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13 Genomic Instability in Medicinal Plants in Response to Heavy Metal Stress 223
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227
Proteomics Response of Medicinal Plants
to Salt Stress
14
L. Sunil , Sadashivaiah, R. Chandrakanth, and Darshan Dorairaj
Abstract
The plant growth, development, and productivity of medicinal plants are signifi-
cantly affected by salt stress. Due to their sensitivity to salt stress, medicinal
plants must develop increased salt tolerance to sustain global agricultural produc-
tivity. To ensure the food supply in the coming decades, it is necessary to produce
medical plants that can withstand salt stress. Saline-adapted plant genomes depict
physiological changes but not the shift in protein levels reflected in matching
gene expression changes at the transcriptome level. However, proteins are a more
reliable predictor since they are directly involved in forming the unique pheno-
type of physiological features adapted to salinity stress. It would be essential to
look at the protein component of the genome. In this chapter, an effort is made to
summarize how proteome studies are used in response to salt stress in medicinal
plants and to understand the plant’s complex mechanisms under salinity stress.
The findings could be used as a benchmark for the species for which there is a
knowledge deficit in this area and are valuable for the proteomic characterization
of phytomedicinal plants. This chapter provides useful proteomic information for
unraveling the molecular mechanisms underlying bioactive compounds in medic-
inal plants under salt stress.
Keywords
Salt stress · Proteomics · Medicinal plants · Food insecurity · Crop improvement
L. Sunil (✉) · D. Dorairaj
Department of Plant Cell Biotechnology, CSIR-Central Food Technological Research Institute,
Mysuru, India
Sadashivaiah · R. Chandrakanth
Department of Molecular Biology, Yuvaraja’s College, University of Mysore, Mysuru, Karnataka,
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_14
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228 L. Sunil et al.
14.1 Introduction
Plants are traditionally used as medicine, especially by poor populations where they
are frequently the only source of primary healthcare. Due to cheaper costs and
additional benefits over standard treatments, using medicinal plants and herbal
medications is an optional treatment for chronic diseases (Pedrete et al.
2019).
Terpenoids (diterpenes and triterpenes), flavonoids, steroids, and tannins are
believed to be responsible for these actions (White et al.
2016). Numerous studies
have investigated their toxicity due to their widespread medicinal usage and research
interest. They demonstrate that the aqueous extract can be used safely and without
causing tissue toxicity (dos Santos and Rieder
2013; Santos et al. 2018). Plant
proteomics has developed and gathered knowledge on various molecular biological
topics. For instance, this science is crucial in assessing and regulating in vitro
morphogenetic processes in plant tissue culture investigations (Dias et al.
2007).
Proteomics can be used to evaluate proteins responsive to biotic or abiotic impacts in
genetically improved plants (Salvato
2010). Evaluation of variations in protein
expression brought on by mutations or even transgenic mechanisms is another
significant application (Dong and Weng
2013). Through protein characterization
by MS and association analyses relating biological plant functions, plant proteome
investigations which were initially conducted on the genera of Arabidopsis and
Oryza are now applied to several species. For example, By using 2DE-Maldi-
TOF, Nogueira and team investigated alterations in Vigna unguiculata defense
proteins in relation to pathogenesis (Nogueira et al.
2007), Datta et al. studied
Arabidopsis thaliana (Datta et al. 2013). The 2DE-LC-MS/MS method has also
been used to identify proteins in Manihot esculenta and Araucaria angustifolia that
are involved in biosynthesis and metabolism as well as to control cell signaling,
oxidative stress regulation, translational processes, oxidative stress regulation, and
energetic metabolic pathways (Li et al. 2010; Balbuena et al. 2011).
Salinity in the soil is a common abiotic stress. High salt concentrations may
disrupt metabolism; lead to mineral nutrient deficiencies, ion imbalance, and osmotic
stress, interfere with water intake; induce oxidative damage; accumulate harmful
ions; and limit growth (Pi et al.
2018). High salinity damages soil structure and
interferes with the ideal air-water balance, which is necessary for the plant’ s typical
biological processes. Stress from salinity has a substantial impact on plant growth,
development, and production. These characteristics include leaf color, area, count
and form, dry mass, and moisture contents (Ismail et al. 2015; Kotagiri and Kollur u
2017). According to available data, plants have evolved sophisticated physiological
and molecular processes that allow for tolerance to the harmful effects of salt stress.
These responses include ion uptake, compartmentalization of Na
+
in vacuoles,
synthesis of compatible solutes, changes in photosynthesis, and regulation of spe-
cific protein expression (Zhang et al. 2012). Our understanding of salinity stress in
species like soybean (Liu et al.
2019a), Jerusalem artichoke (Zhang et al.
2018), and
Arabidopsis (Stanley Kim et al. 2005) has improved because of high-throughput
transcriptomics’ ability to identify salt-responsive genes and molecular regulatory
networks. Additionally, the results from the proteome study could not be fully
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correlated with the transcriptome data because mRNA levels are not always linked to
those of matching proteins in part to posttranscriptional and posttranslational
modifications. Proteomic analysis is a technique that makes it easier to investigate
the overall expression of proteins and gives a wide range of data on the specific
proteins involved in various biological processes.
14 Proteomics Response of Medicinal Plants to Salt Stress 229
The main objective of
the proteome study of plants under salt stress is to discover
potential stress proteins that might be used as protein markers. Under salinity stress
conditions, these markers can gather quantitative data on various physiological
characteristics (Kosová et al.
2011). These approaches have opened up new
opportunities for phytomedicine research to demonstrate the use of medicinal plants
to treat certain chronic health conditions.
14.2 Medicinal Plants Under Salt Stress
Medicinal plants have a long history and are one of the most significant crops for
traditional disease treatment and prevention. According to the World Health Orga-
nization (WHO), 80% of people still use herbal remedies worldwide. Nearly 19% of
adult populations in the United States utilize herbal medicines (Kennedy
2005).
However, environmental factors significantly impact the quantity and quality of
secondary metabolites produced by medicinal plants. Despite the fact that the effects
of salt stress on crops have been thoroughly and extensively studied, there is a lack of
understanding in the case of medicinal plants. These commercially significant crop’s
molecular pathways for secondary metabolism and salt tolerance have not been
studied as thoroughly as other crops. Therefore, it is quite valuable to compare
how medicinal plants react to salt stress with other significant crops. The develop-
ment of medical plants that can withstand salt increases the global production of
essential ingredients for medicines, flavors, fragrances, and spices. The proteomic
approach is a powerful tool to identify the key proteins in secondary metabolism in
medicinal plants since it is vital to know which protein or proteins are involved in the
biosynthetic pathway to increase the synthesis of a particular molecule in a medicinal
plant (Aghaei and Komatsu 2013). Proteomics frequently examines biochemical
processes and intricate plant responses to abiotic stressors. We can identify the
protective mechanisms used by plants by conducting comparative proteome analyses
of plants before and after particular condition s.
Both medicinal plants and crops have similar reactions to salt stress. Many of the
processes discussed for crops also apply to medicinal plants’ mechanisms for salt
tolerance. In medicinal plants under salt stress, the synthesis of suitable solutes like
proline and nitrogen-containing molecules like proteins, carbohydrates, and
polyamines has been shown. These compounds protect cellular macromolecules,
scavenge free radicals, control osmotic pressure, store nitrogen, and maintain cellu-
lar pH. However, because these plants have significant quantities of various aromatic
compounds, some of their response to salt stress is regulated by these aromatic
compounds’ antioxidant characteristics. Important radical scavengers include
phenols formed from phenylpropanoid compounds, such as flavonoids, tannins,
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and hydroxycinnamate esters, created during various stressful conditions (Said-Al
Ahl and Omer 2011).
230 L. Sunil et al.
Plants have developed extens
ive defense mechanisms to tolerate salt stress. In
recent years, many plants like rice, Arabidopsis, wheat, and maize have been the
focus of research into salt tolerance mechanisms (Sharif et al.
2019; Wani et al. 2020;
Zhao et al. 2020). Through whole genome sequencing and comparative
transcriptome analysis, the molecular mechanism behind the salt-tolerant rice culti-
var has been investigated (Chen et al.
2017). More than 2000 genes were discovered
to respond to salt stress during a comparative transcriptional profiling investigation
of a salt-tolerant rice crop and its sensitive parent (Mirdar Mansuri et al. 2019).
According to the findings of another comparative transcriptome study of salt-tolerant
rice under salt stress, ABA signal transduction is intimately related to salt stress (Sun
et al. 2019). Rice with salt tolerance (sd58) and wild-type was subjected to a
proteome study using isobaric tags for relative and absolute quantification
(iTRAQ). According to Kitaake, a heterotrimeric G protein contributes to the salt
response by scavenging ROS (Peng et al.
2019). Salt stress altered 14,172 alternative
splicing (AS) events in cotton, according to RNA-seq data, and Ser/Arg-rich
(SR) proteins associated with AS regulation may be essential in salt stress (Zhu
et al.
2018). According to quantitative proteomics analyses based on maize
SWATH-MS, splicing and transcription factors significantly affect salt stress
(Chen et al. 2021).
The mulberry is a medicinal plant with outstanding nutritional, therapeutic, and
economic benefits and ecological functions (Rosales: Moraceae). Humans can
consume the mulberry fruit, and silkworms depend on the leaves for nourishment.
In addition, mulberry is a valuable genetic resource for abiotic stress and could treat
polluted water and alleviate drought (Liu et al.
2019b). In numerous mulberry
studies, genes related to stress signals were investigated. The generation of oxidative
stress brought on by Cd stress can be stopped by the activities of the enzymes
superoxide dismutase (SOD), catalase (CAT), peroxidase (POD), and ascorbate
peroxidase (APX) in mulberries (Guo et al. 2021).
However, there are still few reports on the molecular processes behind the
synthesis of bioactive components in naturally occurring saline-growing medicinal
plants. We can only develop better-cultivated plants if we fully understand how the
bioactive components of therapeutic plants are formed. Finding differentially
expressed proteins (DEPs) and investigating the major proteins involved in produc-
ing these components in medicinal plants under salt stress become essential as an
outcome (Wang et al. 2020a).
14.3 Proteomic Studies of Medicinal Plants
In general, two methods are used in the proteomic analysis of the sample. The first
one is the most common approach, gel-based proteomic analysis, which depends on
two-dimensional polyacrylamide gel electrophoresis. The other method is gel-free
analysis which includes multidimensional protein identification technology for
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protein separation, isotope-coded affinity technology, isotope-coded prote in label-
ing, and stable isotope labeling of amino acid-coded and other label-free methods in
plant science (Aslam et al. 2017). Information on protein abundances, variations, and
alterations, as well as interaction associates and protein networks, are all part of the
systematic investigation of the proteome (Park
2004). When plants are exposed to
salt stress, their proteomes express themselves differently. Protein profiling research
on plants under stress extensively uses proteomics (Table
1
4.1
). It directly
contributes to identifying genes and proteins responsible for developing tolerance
to salt stress in plants (Kosová et al.
2013; Wang et al. 2014). The sets of proteomes
are separated with a focus on protein quality and quantity by combining differential-
expression proteomics analysis, which focuses on protein identification and relative
quantification (Kosová et al. 2013).
14 Proteomics Response of Medicinal Plants to Salt Stress 231
The individual salt-responsive proteins were identified using gel electrophoresis,
mass spectrometry, and bioinformatics techniques to analyze gel spot patterns and
physiological states (Ahsan et al.
2010; Hwang et al. 2012). Recent developments in
proteomic methods have increased our understanding of the mechanisms and
proteins involved in stre ss response. Most of the time, the expression of related
genes and the relevant proteins are part of a cellular defense system. There are three
main categories of genes that are involved in the stress response: (1) those that are
involved in signaling cascades and transcriptional regulation, (2) those that protect
membranes and proteins, and (3) those that are involved in the uptake and transport
of ions and water molecules. Finding the distinct proteins involved in the stress
response without proteomics was impossible.
Unlike transcripts, proteins directly influence the plant’s stress response. It
includes elements of the transcription and translation machinery and enzymes that
catalyze changes in metabolite levels. Protein plays a direct part in the stress-
acclimation process by altering their level in the tissue of stressed plants. The
phenotypic response of plants to stress, which determines plant tolerance to stress,
is intimately tied to these changes in protein accumulation under stress (Kosová et al.
2013; Potters et al. 2007). On the other hand, changes in gene expres sion at the
transcriptome level do not always accurately reflect changes at the protein level
because of the effects of posttranscriptional regulatory mechanisms (Tian et al. 2004;
Yan et al. 2005, 2006). To better understand the physiological mechanisms, plant
phenotypes, and cellular and molecular processes behind plant stress response and
tolerance, it is crucial to examine the plant proteome under stress at the protein level.
Sorghum (Sorghum bicolor L.) is a natural stress-resistant crop that does rela-
tively well in various challenging conditions. Studying the protein profile of sor-
ghum under salt stress may be very interesting. In line with this, two sorghum
genotypes underwent a proteome analysis to exami ne the proteins that respond to
stress (G-46 and CSV 44 F). The genotypes G-46 and CSV 44F are salt-tolerant and
may produce essential proteins in the salt-stress tolerance (Chakrabarti et al.
2020;
Punia et al.
2020). To generate high-yielding and stress-tolerant sorghum genotypes,
functional analysis of these proteins can be used in breeding programs to help
identify regulatory networks in sorghum under stress (Punia et al. 2020). In another
investigation, wheat seedlings under salinity stress had their proteome profiles
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Species Stress conditions References
performed. This study aimed to examine the alterations that result from salt stress on
the wheat plant at the protein level in the chloroplast. The chloroplast’s interior was
mapped to have 194 DEPs in total. Numerous DEPs were caused by post-salinity
stress treatment and were ascribed to transcription and translation, the Calvin cycle,
232 L. Sunil et al.
Table 14.1 Proteomic profiles have been developed in various species under salt stress
S
No.
1. Aeluropus lagopoides 450 mM NaCl, 10 day Sobhanian et al.
(
2010)
2. Arabidopsis 150 mM NaCl, 48 h Jiang et al.
(
2007)
3. Avena sativa (oats) 150 and 300 mM NaCl, 24 h Xu et al. (2021)
4. Beta vulgaris (beet) 300 mM NaCl, 3 weeks Rasouli et al.
(
2020)
5. Cucumis sativus (cucumber) 200 mM NaCl, 7 days Jiang et al.
(
2020)
6. Cucumis sativus (cucumber) 100 mM NaCl, 72 h Du et al. (2021)
Glycyrrhiza uralensis 200 mM NaCl, 50 days Wang et al.
(
2020a)
7. Hordeum vulgare L. var. nudum 200 mM NaCl, 16 h Lai et al. (2020)
8. Hordeum vulgare (barley) 300 mM NaCl, 6 h Yousefirad et al.
(
2020)
9. Hordeum vulgare (barley) 150 mM NaCl, 14 days Zhu et al. (2020)
10. Jerusalem artichoke 250 mM NaCl, 15 days Zhang et al.
(
2018)
11. Kandelia candel (salt-stressed) 200, 400, 600 mM NaCl Wang et al.
(
2013)
12. Medicago sativa (alfalfa) 50, 100, 200
and 400 mM
NaCl, 14 days
Li et al. (
2020)
13. Oryza sativa L. cv. Bura Rata (rice)
100 mM NaCl, 11 days Ramani and
Apte (
1997)
14. Salicornia europaea 100–800 mM NaCl, 21 days
Wang et al.
(
2009)
15. Sesame 150 mM NaCl, 24 h
Zhang et al.
(
2019)
16. Solanum
lycopersicon cv. Micro-
Tom (tomato)
60 mM NaCl, 22 days
Tang et al.
(2020)
17. Solanum chilense (tomato) 500 mM NaCl, 21 days
Kashyap et al.
(2020)
18. Soybean 150 mM NaCl, 48 h Pi et al. (
2016)
19. Soybean seed genotypes 150 mM NaCl, 144 h
Ma et al. (2014)
20. Sugar beet 50 mM NaCl, 72 h Wu et al. (
2018)
21. Triticum aestivum (wheat) 150 mM NaCl, 24 h Ma et al. (2020)
22. Triticum aestivum (wheat) 200 mM NaCl, 96 h Zhu et al. (2021)
23. Zoysia macrostachya 30 mM NaCl, 24 h Wang et al.
(2020b)
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