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      Halophytes: Potential Resources for Salt Stress Tolerance Genes and Promoters

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          Abstract

          Halophytes have demonstrated their capability to thrive under extremely saline conditions and thus considered as one of the best germplasm for saline agriculture. Salinity is a worldwide problem, and the salt-affected areas are increasing day-by-day because of scanty rainfall, poor irrigation system, salt ingression, water contamination, and other environmental factors. The salinity stress tolerance mechanism is a very complex phenomenon, and some pathways are coordinately linked for imparting salinity tolerance. Though a number of salt responsive genes have been reported from the halophytes, there is always a quest for promising stress-responsive genes that can modulate plant physiology according to the salt stress. Halophytes such as Aeluropus, Mesembryanthemum, Suaeda, Atriplex, Thellungiella, Cakile, and Salicornia serve as a potential candidate for the salt-responsive genes and promoters. Several known genes like antiporters ( NHX, SOS, HKT, VTPase), ion channels (Cl , Ca 2+, aquaporins), antioxidant encoding genes ( APX, CAT, GST, BADH, SOD) and some novel genes such as USP, SDR1, SRP etc. were isolated from halophytes and explored for developing stress tolerance in the crop plants (glycophytes). It is evidenced that stress triggers salt sensors that lead to the activation of stress tolerance mechanisms which involve multiple signaling proteins, up- or down-regulation of several genes, and finally the distinctive or collective effects of stress-responsive genes. In this review, halophytes are discussed as an excellent platform for salt responsive genes which can be utilized for developing salinity tolerance in crop plants through genetic engineering.

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          Most cited references127

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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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            Learning from halophytes: physiological basis and strategies to improve abiotic stress tolerance in crops.

            Global annual losses in agricultural production from salt-affected land are in excess of US$12 billion and rising. At the same time, a significant amount of arable land is becoming lost to urban sprawl, forcing agricultural production into marginal areas. Consequently, there is a need for a major breakthrough in crop breeding for salinity tolerance. Given the limited range of genetic diversity in this trait within traditional crops, stress tolerance genes and mechanisms must be identified in extremophiles and then introduced into traditional crops. This review argues that learning from halophytes may be a promising way of achieving this goal. The paper is focused around two central questions: what are the key physiological mechanisms conferring salinity tolerance in halophytes that can be introduced into non-halophyte crop species to improve their performance under saline conditions and what specific genes need to be targeted to achieve this goal? The specific traits that are discussed and advocated include: manipulation of trichome shape, size and density to enable their use for external Na(+) sequestration; increasing the efficiency of internal Na(+) sequestration in vacuoles by the orchestrated regulation of tonoplast NHX exchangers and slow and fast vacuolar channels, combined with greater cytosolic K(+) retention; controlling stomata aperture and optimizing water use efficiency by reducing stomatal density; and efficient control of xylem ion loading, enabling rapid shoot osmotic adjustment while preventing prolonged Na(+) transport to the shoot.
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              Diversity, distribution and roles of osmoprotective compounds accumulated in halophytes under abiotic stress.

              Osmolytes are low-molecular-weight organic solutes, a broad group that encompasses a variety of compounds such as amino acids, tertiary sulphonium and quaternary ammonium compounds, sugars and polyhydric alcohols. Osmolytes are accumulated in the cytoplasm of halophytic species in order to balance the osmotic potential of the Na(+) and Cl(-) accumulated in the vacuole. The advantages of the accumulation of osmolytes are that they keep the main physiological functions of the cell active, the induction of their biosynthesis is controlled by environmental cues, and they can be synthesized at all developmental stages. In addition to their role in osmoregulation, osmolytes have crucial functions in protecting subcellular structures and in scavenging reactive oxygen species.
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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
                18 May 2017
                2017
                : 8
                : 829
                Affiliations
                [1] 1Marine Biotechnology and Ecology Division, Central Salt and Marine Chemicals Research Institute (CSIR) Bhavnagar, India
                [2] 2Academy of Scientific and Innovative Research, Council of Scientific and Industrial Research New Delhi, India
                Author notes

                Edited by: Oscar Vicente, Universitat Politècnica de València, Spain

                Reviewed by: Salman Gulzar, University of Karachi, Pakistan; Marius-Nicusor Grigore, Alexandru Ioan Cuza University, Romania

                This article was submitted to Plant Abiotic Stress, a section of the journal Frontiers in Plant Science

                Article
                10.3389/fpls.2017.00829
                5435751
                28572812
                69f63886-c079-4e16-b1d2-e210cd68917c
                Copyright © 2017 Mishra and Tanna.

                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) or licensor 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 2017
                : 02 May 2017
                Page count
                Figures: 1, Tables: 1, Equations: 0, References: 136, Pages: 10, Words: 8762
                Funding
                Funded by: Council of Scientific and Industrial Research 10.13039/501100001412
                Funded by: Science and Engineering Research Board 10.13039/501100001843
                Categories
                Plant Science
                Mini Review

                Plant science & Botany
                abiotic stress,halophytes,promoter,salinity,salt responsive genes,salt stress,stress tolerance,transgenic

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