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oxidative modifications to DNA, proteins, and other important biomolecules
resulting in different disease induction . Plants respond to such distress by altering
the gene expression profile toward activating enzymatic and non-enzymatic anti-
oxidant genes. Also, SM biosynthesis genes will be upregulated. Gene expression
studies in Cucumis sativus L. showed elevated mRNA levels of key antioxidant
enzymes like peroxidase (POX), catalase (CAT), and phenyl ammonia l
yase (PAL).
This was further confirmed with a corresponding increase in these enzyme activities.
In addition, there was a corresponding increase in the enzyme activity of SM
biosynthetic pathways (Cantarello et al.
2005)
. F
urther studies have also reported
an increased expression of genes coding transcriptional factors like MYBs and
enzymes like anthocyanidin synthase, which play an important role in protection
from UV-B-induced damages in plants (Cantarello et al.
2005; Ryan et al. 2002).
Also, elevated expression of genes synthesizing SM having the potential to absorb
UV-B radiations and free radical neutralization has been reported (Ravindran and
Stein
2010
). In addition, genes encoding enzymes like glutathione reductase and
glutathione-s-transferase were reported to be elevated, resulting in increased activa-
tion of chalcone synthase, which has been shown to demonstrate UV-B light and
heavy metal-induced oxidative stress in many medicinal plants (Loyall et al.
2000).
Medicinal plants are grown in different climatic regions where chronic temperature
fluctuations occur. When the plants face extremes in environmental temperature,
they find it challenging to maintai n their homeostasis. Slowly they start adapting to
the situation by altering physical, chemical, and biological processes resulting in
structural and anatomical modifications suitable for their existence (Lianopoulou
et al.
2014). Cold stress is a major threat limiting the average growth and develop-
ment of medicinal plants (Eremina et al. 2016). Intricate signaling pathways get
activated and cross-communicate to make the necessary changes at the gene expres-
sion level. This results in tolerance to cold, chilling, or freezing and includes several
metabolic and enzymatic pathways (Ruelland and Zachowski
2010
). The primary
modifications employed by tolerant plants include reduced growth rate, favorable
membrane remodification, activating cold sensing signal transduction pathways,
expression of temperature-sensitive transcription factors, production of proteins
and metabolites having cryoprotective properties, and others. Gene expression
studies have revealed enhanced levels of these genes under cold stress. Genetic
engineering of these genes has conferred be
tter cold-tolerant properties in transgenic
medicinal plants. Specific genes like CAS15 in alfalfa and COR15a in Arabidopsis
thaliana have been demonstrated to confer cold and freeze resistance properties by
different mechanisms. These gene sequences can be transferred to cold susceptible
plants by genetic engineering techniques, thereby enhancing their potential to adapt
to the freezing environment (Monroy et al.
1993; Artus et al. 1996), (Table 6.1).
6 Gene Expression in Medicinal Plants in Stress Conditions 99
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100 Sadashivaiah et al.
Table 6.1 Transcriptomic profile of a few medicinal plants
SL. No. Species name References
1. Vitis amurensis Xu et al. (2014)
2. Aloe vera Choudhri et al. (2018)
3. Paeonia suffruticosa Guo et al. (2018)
4. Trillium govanianum Singh et al. (2017)
5. Polygonum minus Loke et al. (2017)
6. Callery aspeciosa, Xu et al. (2016)
7. Cornus officinalis Hou et al. (2018)
8. Populus alba. Vu et al. (2020)
9. Gastrodia elata Wang et al. (2018)
10. Entada phaseoloides Liao et al. (2020)
11. Lantana camara Shah et al. (2020)
12. Tetrastigma hemsleyanum Yan et al. (2020)
13. Saussurea lappa Bains et al. (2019)
14. Arisaema heterophylum Blume Wang et al. (2018)
15. Ginkgo biloba Wu et al. (2018)
16. Abrus mollis L Yuan et al. (2018)
17. Artemisia argyi Liu et al. (2018)
18. Panax ginseng Jayakodi et al. (2015)
19. Panax notoginseng Luo et al. (2011)
20. Angelica
sinensis Feng et al. (2020)
6.5 Conclusion
Understanding the gene expression pattern is a crucial task. However, understanding
the same under multiple condition s in multiple tissues of medicinal plants is still
challenging. Medicinal plants are primarily grown to harvest increased amounts of
SM they produce, which possess significant therapeutic and prophylactic properties
against a plethora of diseases. Medicinal plants, in particular, and plants in general,
are known to increase their production of SM whenever they face any stress
conditions. This is primarily mediated by the activation of their defense system,
which produces many SM in addition to a significant change in basic biochemical
and molecular processes.
Analyzing gene expression patterns in medicinal plants during normal and
stressed conditions gives us information about the genetic, biochemical, and physi-
ological changes that the plants adapt to overcome the stress. These altered
mechanisms exhibited by the plant s facing diverse stress factors, either individually
or collectively in different combinations provide an enormous amount of informa-
tion. When carefully examined and understood, this data will give multiple ways and
means to be utilized by a biotechnologist or a plant breeder to engineer new plant
cultivars with unique genetic changes which confer them special properties that can
be exploited to produce increased amounts of SM. Further, it enables the scientific
https://t.me/medicina_free
community to develop resistant medicinal and crop plant cultivars with high pro-
ductivity potential under stress conditions. It thus can solve the ever-increasing
global food supply.
6 Gene Expression in Medicinal Plants in Stress Conditions 101
In addition, it can also be exploited in cell and tissue culture technologies to
produce increased amounts
of SM with huge demand in food, pharmaceut ical, and
healthcare sectors without any impact from climate and environmental changes.
All this is possible only with the development of sophisticated tools and
techniques of gene expression studies which otherwise would have been an arduous
task. Thus, understanding functional genomics, transcriptomics, proteomics, and
metabolomics of an organism will tremendously impact the omics era.
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107
Revealing the Epigenetic Mechanisms
Underlying the Stress Response
in Medicinal Plants
7
Chandrashekhar Singh, Rajesh Saini, Richa Upadhyay,
and Kavindra Nath Tiwari
Abstract
Epigenetics involves a heritable change in gene expression patterns in response to
any stress in plants. To cope with various types of environmental stressors, plants
possess intricate physiological and biochemical processes. The epigenetic
alterations are principally responsibl e for the changes in the transcriptome and
proteome. Medicinal plants are a valuable source of new and innovative
medicines. Plants may develop a defense response to combat abiotic environmen-
tal stress by creating a variety of plant secondary metabolites (PSMs) to escape
from cell and tissue damage. In most cases, plants opt for one of two pathways:
either synthesize new compounds from scratch or boost the quantity of already-
existing PSMs. Most PSMs with antioxidant properties, such as terpenoids,
alkaloids, and phenolic compounds, are involved in plant defense. Epigenetic
alterations, such as DNA methylation, acetylation, phosphorylation, and methyl-
ation of histone proteins, primarily regulate transcription and translation. The
current review highlights and examines the role that various epigenetic
modifications play in plants’ reactions to abiotic stress. Although many abiotic
pressures cause comparable initial reactions in plants, new research has revealed
that certain abiotic stresses may cause epigenetic changes in plants. This review
will also provide a thorough explanation of the abiotic stress-related reactions and
underlying biological processes in plants.
C. Singh
Department of Pharmaceutical Engineering and Technology, Indian Institute of Technology, BHU,
Varanasi, India
R. Saini · K. N. Tiwari (
✉)
Department of Botany, MMV, Banaras Hindu University, Varanasi, India
R. Upadhyay
Department of Botany, Mihir Bhoj Postgraduate College, G.B. Nagar, India
#
The Author(s), under exclusive license to Springer Nature Singapore Pte
Ltd. 2023
D. Singh et al. (eds.), Stress-responsive Factors and Molecular Farming
in Medicinal Plants,
https://doi.org/10.1007/978-981-99-4480-4_7
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108 C. Singh et al.
Keywords
Epigenetics · Plant secondary metabolites · Abiotic stress · Biotic stress
7.1 Introduction
In plants, epigenetic changes are crucial for growth, development, and stress adap-
tation (Smulders and De Klerk 2011; Mirouze and Paszkowski 2011). Plant
epigenetics has grown tremendously in recent years, both in fundamental research
and crop improvement (Mirouze and Paszkowski 2011). The development of plant
growth and development processes, such as seed germination, photosynthetic effi-
ciency, and reproduction, are affected by abiotic and biotic factors. Environmental
challenges cause oxidative damage in plants due to the generation of free radicals
(Tripathy and Oelmüller 2012; Dor ion et al. 2021 ). To deal with abiotic and biotic
stresses, plants have evolved a number of adaptations. To detect stress signals, plants
have developed highly complex signal transduction pathways. A variety of signaling
cascades are started by the generation of second messengers like ROS, inositol
phosphates, and Ca
2+
. As a result, the transcriptome is altered, and a variety of
defensive molecules are produced (Desikan et al.
2005; Tuteja and Maharajan 2007).
Together with protective proteins, these substances work either directly or indirectly
to shield plant cells from the damaging effects of abiotic stress. Epigenetics has
become one of the hottest research fields in plant functional genomics due to their
role in stress-adaptiveness in plants (Thiebaut et al.
2019; Kakoulidou et al. 2021).
The epigenetic control of stress-induced phenotypic responses involves gene regu-
lation in plants (Gallo-Franco et al.
2020). DNA methylation plays a crucial role in
the phenotypic variability of plants under stress (Akhter et al. 2021). This mecha-
nism has been connected to siRNA-mediated methylation, transposon mobility, and
host methyltransferase activity (Saze et al.
2012). Several studies reported on the
state of the art in plant stress response epigenetics in order to engineer plants for
stress tolerance. Gene expression in plants is regulated by Si-RNAs, acetylation–
deacetylation, and methylation–demethylation without changing their DNA
sequences (Sudan et al. 2018).
DNA methylation contributes significantly to genome stability and other cellular
and biological processes that help the body adapt to stress. Plants have a specific
DNA methyltransferase called CHRO MOMETHYLASE3 (CMT3) involved in
DNA methylation. CMT3 has a beneficial role in the heat-induced activation of
the retrotransposon ONSEN and is necessary for the complete transcription of
ONSEN (Nozawa et al.
2021). Sessile plants have a collection of signaling channels
that collect, transmit, and eventually rewire transcriptional signals from the environ-
ment. Histone proteins a re a crucial component of the chromatin structure. Numer-
ous histone-modifying enzymes influence the expression of genes in response to
environmental stresses (Kim et al.
2015
). DNA methylation and histone modifica-
tion play important roles in the tissue-specific expression of the gene and their
silencing (Bhadouriya et al.
2021). Different chromatin remodeler genes, such as
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CHD, INO80, SWI/SNF, and ISWI, change the transcriptionally inactive chromatin
into the active one (Tyagi et al. 2016). The chromatin architecture at a certain
promoter determines the transcriptional output (Bhadouriya et al. 2021). Si-RNA,
DNA methylation, and chromatin alterations play important roles in the epigenetic
control of the genes in response to stresses. The regulation of stress-related genes has
also been linked to epigenetic processes. Active DNA demethylation in plants occurs
by base excision repair, by 5-methylcytosine DNA glycosylases (Zhang and Zhu
2012; Li et al. 2015). The present study gives a useful insight into the different
mechanisms of epigenetic control in plants against different stress responses.
7 Revealing the Epigenetic Mechanisms Underlying the Stress Response... 109
7.2 Abiotic Stresses
Extreme cold, heat shock, a lack of water, excessive salinity, a lack of nutrients, and
heavy metal toxicity are the principal abiotic stresses. Abiotic stress can cause a
variety of negative impacts on plants and other organisms, incl uding reduced growth
and production, reduced photosynthesis and respiration, reduced nutrient uptake,
reduced seed and fruit set, increased susceptibility to disease, reduced tolerance to
environmental extremes, reduced plant vigor and yield, and increased mortality.
7.2.1 Heat Stress
Heat stress can also cause changes in histone modifications, which can impact gene
expression. Terpenoids accumulate as a result of warming, and they often serve
protective purposes by reduci ng the effects of environmental oxidative stress on
plants (Copolovici et al.
2012). For example, tomatoes cultivated under heat stress
(at 46 °C) release increased quantities of monoterpenes, such as -thujene, -pinene,
and terpin. Heat stress affects membranes (such as the thylakoid membrane) and
breaks down membrane protein complexes (such as photosystem II), which reduces
the rate of photosynthesis under simulated climatic circumstances. Plants combat
such harm by continuously synthesizing and emitting terpenes (Dekker and
Boekema 2005).
Korankye et al. (2017) postulated that plants shift carbon to generate more
terpenes in stressful situations. to a pathway other than the mevalonate pathway,
which would have normally been employed for photosynthesis. Linalyl acetate,
limonene, and sabinene are examples of monoterpenes. The highest levels of
myrcene, -terpinene, -ocimene, -terpinolene, and -terpinene are created when
photosynthesis in plants is reduced (Korankye et al. 2017).
7.2.2 Cold Stress
Cold stress can affect epigenetic modification. Cold stress has been shown to induce
histone modifications, which can alter gene expression. These epigenetic alterations
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