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Halophytes are the flora of saline soils. They adjust osmotically to soil salinity by accumulating ions and sequestering the vast majority of these (generally Na(+) and Cl(-)) in vacuoles, while in the cytoplasm organic solutes are accumulated to prevent adverse effects on metabolism. At high salinities, however, growth is inhibited. Possible causes are: toxicity to metabolism of Na(+) and/or Cl(-) in the cytoplasm; insufficient osmotic adjustment resulting in reduced net photosynthesis because of stomatal closure; reduced turgor for expansion growth; adverse cellular water relations if ions build up in the apoplast (cell walls) of leaves; diversion of energy needed to maintain solute homeostasis; sub-optimal levels of K(+) (or other mineral nutrients) required for maintaining enzyme activities; possible damage from reactive oxygen species; or changes in hormonal concentrations. This review discusses the evidence for Na(+) and Cl(-) toxicity and the concept of tissue tolerance in relation to halophytes. The data reviewed here suggest that halophytes tolerate cytoplasmic Na(+) and Cl(-) concentrations of 100-200 mm, but whether these ions ever reach toxic concentrations that inhibit metabolism in the cytoplasm or cause death is unknown. Measurements of ion concentrations in the cytosol of various cell types for contrasting species and growth conditions are needed. Future work should also focus on the properties of the tonoplast that enable ion accumulation and prevent ion leakage, such as the special properties of ion transporters and of the lipids that determine membrane permeability.

Most of the water on Earth is seawater, each kilogram of which contains about 35 g of salts, and yet most plants cannot grow in this solution; less than 0·2% of species can develop and reproduce with repeated exposure to seawater. These 'extremophiles' are called halophytes. Improved knowledge of halophytes is of importance to understanding our natural world and to enable the use of some of these fascinating plants in land re-vegetation, as forages for livestock, and to develop salt-tolerant crops. In this Preface to a Special Issue on halophytes and saline adaptations, the evolution of salt tolerance in halophytes, their life-history traits and progress in understanding the molecular, biochemical and physiological mechanisms contributing to salt tolerance are summarized. In particular, cellular processes that underpin the ability of halophytes to tolerate high tissue concentrations of Na+ and Cl−, including regulation of membrane transport, their ability to synthesize compatible solutes and to deal with reactive oxygen species, are highlighted. Interacting stress factors in addition to salinity, such as heavy metals and flooding, are also topics gaining increased attention in the search to understand the biology of halophytes. Halophytes will play increasingly important roles as models for understanding plant salt tolerance, as genetic resources contributing towards the goal of improvement of salt tolerance in some crops, for re-vegetation of saline lands, and as 'niche crops' in their own right for landscapes with saline soils.

As important components in saline agriculture, halophytes can help to provide food for a growing world population. In addition to being potential crops in their own right, halophytes are also potential sources of salt-resistance genes that might help plant breeders and molecular biologists increase the salt tolerance of conventional crop plants. One especially promising halophyte is Suaeda salsa, a euhalophytic herb that occurs both on inland saline soils and in the intertidal zone. The species produces dimorphic seeds: black seeds are sensitive to salinity and remain dormant in light under high salt concentrations, while brown seeds can germinate under high salinity (e.g. 600 mm NaCl) regardless of light. Consequently, the species is useful for studying the mechanisms by which dimorphic seeds are adapted to saline environments. S. salsa has succulent leaves and is highly salt tolerant (e.g. its optimal NaCl concentration for growth is 200 mm). A series of S. salsa genes related to salt tolerance have been cloned and their functions tested: these include SsNHX1, SsHKT1, SsAPX, SsCAT1, SsP5CS and SsBADH. The species is economically important because its fresh branches have high value as a vegetable, and its seed oil is edible and rich in unsaturated fatty acids. Because it can remove salts and heavy metals from saline soils, S. salsa can also be used in the restoration of salinized or contaminated saline land. Because of its economic and ecological value in saline agriculture, S. salsa is one of the most important halophytes in China. In this review, the value of S. salsa as a source of food, medicine and forage is discussed. Its uses in the restoration of salinized or contaminated land and as a source of salt-resistance genes are also considered.

Background and Aims 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. Scope This review discusses the diversity of osmolytes among halophytes and their distribution within taxonomic groups, the intrinsic and extrinsic factors that influence their accumulation, and their role in osmoregulation and osmoprotection. Increasing the osmolyte content in plants is an interesting strategy to improve the growth and yield of crops upon exposure to salinity. Examples of transgenic plants as well as exogenous applications of some osmolytes are also discussed. Finally, the potential use of osmolytes in protein stabilization and solvation in biotechnology, including the pharmaceutical industry and medicine, are considered.

Active removal of Na⁺ from the cytosol into the vacuole plays a critical role in salinity tissue tolerance, but another, often neglected component of this trait is Na⁺ retention in vacuoles. This retention is based on an efficient control of Na⁺-permeable slow- and fast-vacuolar channels that mediate the back-leak of Na⁺ into cytosol and, if not regulated tightly, could result in a futile cycle. This Tansley insight summarizes our current knowledge of regulation of tonoplast Na⁺-permeable channels and discusses the energy cost of vacuolar Na⁺ sequestration, under different scenarios. We also report on a phylogenetic and bioinformatic analysis of the plant two-pore channel family and the difference in its structure and regulation between halophytes and glycophytes, in the context of salinity tolerance.

Ipomoea pes-caprae (Linn.) R. Br. (Convolvulaceae) is a halophytic plant that favorably grows in tropical and subtropical countries in Asia, America, Africa, and Australia. Even though this plant is considered a pan-tropical plant, I. pes-caprae has been found to occur in inland habitats and coasts of wider areas, such as Spain, Anguilla, South Africa, and Marshall Island, either through a purposeful introduction, accidentally by dispersal, or by spreading due to climate change. The plant parts are used in traditional medicine for treating a wide range of diseases, such as inflammation, gastrointestinal disorders, pain, and hypertension. Previous phytochemical analyses of the plant have revealed pharmacologically active components, such as alkaloids, glycosides, steroids, terpenoids, and flavonoids. These phytoconstituents are responsible for the wide range of biological activities possessed by I. pes-caprae plant parts and extracts. This review arranges the previous reports on the botany, distribution, traditional uses, chemical constituents, and biological activities of I. pes-caprae to facilitate further studies that would lead to the discovery of novel bioactive natural products from this halophyte.

In this study, we investigated the seasonal changes in enzyme activities and microbial necromass carbon (C) in the bulk and rhizosphere soils of different halophytes (shrubs, grasses, and forbs) in an inland saline ecosystem in China. We found that the soil enzyme activities and microbial necromass C were 45–646% higher in rhizosphere soils than bulk soils regardless of the season and halophyte functional group. The activities of soil enzymes and bacterial necromass C were highest for forbs (4.9–172.3 nmol g −1  h −1 and 473.2 mg kg −1 , respectively), and grasses had the highest fungal necromass C (FN-C) and total necromass C/soil organic C (TN-C/SOC) ratio. Moreover, the soil enzyme activities were higher in the late season stage (4.0–154.0 nmol g −1  h −1 ) than the early (3.6–132.1 nmol g −1  h −1 ) and middle (3.8–128.9 nmol g −1  h −1 ) season stages. FN-C and TN-C/SOC ratio were significantly higher in the middle season than the early and late seasons. Redundancy analysis showed that the enzyme activities and microbial necromass C mainly depended on the soil physicochemical properties, season, and halophyte functional group. Our results indicate that microbial turnover was consistently faster in rhizosphere soil than bulk soil in an inland saline ecosystem, and forbs could be used as pioneer species to improve microbial turnover during the phytoremediation of saline ecosystems.

IntroductionOptimal planting density (PD) is a critical agronomic practice that directly influences plant growth and yield. Suaeda salsa L. can absorb salts from saline soils and accumulate them in its tissues, thereby playing a significant role in saline-alkali soil remediation. However, studies on the effects of S. salsa PD on soil improvement are limited.MethodsIn this study, a pot experiment was conducted with three planting densities, two plants per pot (D1), four plants per pot (D2), and six plants per pot (D3), to determine their effects on the growth and salt accumulation in S. salsa. The soil moisture in each pot was maintained at 80% of the field capacity and was continuously controlled throughout the growing period using a weighing method.ResultsThe results showed that PD significantly influenced the morphological traits and salt removal capacity. Throughout the growing season, plant height, leaf area, and biomass were highest under D2, exceeding those under D1 by 29–78% and those under D3 by 1–6%. Nitrogen accumulation first increased and then decreased with increasing PD, reaching its maximum under D2 treatment. Although the shoot Na+ concentration decreased over time, the overall biomass continued to increase, resulting in a gradual increase in total salt accumulation, which peaked at the fruiting stage. Under D2, the highest salt removal was observed, reaching nearly 4.17 g pot-1 (84.44 g m-2).DiscussionThe optimal density for S. salsa was 4 plants per pot (81 plants m-2), which led to greater salt removal and higher biomass accumulation.

Mangroves are a group of highly salt-tolerant woody plants. The high water use efficiency of mangroves under saline conditions suggests that regulation of water transport is a crucial component of their salinity tolerance. This review focuses on the processes that contribute to the ability of mangroves to maintain water uptake and limit water loss to the soil and the atmosphere under saline conditions, from micro to macro scales. These processes include: (1) efficient filtering of the incoming water to exclude salt; (2) maintenance of internal osmotic potentials lower than that of the rhizosphere; (3) water-saving properties; and (4) efficient exploitation of less-saline water sources when these become available. Mangroves are inherently plastic and can change their structure at the root, leaf and stand levels in response to salinity in order to exclude salt from the xylem stream, maintain leaf hydraulic conductance, avoid cavitation and regulate water loss (e.g. suberization of roots and alterations of leaf size, succulence and angle, hydraulic anatomy and biomass partitioning). However, much is still unknown about the regulation of water uptake in mangroves, such as how they sense and respond to heterogeneity in root zone salinity, the extent to which they utilize non-stomatally derived CO2 as a water-saving measure and whether they can exploit atmospheric water sources.