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      Halophytes as new model plant species for salt tolerance strategies

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          Abstract

          Soil salinity is becoming a growing issue nowadays, severely affecting the world’s most productive agricultural landscapes. With intersecting and competitive challenges of shrinking agricultural lands and increasing demand for food, there is an emerging need to build resilience for adaptation to anticipated climate change and land degradation. This necessitates the deep decoding of a gene pool of crop plant wild relatives which can be accomplished through salt-tolerant species, such as halophytes, in order to reveal the underlying regulatory mechanisms. Halophytes are generally defined as plants able to survive and complete their life cycle in highly saline environments of at least 200-500 mM of salt solution. The primary criterion for identifying salt-tolerant grasses (STGs) includes the presence of salt glands on the leaf surface and the Na + exclusion mechanism since the interaction and replacement of Na + and K + greatly determines the survivability of STGs in saline environments. During the last decades or so, various salt-tolerant grasses/halophytes have been explored for the mining of salt-tolerant genes and testing their efficacy to improve the limit of salt tolerance in crop plants. Still, the utility of halophytes is limited due to the non-availability of any model halophytic plant system as well as the lack of complete genomic information. To date, although Arabidopsis ( Arabidopsis thaliana) and salt cress ( Thellungiella halophila) are being used as model plants in most salt tolerance studies, these plants are short-lived and can tolerate salinity for a shorter duration only. Thus, identifying the unique genes for salt tolerance pathways in halophytes and their introgression in a related cereal genome for better tolerance to salinity is the need of the hour. Modern technologies including RNA sequencing and genome-wide mapping along with advanced bioinformatics programs have advanced the decoding of the whole genetic information of plants and the development of probable algorithms to correlate stress tolerance limit and yield potential. Hence, this article has been compiled to explore the naturally occurring halophytes as potential model plant species for abiotic stress tolerance and to further breed crop plants to enhance salt tolerance through genomic and molecular tools.

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          Most cited references205

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          Salt tolerance and salinity effects on plants: a review.

          Plants exposed to salt stress undergo changes in their environment. The ability of plants to tolerate salt is determined by multiple biochemical pathways that facilitate retention and/or acquisition of water, protect chloroplast functions, and maintain ion homeostasis. Essential pathways include those that lead to synthesis of osmotically active metabolites, specific proteins, and certain free radical scavenging enzymes that control ion and water flux and support scavenging of oxygen radicals or chaperones. The ability of plants to detoxify radicals under conditions of salt stress is probably the most critical requirement. Many salt-tolerant species accumulate methylated metabolites, which play crucial dual roles as osmoprotectants and as radical scavengers. Their synthesis is correlated with stress-induced enhancement of photorespiration. In this paper, plant responses to salinity stress are reviewed with emphasis on physiological, biochemical, and molecular mechanisms of salt tolerance. This review may help in interdisciplinary studies to assess the ecological significance of salt stress.
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            Salinity tolerance in halophytes.

            Halophytes, plants that survive to reproduce in environments where the salt concentration is around 200 mm NaCl or more, constitute about 1% of the world's flora. Some halophytes show optimal growth in saline conditions; others grow optimally in the absence of salt. However, the tolerance of all halophytes to salinity relies on controlled uptake and compartmentalization of Na+, K+ and Cl- and the synthesis of organic 'compatible' solutes, even where salt glands are operative. Although there is evidence that different species may utilize different transporters in their accumulation of Na+, in general little is known of the proteins and regulatory networks involved. Consequently, it is not yet possible to assign molecular mechanisms to apparent differences in rates of Na+ and Cl- uptake, in root-to-shoot transport (xylem loading and retrieval), or in net selectivity for K+ over Na+. At the cellular level, H+-ATPases in the plasma membrane and tonoplast, as well as the tonoplast H+-PPiase, provide the trans-membrane proton motive force used by various secondary transporters. The widespread occurrence, taxonomically, of halophytes and the general paucity of information on the molecular regulation of tolerance mechanisms persuade us that research should be concentrated on a number of 'model' species that are representative of the various mechanisms that might be involved in tolerance.
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              Mechanism of Salinity Tolerance in Plants: Physiological, Biochemical, and Molecular Characterization

              Salinity is a major abiotic stress limiting growth and productivity of plants in many areas of the world due to increasing use of poor quality of water for irrigation and soil salinization. Plant adaptation or tolerance to salinity stress involves complex physiological traits, metabolic pathways, and molecular or gene networks. A comprehensive understanding on how plants respond to salinity stress at different levels and an integrated approach of combining molecular tools with physiological and biochemical techniques are imperative for the development of salt-tolerant varieties of plants in salt-affected areas. Recent research has identified various adaptive responses to salinity stress at molecular, cellular, metabolic, and physiological levels, although mechanisms underlying salinity tolerance are far from being completely understood. This paper provides a comprehensive review of major research advances on biochemical, physiological, and molecular mechanisms regulating plant adaptation and tolerance to salinity stress.
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                Author and article information

                Contributors
                Journal
                Front Plant Sci
                Front Plant Sci
                Front. Plant Sci.
                Frontiers in Plant Science
                Frontiers Media S.A.
                1664-462X
                11 May 2023
                2023
                : 14
                : 1137211
                Affiliations
                [1] 1 ICAR-Central Soil Salinity Research Institute , Karnl, Haryana, India
                [2] 2 ICAR-Indian Institute of Wheat and Barley Research , Shimla, Himachal Pardesh, India
                [3] 3 Department of Biochemistry, Eternal University, Baru Sahib, Himachal Pardesh , Ludhiana, India
                [4] 4 ICAR-Agriculture Technology Application Research Center , Ludhiana, India
                Author notes

                Edited by: Rakesh Kumar Singh, International Center for Biosaline Agriculture (ICBA), United Arab Emirates

                Reviewed by: G Gururaja Rao, Indian Council of Agricultural Research (ICAR), India; Rajeev Nayan Bahuguna, Dr. Rajendra Prasad Central Agricultural University, India

                *Correspondence: Anita Mann, Anita.mann@ 123456icar.gov.in ; Charu Lata, charu.sharma@ 123456icar.gov.in
                Article
                10.3389/fpls.2023.1137211
                10211249
                37251767
                584cc88f-6360-466b-ae74-1b039f829c66
                Copyright © 2023 Mann, Lata, Kumar, Kumar, Kumar and Sheoran

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 04 January 2023
                : 11 April 2023
                Page count
                Figures: 2, Tables: 2, Equations: 0, References: 205, Pages: 18, Words: 10281
                Funding
                The work on the halophytes Urochondra setulosa, Dichanthium annulatum, Aeluropus lagopoides, Suaeda nudiflora, Sporobolus marginatus, and Leptochloa fusca was funded by ICAR-National Agricultural Science Fund (NASF), New Delhi.
                Categories
                Plant Science
                Review
                Custom metadata
                Plant Breeding

                Plant science & Botany
                halophytes,transcriptomics,salinity,degs (differentially expressed genes),gene transformation,salt tolerance,osmoregulation

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