s defence against pathogens.2, 9, 10, 53, 36, 39, 60, 61, 62

These adaptive strategies highlight the versatility of plant responses to stress, enabling them to synthesize bioactive compounds that serve multiple functions, from defence against herbivores and pathogens to protection against environmental stressors. The scientific evidence underscores the importance of these strategies in plant survival and adaptation.

Case Studies and Comparative Analyses

The responses of medicinal plants to stressful environments often involve the synthesis of bioactive compounds, which are secondary metabolites that play a crucial role in the plants’ adaptation and defence mechanisms. Here are some examples of medicinal plants and their bioactive compound responses to stress:

Turmeric (Curcuma longa)

Bioactive Compounds: Curcuminoids, with curcumin being the most prominent.

Stress Response

Turmeric synthesizes curcuminoids, particularly curcumin, as a response to stress. Curcumin is well-known for its anti-inflammatory and antioxidant properties, making turmeric a valuable medicinal herb.81

Holy Basil (Ocimum sanctum):

Bioactive Compounds: Eugenol, ursolic acid, rosmarinic acid.

Stress Response

Holy Basil, or Tulsi, is an adaptogenic herb that produces higher levels of bioactive compounds like eugenol and ursolic acid in response to stress. These compounds contribute to its medicinal properties, including antioxidant and anti-inflammatory effects.82

Aloe Vera (Aloe barbadensis miller)

Bioactive Compounds: Aloin, polysaccharides.

Stress Response

Aloe vera produces higher levels of aloin and polysaccharides in response to environmental stress. These compounds contribute to the plant’s ability to survive in arid conditions and have medicinal properties, including wound healing and anti-inflammatory effects.83

Moringa (Moringa oleifera)

Bioactive Compounds: Quercetin, chlorogenic acid, beta-carotene.

Stress Response

Moringa is known for its nutritional value and produces bioactive compounds like quercetin and chlorogenic acid in response to stress. These compounds contribute to its antioxidant and anti-inflammatory properties.84

Lavender (Lavandula angustifolia):

Bioactive Compounds: Linalool, linalyl acetate.

Stress Response

Lavender synthesizes higher levels of essential oils, including linalool and linalyl acetate, in response to stress. These compounds are responsible for the plant’s aromatic and stress-relieving properties.85

Understanding how these medicinal plants respond to stress by producing specific bioactive compounds provides insights into their adaptation mechanisms and the potential enhancement of their medicinal properties under certain cultivation conditions. Additionally, it highlights the importance of considering environmental factors in optimizing the cultivation of these plants for medicinal use.

Comparisons Across Stress Conditions

Evaluating how different stressors influence the bioactive compound profiles of medicinal plants provides valuable insights into the adaptability of these plants to various environmental challenges. Here’s a more detailed exploration:

Drought Stress

Influence on Bioactive Compounds

Many medicinal plants, such as Aloe vera and Ginseng, respond to drought stress by increasing the synthesis of specific compounds like polysaccharides and ginsenosides. These compounds play a role in water retention, osmotic regulation, and overall stress tolerance.1, 6, 7, 19, 86

Heat Stress

Influence on Bioactive Compounds

Heat stress can lead to the production of secondary metabolites with antioxidant properties. For example, Curcuma domistica (Turmeric) responds to heat stress by increasing the concentration of curcuminoids, enhancing its antioxidant and anti-inflammatory effects.1, 43

Pathogen or Pest Infestation

Influence on Bioactive Compounds

Medicinal plants often respond to pathogen or pest attacks by producing compounds with antimicrobial properties. For instance, Holy Basil increases the production of eugenol in response to pest stress, contributing to its insect-repelling and medicinal qualities.9, 10, 11, 12, 37, 43, 69

Salinity Stress

Influence on Bioactive Compounds

Plants exposed to high salinity may produce bioactive compounds like flavonoids and phenolics, contributing to antioxidant defences. This adaptation is observed in plants like Rhodiola, which may enhance the synthesis of rosavin and salidroside.1, 7, 12, 19, 43, 55, 87

Nutrient Deficiency

Influence on Bioactive Compounds

Nutrient deficiencies can lead to altered bioactive compound profiles. For example, Moringa subjected to nutrient stress may exhibit changes in the levels of quercetin and chlorogenic acid, impacting its overall nutritional and medicinal content.30, 31, 69, 70, 88

Light Stress

Influence on Bioactive Compounds 

Light stress, such as exposure to UV radiation, can stimulate the production of protective compounds. Reishi mushrooms respond to light stress by increasing the levels of triterpenes and polysaccharides, potentially enhancing their medicinal properties.2, 34, 46

Understanding these stress-induced changes in bioactive compound profiles is crucial for several reasons:

Medicinal Efficacy

Different stressors can impact the concentration and types of bioactive compounds in medicinal plants. Understanding these variations is essential for ensuring the consistent medicinal efficacy of plant-derived products.3, 34, 60, 62, 81, 83, 89

Cultivation Strategies

Insights into how medicinal plants adapt to stress can guide cultivation practices. Farmers and cultivators can optimize growing conditions to promote the synthesis of specific bioactive compounds, enhancing the overall quality of the medicinal plant.90, 91, 92

Pharmacological Applications 

Knowledge of stress-induced changes in bioactive compounds can inform pharmacological research. Researchers can explore stress-induced variations to develop more potent and targeted medicinal products.3, 34, 60, 62, 81, 83, 89

Oxidative Stress

Influence on Bioactive Compounds

Exposure to oxidative stress, often induced by environmental pollutants, can lead to increased production of antioxidants. Plants like Green Tea (Camellia sinensis) respond by synthesizing higher levels of catechins, particularly epigallocatechin gallate (EGCG), which has potent antioxidant properties.2, 7, 8, 9, 10, 17, 21, 22, 28, 29, 34, 45, 46

Cold Stress

Influence on Bioactive Compounds

Cold stress prompts plants to produce cryoprotective compounds. Echinacea (Echinacea purpurea), for instance, responds to cold stress by increasing the concentration of alkamides and polysaccharides, potentially enhancing its immune-stimulating effects.2, 7, 8, 9, 10, 17, 22, 30, 31, 34, 45, 46

Heavy Metal Stress

Influence on Bioactive Compounds

Plants exposed to heavy metal stress may produce chelating compounds to detoxify and mitigate the effects. Coriander (Coriandrum sativum) responds to heavy metal stress by increasing the production of coriandrin and other bioactive compounds with potential detoxification properties.2, 7, 8, 9, 10, 17, 22, 34, 45, 46, 93

Waterlogging Stress

Influence on Bioactive Compounds

Waterlogged conditions can induce changes in the bioactive compound profile. Ginkgo biloba responds to waterlogging stress by altering the levels of ginkgolides and bilobalide, which are believed to play a role in stress adaptation.2, 7, 8, 9, 10, 17, 22, 34, 45, 46, 94

Air Pollution Stress

Influence on Bioactive Compounds

Exposure to air pollution can lead to the production of compounds with antioxidant and detoxifying properties. Rosemary (Rosmarinus officinalis) responds to air pollution stress by increasing the concentration of rosmarinic acid and other antioxidants.2, 7, 8, 9, 10, 17, 22, 34, 45, 46, 95

Fungal Infections

Influence on Bioactive Compounds

Medicinal plants facing fungal infections may enhance the production of antifungal compounds. For example, Garlic (Allium sativum) responds to fungal stress by increasing allicin content, known for its antimicrobial properties. 2, 7, 8, 9, 10, 11, 12, 17, 22, 26, 34, 45, 46, 73, 74, 96

Understanding these nuanced responses to various stressors provides a comprehensive view of the biochemical adaptations of medicinal plants. It also emphasizes the dynamic nature of plant secondary metabolism and the potential for eliciting specific bioactive compounds under different environmental conditions. This knowledge is instrumental for sustainable cultivation practices, pharmaceutical development, and maximizing the therapeutic potential of medicinal plants in diverse ecological settings.

Integration With Medicinal Potential

Linking Bioactive Compounds to Medicinal Properties

The stress-induced changes in bioactive compounds in medicinal plants are closely tied to potential enhancements in their medicinal properties. These adaptations represent the plant’s response to environmental challenges and can result in the synthesis of compounds that contribute to the plant’s resilience and therapeutic efficacy.2, 7, 8, 9, 10, 11, 12, 17, 22, 38, 45, 50, 51, 52, 53, 60, 61, 62   Here’s a closer look at how these changes connect to enhancements in medicinal properties:

Increased Antioxidant Content

Connection to Medicinal Properties

Stressors such as UV radiation, air pollution, and oxidative stress can trigger the production of antioxidants in medicinal plants. Higher levels of antioxidants, like polyphenols in Green Tea (Camellia sinensis), contribute to enhanced medicinal properties by providing increased protection against oxidative damage, supporting overall health, and potentially preventing chronic diseases.2, 7, 8, 9, 10, 17, 21, 22, 28, 29, 34, 45, 46

Elevated Antimicrobial Compounds

Connection to Medicinal Properties

Fungal infections and pest infestations often lead to the production of antimicrobial compounds in medicinal plants. For example, Garlic (Allium sativum) responds to fungal stress by increasing allicin content, contributing to enhanced antimicrobial and immune-boosting properties.2, 7, 8, 9, 10, 11, 12, 17, 22, 26, 34, 45, 46, 73, 74, 96

Adaptogenic and Stress-Relieving Compounds

Connection to Medicinal Properties:

Plants like Ginseng and Rhodiola respond to various stressors by producing adaptogenic compounds, such as ginsenosides and rosavin. These compounds are associated with stress relief, immune modulation, and improved resilience. When consumed, they may contribute to the plant’s traditional uses for supporting overall well-being and mitigating the effects of stress-related conditions.1, 7, 12, 19, 43, 55, 87

Detoxification Compounds

Connection to Medicinal Properties

Heavy metal stress and pollution can induce the synthesis of detoxifying compounds in medicinal plants. Coriander (Coriandrum sativum), for instance, responds to heavy metal stress by increasing the production of coriandrin. This compound may contribute to the plant’s medicinal properties by assisting in the detoxification process when consumed.2, 7, 8, 9, 10, 17, 22, 34, 45, 46, 93

Enhanced Immune-Stimulating Compounds

Connection to Medicinal Properties

Cold stress in plants like Echinacea can lead to an increase in immune-stimulating compounds such as alkamides and polysaccharides. These compounds contribute to the plant’s traditional use for immune support and can potentially enhance the medicinal properties of Echinacea-based products.2, 7, 8, 9, 10, 17, 22, 30, 31, 34, 45, 46

Optimized Secondary Metabolism

Connection to Medicinal Properties

Stress-induced changes in bioactive compounds often reflect a redirection of the plant’s secondary metabolism. This can result in an optimized balance of specific compounds, such as terpenes, alkaloids, and flavonoids, which collectively contribute to the medicinal properties of the plant. For example, Ginkgo biloba responds to waterlogging stress by altering the levels of ginkgolides and bilobalide, which are associated with cognitive and circulatory benefits.2, 7, 8, 9, 10, 11, 12, 17, 22, 38, 45, 50, 51, 52, 53, 60, 61, 62

Understanding these connections is vital for practitioners of traditional medicine, herbalists, and researchers aiming to harness the full potential of medicinal plants. It also guides sustainable cultivation practices that promote stress-induced adaptations, resulting in plants with optimal medicinal properties. Additionally, this knowledge informs the development of standardized herbal preparations and pharmaceuticals, ensuring consistency and efficacy in medicinal plant-based products.

Consideration of Traditional Uses

The examination of stress-induced alterations in medicinal plants and their alignment or deviation from traditional uses is crucial for understanding the dynamic nature of these plants’ biochemistry and their potential impact on therapeutic efficacy.1, 3, 5, 36, 37, 38 Here’s a closer look at how stress-induced changes may align with or deviate from traditional uses:

Alignment with Traditional Uses

Enhanced Bioactive Compounds

Stress-induced changes that lead to an increase in bioactive compounds associated with traditional therapeutic effects can align with traditional uses. For example, if a medicinal plant traditionally used for its anti-inflammatory properties responds to stress by increasing the production of anti-inflammatory compounds, this aligns with its traditional application.1, 3,5, 7, 8, 9, 10, 17, 22, 36, 37, 38, 43, 45, 50, 51, 52, 57, 79

Optimization of Medicinally Relevant Compounds

Stress-induced alterations that optimize the concentration of medicinally relevant compounds can align with traditional uses. Traditional herbal knowledge often emphasizes specific parts of plants or certain growth conditions for optimal medicinal properties. Stress-induced adaptations may naturally enhance the concentration of these compounds, aligning with traditional practices.1, 3,5, 7, 8, 9, 10, 17, 22, 36, 37, 38, 43, 45, 50, 51, 52, 57, 79

Adaptogenic Responses 

Stress-induced changes leading to the production of adaptogenic compounds can align with traditional uses of medicinal plants known for their adaptogenic effects. For example, if a plant traditionally used to combat stress and promote overall well-being responds to environmental stress by increasing adaptogenic compounds, it reinforces its traditional application.1, 3,5, 7, 8, 9, 10, 17, 22, 36, 37, 38, 43, 45, 50, 51, 52, 57, 79

Deviation from Traditional Uses

Changes in Bioactive Composition

 If stress-induced changes result in a significant alteration in the bioactive composition, especially if it introduces compounds not traditionally associated with the plant’s medicinal properties, it may deviate from traditional uses. This could impact the overall therapeutic effects and safety of the plant.1, 3,5, 7, 8, 9, 10, 17, 22, 36, 37, 38, 43, 45, 50, 51, 52, 57, 79

Shifts in Secondary Metabolism

Stress may lead to shifts in secondary metabolism, potentially changing the balance of different bioactive compounds. If these shifts deviate from the traditionally recognized composition of the plant, it may impact its therapeutic profile and how it is traditionally used.1, 3,5, 7, 8, 9, 10, 17, 22, 36, 37, 38, 43, 45, 50, 51, 52, 57, 79

Loss of Medicinally Relevant Compounds

In some cases, stress-induced alterations might lead to a decrease or loss of certain medicinally relevant compounds. If these compounds are central to the traditional use of the plant, such deviations may impact its efficacy and alignment with traditional practices.1, 3,5, 7, 8, 9, 10, 17, 22, 36, 37, 38, 43, 45, 50, 51, 52, 57, 79

Environmental Contaminants

Stress-induced responses to environmental challenges, such as pollution or heavy metal stress, might introduce contaminants that deviate from the traditional purity standards. This can impact the safety and effectiveness of the plant for traditional uses.1, 3,5, 7, 8, 9, 10, 17, 22, 36, 37, 38, 43, 45, 50, 51, 52, 57, 79

Consideration of Ecotypes and Cultivars

Natural Variation

Different ecotypes or cultivars of a medicinal plant may respond differently to stress, introducing natural variations in bioactive compound profiles. Traditional uses might be rooted in the specific ecotype or cultivar, and deviations may arise when different variants respond uniquely to stressors.62, 97

During stress-induced alterations can align with traditional uses by enhancing relevant bioactive compounds and adaptogenic responses, they can also deviate by introducing new compounds or changing the balance of existing ones. It is essential to consider these changes in the context of traditional knowledge, taking into account the specific plant varieties, growth conditions, and traditional preparation methods. Additionally, scientific research can help validate and refine traditional practices in light of stress-induced adaptations in medicinal plants.62, 97, 98

Conclusion

Exploring the biochemical compositions of medicinal plants cultivated in demanding environments offers a nuanced understanding of the intricate interplay between these plants and their surroundings. These plants exhibit varied responses to stressors, resulting in complex changes in their bioactive compound profiles that influence adaptability and potential medicinal properties.

The alterations in bioactive compounds induced by stress serve as a reflection of the plants’ adaptive strategies, showcasing their capacity to flourish and endure in adverse conditions. This adaptability assumes particular significance in traditional medicine, where the efficacy of medicinal plants is deeply rooted in centuries-old practices and observations.

The alignment or diversion of stress-induced changes from traditional uses underscores the intricate interplay between environmental factors and the traditional knowledge surrounding medicinal plants. While some stress-induced alterations may fortify and augment the traditional applications of these plants, others may introduce variations that necessitate careful consideration.

The optimization of specific bioactive compounds, the synthesis of adaptogenic responses, and the potential enhancement of therapeutic properties underscore the resilience of medicinal plants under stress. These discoveries bear practical implications for sustainable cultivation practices, pharmaceutical development, and the formulation of herbal remedies.

Nevertheless, a cautious approach is imperative when integrating stress-induced changes into traditional medicine. Deviations from established traditional uses may introduce uncertainties regarding the safety and efficacy of medicinal plants. Rigorous scientific research, in conjunction with traditional knowledge, is essential to validate and refine our comprehension of these complex relationships.

Exploring the biochemical profiles of medicinal plants in stressful environments not only enhances our understanding of plant biology but also provides avenues for harnessing the full therapeutic potential of these valuable resources. This interdisciplinary approach, marrying traditional wisdom with modern scientific inquiry, is key to unravelling the intricate mechanisms by which medicinal plants adapt, thrive, and contribute to human health in diverse ecological contexts.

Future Perspectives

Medicinal plants, despite being a rich source of bioactive compounds, are not fully explored, with only a small fraction of their properties investigated. Many medicinal plants remain untapped, offering crucial natural resources for discovering novel compounds that modify resistance and could serve as valuable therapeutic tools. A significant portion of these natural compound mixtures, though complex, await thorough chemical investigation, presenting a resource with substantial potential for further scientific exploration. Subsequent steps may involve in vivo testing in animal models of infection to ascertain the clinical relevance of these compounds and establish a valid correlation with in vitro efficacy results.

Prospective investigations should encompass the exploration of structural modifications in compounds aimed at optimizing pharmacokinetic and pharmacodynamic properties. Additionally, a comprehensive analysis of structure-activity relationships is warranted. The exploration of synergistic interactions within medicinal plant extracts, as well as interactions between compounds and antibiotics, is indispensable to elucidate the mechanisms underpinning the antimicrobial efficacy of these compounds and discern multiple pathways for targeted intervention. It is crucial to emphasize that interactions between medicinal plant extracts and antimicrobial agents can manifest as either advantageous, such as synergism, or detrimental, exemplified by antagonism. Consequently, further research, particularly through in vivo studies and evaluations of product toxicity, is imperative to substantiate their classification as biomedical agents.

Acknowledgement

The author would like to thank the Department of Botany, J V College, Baraut, for providing the necessary laboratory facilities. The author is also grateful to the Department’s personnel for assisting with several of the analyses.

Funding Sources

This is an academic research initiative. All of the author’s work is self-sponsored; neither the institution nor any outside financial source provides funding or scholarships. 

Conflict of Interest

There is no conflict of interest.

Authors’ Contribution

There is a single author, so no need for a contribution letter.

References

  1. dos Santos, T.B.; Ribas, A.F.; de Souza, S.G.H.; Budzinski, I.G.F.; Domingues, D.S. Physiological Responses to Drought, Salinity, and Heat Stress in Plants: A Review. Stresses. 2022, 2, 113-135. https://doi.org/10.3390/stresses2010009
    CrossRef
  2. Altemimi A, Lakhssassi N, Baharlouei A, Watson DG, Lightfoot DA. Phytochemicals: Extraction, Isolation, and Identification of Bioactive Compounds from Plant Extracts. Plants (Basel). 2017 Sep 22;6(4):42. doi: 10.3390/plants6040042.
    CrossRef
  3. Sofowora A, Ogunbodede E, Onayade A. The role and place of medicinal plants in the strategies for disease prevention. Afr J Tradit Complement Altern Med. 2013 Aug 12;10(5):210-29. doi: 10.4314/ajtcam.v10i5.2.
    CrossRef
  4. Paul D, Sanap G, Shenoy S, Kalyane D, Kalia K, Tekade RK. Artificial intelligence in drug discovery and development. Drug Discov Today. 2021 Jan;26(1):80-93. doi: 10.1016/j.drudis.2020.10.010. Epub 2020 Oct 21.
    CrossRef
  5. Sharma M. K. Plants Stress: Salt Stress and Mechanisms of Stress Tolerance. Curr Agri Res 2023; 11(2). doi : http://dx.doi.org/10.12944/CARJ.11.2.03
    CrossRef
  6. Gull, A., Ahmad Lone, A., & Ul Islam Wani, N. Biotic and Abiotic Stresses in Plants. IntechOpen.2019,  doi: 10.5772/intechopen.85832
    CrossRef
  7. Bousba R., Rached-Kanouni M., Benghersallah N., Djekoune A., Ykhlef N.  Role of exogenous application of abscisic acid ABA in drought tolerance and evaluation of antioxidant activity in durum wheat genotypes. Acta Sci. Nat. 2020, 7 44–60. 10.2478/asn-2020-0019
    CrossRef
  8. Fatma M., Iqbal N., Gautam H., Sehar Z., Sofo A., D’Ippolito I., et al. Ethylene and sulfur coordinately modulate the antioxidant system and ABA accumulation in mustard plants under salt stress. Plants. 2021,  10:180. 10.3390/plants10010180
    CrossRef
  9. Alam M. M., Nahar K., Hasanuzzaman M., Fujita M. . Exogenous jasmonic acid modulates the physiology, antioxidant defense and glyoxalase systems in imparting drought stress tolerance in different Brassica species. Plant Biotechnol. 2014, Rep. 8 279–293. 10.1007/s11816-014-0321-8
    CrossRef
  10. Ghaffari H., Tadayon M. R., Nadeem M., Razmjoo J., Cheema M.  Foliage applications of jasmonic acid modulate the antioxidant defense under water deficit growth in sugar beet. Spanish J. Agric. Res.2020,  17:0805. 10.5424/sjar/2019174-15380
    CrossRef
  11. Biareh V., Shekari F., Sayfzadeh S., Zakerin H., Hadidi E., Beltrão J. G. T., et al. Physiological and qualitative response of Cucurbita pepo L. to salicylic acid under controlled water stress conditions. Horticulturae.2022, 8:79. 10.3390/horticulturae8010079
    CrossRef
  12. Jayakannan M., Bose J., Babourina O., Rengel Z., Shabala S. Salicylic acid improves salinity tolerance in Arabidopsis by restoring membrane potential and preventing salt-induced K+ loss via a GORK channel. J. Exp. Bot. 2013, 64 2255–2268. 10.1093/jxb/ert085
    CrossRef
  13. Alexander Plotnikov, Eldar Zehorai, Shiri Procaccia, Rony Seger. The MAPK cascades: Signaling components, nuclear roles and mechanisms of nuclear translocation, Biochimica et Biophysica Acta (BBA) – Molecular Cell Research,2011. 1813 (9), 1619-1633,https://doi.org/10.1016/j.bbamcr.2010.12.012.
    CrossRef
  14. Cargnello M, Roux PP. Activation and function of the MAPKs and their substrates, the MAPK-activated protein kinases. Microbiol Mol Biol Rev. 2011 Mar;75(1):50-83. doi: 10.1128/MMBR.00031-10.
    CrossRef
  15. Alberts B, Bray D, Hopkin K, Johnson A, Lewis J, Raff MC, et al.  Essential Cell Biology (4th ed.). New York, NY: Garland Science. 2014. pp. 548–549.
    CrossRef
  16. Reddy AS, Ali GS, Celesnik H, Day IS. Coping with stresses: roles of calcium- and calcium/calmodulin-regulated gene expression. Plant Cell. 2011 Jun;23(6):2010-32. doi: 10.1105/tpc.111.084988.
    CrossRef
  17. Mittler, R. Oxidative stress, antioxidants and stress tolerance. Trends in Plant Science, 2002, 7(9), 405-410.
    CrossRef
  18. Tripathy BC, Oelmüller R. Reactive oxygen species generation and signaling in plants. Plant Signal Behav. 2012 Dec;7(12):1621-33. doi: 10.4161/psb.22455.
    CrossRef
  19. Han X, Yang Y. Phospholipids in Salt Stress Response. Plants (Basel). 2021 Oct 17;10(10):2204. doi: 10.3390/plants10102204.
    CrossRef
  20. Bill CA, Vines CM. Phospholipase C. Adv Exp Med Biol. 2020;1131:215-242. doi: 10.1007/978-3-030-12457-1_9.
    CrossRef
  21. Mittler R.  ROS are good. Trends Plant Sci. 2017,22. 11–19. 10.1016/j.tplants.2016.08.002
    CrossRef
  22. Gill, S. S., & Tuteja, N. Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants. Plant Physiology and Biochemistry, 2010. 48(12), 909-930.
    CrossRef
  23. Ku Y.-S., Sintaha M., Cheung M.-Y., Lam H.-M.  Plant hormone signaling crosstalks between biotic and abiotic stress responses. Int. J. Mol. Sci. 2018, 19:3206. 10.3390/ijms19103206
    CrossRef
  24. Tuteja N. Abscisic Acid and abiotic stress signaling. Plant Signal Behav. 2007 May;2(3):135-8. doi: 10.4161/psb.2.3.4156.
    CrossRef
  25. Muhammad Aslam M, Waseem M, Jakada BH, Okal EJ, Lei Z, Saqib HSA, Yuan W, Xu W, Zhang Q. Mechanisms of Abscisic Acid-Mediated Drought Stress Responses in Plants. Int J Mol Sci. 2022 Jan 19;23(3):1084. doi: 10.3390/ijms23031084.
    CrossRef
  26. Diwaker Tripathi , Gaurav Raikhy , Dhirendra Kumar. Chemical elicitors of systemic acquired resistance—Salicylic acid and its functional analogs. Current Plant Biology. 2019, 17, 48-59. https://doi.org/10.1016/j.cpb.2019.03.002
    CrossRef
  27. D’Haeze W., De Rycke R., Mathis R., Goormachtig S., Pagnotta S., Verplancke C., et al.  Reactive oxygen species and ethylene play a positive role in lateral root base nodulation of a semiaquatic legume. PNAS. 2003, 100 11789–11794. 10.1073/pnas.1333899100
    CrossRef
  28. Chang Z., Liu Y., Dong H., Teng K., Han L., Zhang X.  Effects of cytokinin and nitrogen on drought tolerance of creeping bent grass. PLoS One . 2016, 11:e0154005. 10.1371/journal.pone.0154005
    CrossRef
  29. Hai NN, Chuong NN, Tu NHC, Kisiala A, Hoang XLT, Thao NP. Role and Regulation of Cytokinins in Plant Response to Drought Stress. Plants (Basel). 2020 Mar 31;9(4):422. doi: 10.3390/plants9040422.
    CrossRef
  30. Hedden P. Gibberellin biosynthesis in higher plants. Ann. Plant Rev. 2018, 49 37–71. 10.1002/9781119312994.apr0531
    CrossRef
  31. Weiss D, Ori N. Mechanisms of cross talk between gibberellin and other hormones. Plant Physiol. 2007 Jul;144(3):1240-6. doi: 10.1104/pp.107.100370.
    CrossRef
  32. Hafeez M. B., Zahra N., Zahra K., Raza A., Khan A., Shaukat K., et al.  Brassinosteroids: molecular and physiological responses in plant growth and abiotic stresses. Plant Stress. 2021. 2:100029. 10.1016/j.stress.2021.100029
    CrossRef
  33. Manghwar H, Hussain A, Ali Q, Liu F. Brassinosteroids (BRs) Role in Plant Development and Coping with Different Stresses. Int J Mol Sci. 2022 Jan 18;23(3):1012. doi: 10.3390/ijms23031012.
    CrossRef
  34. Tungmunnithum D, Thongboonyou A, Pholboon A, Yangsabai A. Flavonoids and Other Phenolic Compounds from Medicinal Plants for Pharmaceutical and Medical Aspects: An Overview. Medicines (Basel). 2018 Aug 25;5(3):93. doi: 10.3390/medicines5030093.
    CrossRef
  35. Ninkuu V, Zhang L, Yan J, Fu Z, Yang T, Zeng H. Biochemistry of Terpenes and Recent Advances in Plant Protection. Int J Mol Sci. 2021 May 27;22(11):5710. doi: 10.3390/ijms22115710.
    CrossRef
  36. War AR, Paulraj MG, Ahmad T, Buhroo AA, Hussain B, Ignacimuthu S, Sharma HC. Mechanisms of plant defense against insect herbivores. Plant Signal Behav. 2012 Oct 1;7(10):1306-20. doi: 10.4161/psb.21663.
    CrossRef
  37. Lv Q, Li X, Fan B, Zhu C, Chen Z. The Cellular and Subcellular Organization of the Glucosinolate-Myrosinase System against Herbivores and Pathogens. Int J Mol Sci. 2022 Jan 29;23(3):1577. doi: 10.3390/ijms23031577.
    CrossRef
  38. Divekar PA, Narayana S, Divekar BA, Kumar R, Gadratagi BG, Ray A, Singh AK, Rani V, Singh V, Singh AK, Kumar A, Singh RP, Meena RS, Behera TK. Plant Secondary Metabolites as Defense Tools against Herbivores for Sustainable Crop Protection. Int J Mol Sci. 2022 Feb 28;23(5):2690. doi: 10.3390/ijms23052690.
    CrossRef
  39. Chung KT, Wong TY, Wei CI, Huang YW, Lin Y. Tannins and human health: a review. Crit Rev Food Sci Nutr. 1998 Aug;38(6):421-64. doi: 10.1080/10408699891274273. PMID: 9759559.
    CrossRef
  40. Zulfiqar, Faisal & Aisha, Nudrat & Ashraf, Muhammad. Osmoprotection in plants under abiotic stresses: new insights into a classical phenomenon. Planta. 2019.251. 10.1007/s00425-019-03293-1.
    CrossRef
  41. Seleiman MF, Al-Suhaibani N, Ali N, Akmal M, Alotaibi M, Refay Y, Dindaroglu T, Abdul-Wajid HH, Battaglia ML. Drought Stress Impacts on Plants and Different Approaches to Alleviate Its Adverse Effects. Plants (Basel). 2021 Jan 28;10(2):259. doi: 10.3390/plants10020259.
    CrossRef
  42. M. M. Chaves, M. M. Oliveira, Mechanisms underlying plant resilience to water deficits: prospects for water-saving agriculture, Journal of Experimental Botany, 2004. 55, Issue 407, 2365–2384, https://doi.org/10.1093/jxb/erh269
    CrossRef
  43. Park CJ, Seo YS. Heat Shock Proteins: A Review of the Molecular Chaperones for Plant Immunity. Plant Pathol J. 2015 Dec;31(4):323-33. doi: 10.5423/PPJ.RW.08.2015.0150.
    CrossRef
  44. Delavari A, Zinati Z, Sazegari S, Tahmasebi A. Integrating expression data and genomic sequences to investigate transcriptional regulation in barley in response to abiotic stress. BioTechnologia (Pozn). 2021 Mar 31;102(1):21-32. doi: 10.5114/bta.2021.103759.
    CrossRef
  45. Pallavi Sharma, Ambuj Bhushan Jha, Rama Shanker Dubey, Mohammad Pessarakli, “Reactive Oxygen Species, Oxidative Damage, and Antioxidative Defense Mechanism in Plants under Stressful Conditions”, Journal of Botany, 2012, Article ID 217037, 26 pages, 2012. https://doi.org/10.1155/2012/217037
    CrossRef
  46. Agati G, Azzarello E, Pollastri S, Tattini M. Flavonoids as antioxidants in plants: location and functional significance. Plant Sci. 2012, Nov;196:67-76. doi: 10.1016/j.plantsci.2012.07.014.
    CrossRef
  47. Kamran M., Wang D., Alhaithloul H. A. S., Alghanem S. M., Aftab T., Xie K., et al.  Jasmonic acid-mediated enhanced regulation of oxidative, glyoxalase defense system and reduced chromium uptake contributes to alleviation of chromium (VI) toxicity in choysum (Brassica parachinensis L.). Ecotoxicol. Environ. Saf. 2021b, 208:111758. 10.1016/j.ecoenv.2020.111758
    CrossRef
  48. Khosravi-nejad F., Khavari-nejad R. A., Moradi F., Najafi F.  Cytokinin and abscisic acid alleviate drought stress through changing organic acids profile, ion immolation, and fatty acid profile to improve yield of wheat (Triticum aestivum L.) cultivars. Physiol. Mol. Biol. Plants. 2022, 28 1119–1129. 10.1007/s12298-022-01173-9
    CrossRef
  49. Khalid A., Aftab F.  Effect of exogenous application of IAA and GA3 on growth, protein content, and antioxidant enzymes of Solanum tuberosum L. grown in vitro under salt stress. In vitro Cell Dev. Biol. Plant. 2020, 56 377–389. 10.1007/s11627-019-10047-x
    CrossRef
  50. Apel, K., & Hirt, H. Reactive oxygen species: metabolism, oxidative stress, and signal transduction. Annual Review of Plant Biology,2004, 55, 373-399.
    CrossRef
  51. Foyer, C. H., & Noctor, G. Redox homeostasis and antioxidant signaling: a metabolic interface between stress perception and physiological responses. The Plant Cell, 2005, 17(7), 1866-1875.
    CrossRef
  52. Ramakrishna A, Ravishankar GA. Influence of abiotic stress signals on secondary metabolites in plants. Plant Signal Behav. 2011 Nov;6(11):1720-31. doi: 10.4161/psb.6.11.17613.
    CrossRef
  53. Li Zhang, Liangwei Zhang, Xia Zhang, Yang Zhao, Shujing Fang, Jinmao You, Lingxin Chen. Responsive fluorescent probes for cellular microenvironment and redox small biomolecules, TrAC Trends in Analytical Chemistry, 2023, 169, https://doi.org/10.1016/j.trac.2023.117377.
    CrossRef
  54. Ji-Ting Hou, Kang-Kang Yu, Kyoung Sunwoo, Won Young Kim, Seyoung Koo, Jinyu Wang, Wen Xiu Ren, Shan Wang, Xiao-Qi Yu, Jong Seung Kim. Fluorescent Imaging of Reactive Oxygen and Nitrogen Species Associated with Pathophysiological Processes, Chem, 2020. 6(4), 832-866, https://doi.org/10.1016/j.chempr.2019.12.005.
    CrossRef
  55. Gupta B, Huang B. Mechanism of salinity tolerance in plants: physiological, biochemical, and molecular characterization. Int J Genomics. 2014;2014:701596. doi: 10.1155/2014/701596.
    CrossRef
  56. Zheng Y, Wang X, Cui X, Wang K, Wang Y, He Y. Phytohormones regulate the abiotic stress: An overview of physiological, biochemical, and molecular responses in horticultural crops. Front Plant Sci. 2023 Jan 6;13:1095363. doi: 10.3389/fpls.2022.1095363.
    CrossRef
  57. Awais, Muhammad & Li, Wei & Cheema, Muhammad Jehanzeb & Zaman, Q. & Shah, Ash & Aslam, Bilal & Zhu, W. & Ajmal, Muhammad & Faheem, Muhammad & Hussain, Sajjad & Nadeem, Adeel & Afzal, Muhammad & Liu, C. UAV-based remote sensing in plant stress imagine using high-resolution thermal sensor for digital agriculture practices: a meta-review. International Journal of Environmental Science and Technology. 2022. 20. 10.1007/s13762-021-03801-5.
    CrossRef
  58. Roychowdhury R, Das SP, Gupta A, Parihar P, Chandrasekhar K, Sarker U, Kumar A, Ramrao DP, Sudhakar C. Multi-Omics Pipeline and Omics-Integration Approach to Decipher Plant’s Abiotic Stress Tolerance Responses. Genes (Basel). 2023 Jun 16;14(6):1281. doi: 10.3390/genes14061281.
    CrossRef
  59. Verma V, Ravindran P, Kumar PP. Plant hormone-mediated regulation of stress responses. BMC Plant Biol. 2016 Apr 14;16:86. doi: 10.1186/s12870-016-0771-y.
    CrossRef
  60. Anand U, Jacobo-Herrera N, Altemimi A, Lakhssassi N. A Comprehensive Review on Medicinal Plants as Antimicrobial Therapeutics: Potential Avenues of Biocompatible Drug Discovery. Metabolites. 2019, Nov 1;9(11):258. doi: 10.3390/metabo9110258.
    CrossRef
  61. Pandey A., Kumar S. Perspective on Plant Products as Antimicrobials Agents: A Review. Pharmacologia. 2013;4:469–480. doi: 10.5567/pharmacologia.2013.469.480.
    CrossRef
  62. Vaou N, Stavropoulou E, Voidarou C, Tsigalou C, Bezirtzoglou E. Towards Advances in Medicinal Plant Antimicrobial Activity: A Review Study on Challenges and Future Perspectives. Microorganisms. 2021 Sep 27;9(10):2041. doi: 10.3390/microorganisms9102041. P
    CrossRef
  63. Mithöfer A, Boland W. Plant defense against herbivores: chemical aspects. Annu Rev Plant Biol. 2012;63:431-50. doi: 10.1146/annurev-arplant-042110-103854.
    CrossRef
  64. Steppuhn A, Gase K, Krock B, Halitschke R, Baldwin IT. Nicotine’s defensive function in nature. PLoS Biol. 2004 Aug;2(8):E217. doi: 10.1371/journal.pbio.0020217.
    CrossRef
  65. Wink, M. Production and Application of Phytochemicals from an Agricultural Perspective. In: van Beek, T.A. and Breteler, H., Eds., Phytochemistry and Agriculture, Clarendon Press, Oxford, 1993. 171-213.
    CrossRef
  66. Nguyen H. C., Lin K. H., Ho S. L., Chiang C. M., Yang C. M. Enhancing the abiotic stress tolerance of plants: from chemical treatment to biotechnological approaches. Physiol. Plan. 2018, 164. 452–466. 10.1111/ppl.12812
    CrossRef
  67. Lu, W.; Zhao, Y.; Liu, J.; Zhou, B.; Wei, G.; Ni, R.; Zhang, S.; Guo, J. Comparative Analysis of Antioxidant System and Salt-Stress Tolerance in Two Hibiscus Cultivars Exposed to NaCl Toxicity. Plants. 2023, 12, 1525. https://doi.org/10.3390/plants12071525
    CrossRef
  68. Ashraf, M., Harris, P.J.C. Photosynthesis under stressful environments: An overview. Photosynthetica. 2013, 51, 163–190 . https://doi.org/10.1007/s11099-013-0021-6
    CrossRef
  69. S.V. Leontopoulos, I. Giavasis, K. Petrotos, M. Kokkora, Ch. Makridis. Effect of Different Formulations of Polyphenolic Compounds Obtained from OMWW on the Growth of Several Fungal Plant and Food Borne Pathogens. Studies in vitro and in vivo. Agriculture and Agricultural Science Procedia, 2015. 4, 327-337, https://doi.org/10.1016/j.aaspro.2015.03.037.
    CrossRef
  70. Jones Clive G. and Firn R. D. On the evolution of plant secondary chemical diversity. Phil. Trans. R. Soc. Lond. 1991. B333273–280. http://doi.org/10.1098/rstb.1991.0077
    CrossRef
  71. Matus, J.T., Cavallini, E., Loyola, R., Höll, J., Finezzo, L., Dal Santo, S., Vialet, S., Commisso, M., Roman, F., Schubert, A., Alcalde, J.A., Bogs, J., Ageorges, A., Tornielli, G.B. and Arce-Johnson, P. A group of grapevine MYBA transcription factors located in chromosome 14 control anthocyanin synthesis in vegetative organs with different specificities compared with the berry color locus. Plant J, 2017. 91: 220-236. https://doi.org/10.1111/tpj.13558
    CrossRef
  72. Hasanuzzaman M., Bhuyan M. H. M. B., Zulfiqar F., Raza A., Mohsin S. M., Mahmud J. A., et al.  Reactive oxygen species and antioxidant defense in plants under abiotic stress: revisiting the crucial role of a universal defense regulator. Antioxidants, 2020a,  9:681. 10.3390/antiox9080681
    CrossRef
  73. Khan MI, Fatma M, Per TS, Anjum NA, Khan NA. Salicylic acid-induced abiotic stress tolerance and underlying mechanisms in plants. Front Plant Sci. 2015 Jun 30;6:462. doi: 10.3389/fpls.2015.00462.
    CrossRef
  74. Abid M, Ali S, Qi LK, Zahoor R, Tian Z, Jiang D, Snider JL, Dai T. Physiological and biochemical changes during drought and recovery periods at tillering and jointing stages in wheat (Triticum aestivum L.). Sci Rep. 2018 Mar 15;8(1):4615. doi: 10.1038/s41598-018-21441-7.
    CrossRef
  75. Hectors K, Prinsen E, De Coen W, Jansen MAK, Guisez Y. Arabidopsis thaliana plants acclimated to low dose rates of ultraviolet B radiation show specific changes in morphology and gene expression in the absence of stress symptoms. New Phytol. 2007;175(2):255-270. doi: 10.1111/j.1469-8137.2007.02092.x.
    CrossRef
  76. Muhammad N., Hakim Quraishi U., Chaudhary H. J., Munis M. F. H. Indole-3-acetic acid induces biochemical and physiological changes in wheat under drought stress conditions. Philipp. Agric. Sci. 2016, 99 19–24.
  77. Yang L, Qiao L, Su X, Ji B, Dong C. Drought Stress Stimulates the Terpenoid Backbone and Triterpenoid Biosynthesis Pathway to Promote the Synthesis of Saikosaponin in Bupleurum chinense DC. Roots. Molecules. 2022 Aug 25;27(17):5470. doi: 10.3390/molecules27175470.
    CrossRef
  78. Mierziak J, Kostyn K, Kulma A. Flavonoids as important molecules of plant interactions with the environment. Molecules. 2014 Oct 10;19(10):16240-65. doi: 10.3390/molecules191016240.
    CrossRef
  79. Verma, V., Ravindran, P. & Kumar, P.P. Plant hormone-mediated regulation of stress responses. BMC Plant Biol 16, 86 (2016). https://doi.org/10.1186/s12870-016-0771-y
    CrossRef
  80. Choudhury F. K., Rivero R. M., Blumwald E., Mittler R.  Reactive oxygen species, abiotic stress and stress combination. Plant J. 2017, 90 856–867. 10.1111/tpj.13299
    CrossRef
  81. Sharifi-Rad J, Rayess YE, Rizk AA, Sadaka C, Zgheib R, Zam W, Sestito S, Rapposelli S, Neffe-Skocińska K, Zielińska D, Salehi B, Setzer WN, Dosoky NS, Taheri Y, El Beyrouthy M, Martorell M, Ostrander EA, Suleria HAR, Cho WC, Maroyi A, Martins N. Turmeric and Its Major Compound Curcumin on Health: Bioactive Effects and Safety Profiles for Food, Pharmaceutical, Biotechnological and Medicinal Applications. Front Pharmacol. 2020 Sep 15;11:01021. doi: 10.3389/fphar.2020.01021.
    CrossRef
  82. Cohen MM. Tulsi – Ocimum sanctum: A herb for all reasons. J Ayurveda Integr Med. 2014 Oct-Dec;5(4):251-9. doi: 10.4103/0975-9476.146554.
    CrossRef
  83. Sánchez M, González-Burgos E, Iglesias I, Gómez-Serranillos MP. Pharmacological Update Properties of Aloe Vera and its Major Active Constituents. Molecules. 2020 Mar 13;25(6):1324. doi: 10.3390/molecules25061324.
    CrossRef
  84. Vergara-Jimenez M, Almatrafi MM, Fernandez ML. Bioactive Components in Moringa Oleifera Leaves Protect against Chronic Disease. Antioxidants (Basel). 2017 Nov 16;6(4):91. doi: 10.3390/antiox6040091.
    CrossRef
  85. Bavarsad NH, Bagheri S, Kourosh-Arami M, Komaki A. Aromatherapy for the brain: Lavender’s healing effect on epilepsy, depression, anxiety, migraine, and Alzheimer’s disease: A review article. Heliyon. 2023 Jul 20;9(8):e18492. doi: 10.1016/j.heliyon.2023.e18492.
    CrossRef
  86. Farooq, M., Wahid, A., Kobayashi, N. Fujita, D., & Basra, S. M. A. Plant drought stress: effects, mechanisms and management. Agron. Sustain. Dev. 2009. 29, 185–212 https://doi.org/10.1051/agro:2008021
    CrossRef
  87. Balasubramaniam, T.; Shen, G.; Esmaeili, N.; Zhang, H. Plants’ Response Mechanisms to Salinity Stress. Plants 2023, 12, 2253. https://doi.org/10.3390/plants12122253
    CrossRef
  88. Kashyap P, Kumar S, Riar CS, Jindal N, Baniwal P, Guiné RPF, Correia PMR, Mehra R, Kumar H. Recent Advances in Drumstick (Moringa oleifera) Leaves Bioactive Compounds: Composition, Health Benefits, Bioaccessibility, and Dietary Applications. Antioxidants (Basel). 2022 Feb 16;11(2):402. doi: 10.3390/antiox11020402.
    CrossRef
  89. Ody P. The Complete Medicinal Herbal: A Practical Guide to the Healing Properties of Herbs. Skyhorse Publishing Inc.; New York, NY, USA: 2017. pp. 1–271.
  90. Ali A. Y. A., Ibrahim M. E. H., Zhou G., Nimir N. E. A., Elsiddig A. M. I., Jiao X., et al. Gibberellic acid and nitrogen efficiently protect early seedlings growth stage from salt stress damage in Sorghum. Sci. Rep. 2021, 11:6672. 10.1038/s41598-021-84713-9
    CrossRef
  91. Mariana Rivas-San Vicente, Javier Plasencia. Salicylic acid beyond defence: its role in plant growth and development, Journal of Experimental Botany, 2011, 62 (10), 3321–3338, https://doi.org/10.1093/jxb/err031
    CrossRef
  92. Vaghela N, Gohel S. Medicinal plant-associated rhizobacteria enhance the production of pharmaceutically important bioactive compounds under abiotic stress conditions. J Basic Microbiol. 2023 Mar;63(3-4):308-325. doi: 10.1002/jobm.202200361.
    CrossRef
  93. Ahmad Cheikhyoussef, Natascha Cheikhyoussef, Aiman K. H. Bashir, Ahmed A. Hussein. Heavy Metals Detoxification Using Coriander from: Handbook of Coriander (Coriandrum sativum), Chemistry, Functionality, and Applications CRC Press Accessed on: 29 Nov 2023
    CrossRef
  94. Lima J., Lobato A. Brassinosteroids improve photosystem II efficiency, gas exchange, antioxidant enzymes and growth of cowpea plants exposed to water deficit. Physiol. Mol. Biol. Plants. 2017, 23 59–72. 10.1007/s12298-016-0410-y
    CrossRef
  95. de Oliveira JR, Camargo SEA, de Oliveira LD. Rosmarinus officinalis L. (rosemary) as therapeutic and prophylactic agent. J Biomed Sci. 2019 Jan 9;26(1):5. doi: 10.1186/s12929-019-0499-8.
    CrossRef
  96. Hayat S, Cheng Z, Ahmad H, Ali M, Chen X, Wang M. Garlic, from Remedy to Stimulant: Evaluation of Antifungal Potential Reveals Diversity in Phytoalexin Allicin Content among Garlic Cultivars; Allicin Containing Aqueous Garlic Extracts Trigger Antioxidants in Cucumber. Front Plant Sci. 2016 Aug 25;7:1235. doi: 10.3389/fpls.2016.01235.
    CrossRef
  97. Vaou N, Stavropoulou E, Voidarou CC, Tsakris Z, Rozos G, Tsigalou C, Bezirtzoglou E. Interactions between Medical Plant-Derived Bioactive Compounds: Focus on Antimicrobial Combination Effects. Antibiotics (Basel). 2022 Jul 28;11(8):1014. doi: 10.3390/antibiotics11081014.
    CrossRef
  98. Panossian AG, Efferth T, Shikov AN, Pozharitskaya ON, Kuchta K, Mukherjee PK, Banerjee S, Heinrich M, Wu W, Guo DA, Wagner H. Evolution of the adaptogenic concept from traditional use to medical systems: Pharmacology of stress- and aging-related diseases. Med Res Rev. 2021 Jan;41(1):630-703. doi: 10.1002/med.21743.
    CrossRef
scroll to top