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and carbon and nitrogen metabolism (Zhu et al. 2021). During a proteomic investi-
gation, a collection of proteins found in sensitive and tolerant lines were identified.
Most of the highly expressed proteins in the tolerant lines were related to photosyn-
thesis, ROS scavenging, and ATP generation (Zhu et al.
2020
). The findings
indicated the essential significance of chloroplast and photosynthesis-related
activities since the ATP-mediated energy production reaction is mainly carried out
in chloroplasts. These ATP energy packets are essential for preserving the steady
state of ion homeostasis, scavenging potentially damaging ROS, and maybe decid-
ing the course of overall growth. As a result, the summarized proteins may aid the
research community in producing crops that ar
e functionally resistant to stress
(Munns et al.
2020).
14 Proteomics Response of Medicinal Plants to Salt Stress 233
14.4 Proteomics Approaches in Response to Salt Stress
A biological field of study, “omics” examines cellular, DNA, protein, and metabolite
structure variations. The goal of omics approaches is to map plant genomes and
define their operat ional functions, changes, and biological processes. Omics
approaches include genomics, transcriptomics, proteomics, and metabolomics.
Multi-omics offers the molecular understanding to produce crops that can withstand
stress (Arif et al.
2020; Raza et al. 2021, 2022). The control of genes determines how
plants respond to salt stress in upregulation or downregulation. A revolutionary idea
in this framework, which deals with a complete understanding of the molecular level
of salinity-stressed plants, is the integration of datasets from multi-omics
investigations. The field of proteomics studies the role, stru cture, function, location,
interactions with other proteins, and application of functional proteins during stress
reactions. We can examine posttranscriptional modifications and changes at the
protein level in more detail owing to proteomics. Proteomics thus becomes a crucial
method for locating important stress protein markers that may aid in developing
resilient crops. Proteomics has improved in feasibility, cost, and value for scientific
research facilities worldwide due to advances in biotechnological tools.
In a recent study, the proteome of rice plants exposed to salinity was
characterized. Two varieties of rice were used: cv. Jhelum, which is salt-sensitive,
and cv. Vytilla-4, which is very salt-tolerant. The research revealed numerous
distinct protein expres sion patterns and reactions in rice plant tissues to salt stress;
notably during salinity stress in cv. Vytilla, proteins involved with photosynthesis
were activated (Frukh et al. 2020).
On the other hand, the PS-II CP47 was suppressed in the Jhelum case. Other
carbon fixation-related proteins such as cytochrome c oxidase subunit 5C, peroxi-
somal (S)-2-hydroxy-acid oxidase GLO1, and peroxisomal (S)-2-hydroxy-acid oxi-
dase GLO5) that are essential for energy metabolism were significantly elevated in
Vytilla (Frukh et al. 2020). This clearly shows that these proteins have the potential
to function as biomarkers in controlling the salinity response of plants without
affecting growth. Hormones play a crucial role in imitating salinity in all plants
(Fahad et al. 2015).
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234 L. Sunil et al.
In another study, the germination of two cultivars (the salt-tolerant lk573 and the
salt-sensitive landrace lk621) was examined under salinity stress. In contrast to the
impaired germination rate of the LK621 cultivar, the LK573 cultivar germinated
properly in saline circumstances. In the salt-tolerant cultivar, 171 differentially
expressed proteins (DEPs) were found. Several of them were expressed at 4 h
following salt stress, including proteins involved in nitrogen metabolism, ascorbate
and alternate metabolism, ABC transporters, and another terpenoid-quinone biosyn-
thesis (Lai et al. 2020). On the other hand, in lk621, most expressed proteins were
linked to fatty acid and terpe noid backbone production. It suggests that to control
barley seed germination under salinity stress, these DEPs interact with plant
hormones (Lai et al.
2020).
14.5 Molecular Mechanism of Medicinal Plants Under Salt
Stress
Salinity-induced gene and protein activation changes in plants result in various gene
modifications and protein activation under salt stress in plant metabolism. The
transcription of the primary gene frequently causes these pathways to divert from
the main metabolic pathways (do Nascimento and Fett-Neto
2010). Medicinal plants
accumulate enzymatic antioxidants like glutathione reductase, antioxidant enzymes
catalase, peroxidase, superoxide dismutase, and ascorbate peroxidase components
during salt stress to prevent oxidative damage (Ozkur et al.
2009; Sarker and Oba
2020). The salicylic acid molecule, a phenolic component in plants, is a growth
regulator essential for regulating plant physiological processes, safeguarding plants
from environmental stress, and enhancing plant growth (Hayat et al.
2008; Liu et al.
2016). Salicylic acid increases pigment content, inhibits ethylene generation, and
promotes growth to counteract the adverse effects of salinity stress (Srivastava and
Dwivedi 2000; Fariduddin et al. 2003; Tuna et al. 2007). The efficiency of
antioxidants in the plant is increased by exogenous salicylic acid at moderate
concentrations (1.0 mM), according to research (Wang et al.
2015; Blanch et al.
2020).
The plant must adapt to salt stress in order to change gene expression and the
protein profile, according to the recent study and this process is complex at the whole
plant and cellular levels (Aghaei and Komatsu
2013). It will be useful in breeding
salt-tolerant plant cultivars to understand the molecular mechanism and innovative
approaches that cause salt stress in plants. Plant roots are discovered to be more
sensitive to salt stress than leaves as a result of the direct effects of soil salt stress on
plant roots (Bernstein et al.
2004). Together, these activities help roots achieve
cellular homeostasis and establish plant growth and development phenotype in
response to salt stress (Zhao et al.
2013). As a result, the proteins identified as
being differentially expressed in plant seedling roots under salt stress may play more
significant roles in aiding our knowledge of how to plant seedlings react to salt
stress. However, there is currently a lack of understanding regarding the proteins in
https://t.me/medicina_free
medicinal plants that respond to salt. Our knowledge of the molecular mechanism
used by this significant crop to respond to salt stress has been limited as a result.
14 Proteomics Response of Medicinal Plants to Salt Stress 235
To adapt to harsh conditions,
plants under salt stress fundamentally alter their
gene expression, which modifies the makeup of the plant transcriptome, proteome,
and metabolome. Numerous studies revealed that in stressful conditions, protein
accumulation varied dramatically (Wang et al. 2014; Ghaffari et al. 2014; Liu et al.
2014). Salinity-tolerant phenot ypes can be conferred on salt-sensitive plants through
genes encoding proteins for osmolyte production, ion channels, receptors, and
salt-responsive signaling factors or enzymes (Tuteja
2007). Therefore, the critical
building blocks for creating salt-tolerant plant types are the identification and
characterization of salt-responsive proteins using high-throughput proteomics
investigations.
Salt stress has an impact on gene expression as well. Salt stress, for instance,
significantly affects genes implicated in the metabolic pathways of nitrogen reduc-
tion and fixation and methionine production (Ouyang et al.
2007). Higher levels of
salt stre ss cause Na
+
and Cl
-
to direc tly damage membrane and enzyme structure,
which causes an unregulated inflow or outflow of minerals and nutrients (Qureshi
et al.
2007). Genome sequence information alone is insufficient to show the facts
regarding the appropriate functioning of genes and the biochemical kinetics of plants
under salt stress, making it impossible to identify the precise response mechanisms.
More comprehensive approaches are required at the transcriptome, metabolome, and
proteome levels. Several studies have shown that certain environmental stresses,
such as cold, salt, and water stress, which cause cellular dehydration, frequently
result in changes in plant gene expression and metabolism, and their signaling
pathways exhibit significant cross-link (Rabbani et al. 2003).
The biosynthesis and regulation of biosynthetic pathway genes are crucial since
they increase secondary metabolite production (Sunil and Shetty 2022). The second
strategy agains t salinity is to increase Na
+
export from cells and thereby achieve
equilibrium. Additionally, Zhu and the group revealed that Arabidopsis could
tolerate salt when a plasma membrane Na
+
/H
+
antiporter is overexpressed (Zhu
2001). The capability for selective ion uptake, ion exclusion, and compartmentaliza-
tion of Na
+
in vacuoles can help maintain low cytosolic salt concentrations in crops
(Liu et al. 2012). For plant cells to withstand salt stress, malate dehydrogenase
(NADP+) and pyruvate phospha te dikinase can be raised, reduced, stimulated, or
completely repressed (Ngara et al.
2012). It is known that the reaction of crops to salt
stress is correlated with different families of proteins that have been freshly pro-
duced, accumulated, or reduced (Ngara et al. 2012).
14.6 Limitations of Proteomics Studies
A comprehensive proteome has not yet been established, despite identifying large-
scale crops’ protein even for the model plant A. thaliana. Even the most cutting-edge
MS instruments cannot detect low-abundance proteins in response to biotic and
abiotic stress. In most cases, these proteins serve as the primary stress regulation
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proteins. Protein fractionation and enrichment enable biological samples to be more
variable. However, it also cannot be used with high-throughput quantitative
proteomic techniques. It is essential to establish the broader importance of posttrans-
lational modifications for crop breeding purposes. Because protein phosphorylation
is typically quick and short, characterizing phosphoproteomics plays a significant
role in crop breeding and enhancement efforts.
It is essential to find more affordable
molecular and functional markers, which can be achieved through proteome
investigations; nevertheless, crop proteomics promise to create novel markers for
plant breeding has not yet been completely exploited (Hossain
2020).
236 L. Sunil et al.
Salt stress is one
of the more well-researched abiotic stresses, and proteomic-
based approaches have given researchers a valuable tool for elucidating the molecu-
lar mechanisms behind salt stress responses. Shortly, it will be feasible to convert
salt-sensitive crops into salt-tolerant crops employing several salt-responsi ve
proteins that have been proposed for crops using these techniques. Despite these
developments in proteomics-based research on crop stress tolerance, there are still
just a few preliminary studies in proteome analysis of medicinal plants under salt
stress or other conditions. The molecular mechanisms underlying the formation of
these molecules are still unknown, even though secondary metabolites from medici-
nal plants, which are utilized for herbal medications, tastes, scents, spices, and other
purposes, are essential. The lack of genomic data on medicinal plants is one of the
primary limits in this field, in addition to these issues. Many of the proteins found in
medicinal plants while performing proteome research have remained unidentified.
As a result, developments in the genome-based knowledge of medicinal plants will
open up new perspectives in studying medicinal plant’ s proteomes. We hope that
identifying key proteins involved in synthesizing secondary metabolites in medicinal
plants will increase the production of highly valued anticancer compound s and other
valuable herbal medicines that are essential for people’s health worldwide.
14.7 Conclusion
Salinity stress is the second largest yield-limiting abiotic factor that significantly
threatens worldwide sustainable agricultural production and undermines efforts to
end “hunger for everyone.” To create future plants that are tolerant to salinity and
reach “zero hunger,” this chapter suggests that recent developments in various
proteomic approaches should be considered a safer process. To adapt to their
saltwater environment and thrive against stressful situations, plants modify their
stress-responsive proteins and accumulate essential metabolites. Therefore, thorough
proteomics studies are essential to dealing with salinity in the current technology and
fast-changing climate period. Proteomic investigations help provide hints regarding
proteins with anonymous functions. Understanding the involvement of these
proteins in the salt tolerance process requires the identification of salt stress-
responsive proteins, particularly defense proteins, and the assessment of their
expression patterns in response to stress. Practical solutions to increase the capacity
of plants to tolerate salt would be based on understanding these protein functions to
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stress adaption by proteomics analysis. In salt stress signaling and metabolic
pathways, novel regulatory mechanisms and low-abundant proteins could also be
found.
14 Proteomics Response of Medicinal Plants to Salt Stress 237
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243
Regulation of PGPR-Related Genes
in Medicinal Plants in Adverse Conditions
15
Kanti Kiran and Gunjan Sharma
Abstract
Bioaugmentation with plant growth-promoting rhizobacteria (PGPR) has the
ability to alleviate the adverse impacts of environmental stresses on plants.
Therefore, this is known as a promising and eco-friendly approach to ensure
sustainable and long-term plant secondary metabolites production under adverse
circumstances. Such microbes have enormous plant growth-promoting (PGP)
properties which can induce drought and heat tolerance in plants by directly or
indirectly impacting molecular, metabolic, and physiological stress responses of
plants. This review aims to garner the current knowledge regarding the mecha-
nism of PGPR through gene expression to induce environmental stress tolerance
in medicinal plants.
Keywords
PGPR · Medicinal plants · Phytobeneficial genes · Genomics
15.1 Introduction
Plant growth-promoting rhizobacteria (PGPR) belong to a diverse group of
terricolous bacteria. Bacteria are the most abundant microorganism though many
other microorganisms such as fungi, algae, and protozoa coexist in the rhizosphere.
These groups of microorganisms can competently colonize the plant root system by
maintaining a symbiotic relationship with the plant roots and promote plant growth
K. Kiran (✉) · G. Sharma
Department of Plant Biotechnology, Gujarat Biotechnology University, Gandhinagar, Gujarat,
India
e-mail: kanti.kiran@gbu.edu.in
#
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_15
https://t.me/medicina_free
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