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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.

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.

Plant roots must exclude almost all of the Na⁺ and Cl⁻ in saline soil while taking up water, otherwise these ions would build up to high concentrations in leaves. Plants evaporate c. 50 times more water than they retain, so 98% exclusion would result in shoot NaCl concentrations equal to that of the external medium. Taking up just 2% of the NaCl allows a plant to osmotically adjust the Na⁺ and Cl⁻ in vacuoles, while organic solutes provide the balancing osmotic pressure in the cytoplasm. We quantify the costs of this exclusion by roots, the regulation of Na⁺ and Cl⁻ transport through the plant, and the costs of osmotic adjustment with organic solutes in roots.

Knowledge of the genetic regulation of adventitious roots, aerenchyma, and radial oxygen loss barrier formation, and the signaling for acclimation, will assist the development of waterlogging-tolerant crops.

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${\\rm H}^{+}\\text{-}{\\rm PP}_{{\\rm i}}{\\rm ase}$, provide the transmembrane 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.

Seagrasses grow submerged in aerated seawater but often in low O2 sediments. Elevated temperatures and low O2 are stress factors. Internal aeration was measured in two tropical seagrasses, Thalassia hemprichii and Enhalus acoroides, growing with extreme tides and diel temperature amplitudes. Temperature effects on net photosynthesis (P N) and dark respiration (R D) of leaves were evaluated. Daytime low tide was characterized by high pO2 (54 kPa), pH (8.8) and temperature (38°C) in shallow pools. As P N was maximum at 33°C (9.1 and 7.2 μmol O2 m−2 s−1 in T. hemprichii and E. acoroides, respectively), the high temperatures and reduced CO2 would have diminished P N, whereas R D increased (Q10 of 2.0–2.7) above that at 33°C (0.45 and 0.33 μmol O2 m−2 s−1, respectively). During night-time low tides, O2 declined resulting in shoot base anoxia in both species, but incoming water containing c. 20 kPa O2 relieved the anoxia. Shoots exposed to 40°C for 4 h showed recovery of P N and R D, whereas 45°C resulted in leaf damage. These seagrasses are ‘living near the edge’, tolerant of current diel O2 and temperature extremes, but if temperatures rise both species may be threatened in this habitat.

Drought tolerance is a complex trait that involves numerous genes. Identifying key causal genes or linked molecular markers can facilitate the fast development of drought tolerant varieties. Using a whole-genome resequencing approach, we sequenced 132 chickpea varieties and advanced breeding lines and found more than 144,000 single nucleotide polymorphisms (SNPs). We measured 13 yield and yield-related traits in three drought-prone environments of Western Australia. The genotypic effects were significant for all traits, and many traits showed highly significant correlations, ranging from 0.83 between grain yield and biomass to -0.67 between seed weight and seed emergence rate. To identify candidate genes, the SNP and trait data were incorporated into the SUPER genome-wide association study (GWAS) model, a modified version of the linear mixed model. We found that several SNPs from auxin-related genes, including auxin efflux carrier protein (PIN3), p-glycoprotein, and nodulin MtN21/EamA-like transporter, were significantly associated with yield and yield-related traits under drought-prone environments. We identified four genetic regions containing SNPs significantly associated with several different traits, which was an indication of pleiotropic effects. We also investigated the possibility of incorporating the GWAS results into a genomic selection (GS) model, which is another approach to deal with complex traits. Compared to using all SNPs, application of the GS model using subsets of SNPs significantly associated with the traits under investigation increased the prediction accuracies of three yield and yield-related traits by more than twofold. This has important implication for implementing GS in plant breeding programs.

The leaf economics spectrum (LES) describes consistent correlations among a variety of leaf traits that reflect a gradient from conservative to acquisitive plant strategies. So far, whether the LES holds in wetland plants at a global scale has been unclear. Using data on 365 wetland species from 151 studies, we find that wetland plants in general show a shift within trait space along the same common slope as observed in non-wetland plants, with lower leaf mass per area, higher leaf nitrogen and phosphorus, faster photosynthetic rates, and shorter leaf life span compared to non-wetland plants. We conclude that wetland plants tend to cluster at the acquisitive end of the LES. The presented global quantifications of the LES in wetland plants enhance our understanding of wetland plant strategies in terms of resources acquisition and allocation, and provide a stepping-stone to developing trait-based approaches for wetland ecology. Leaf economics spectrum theory has greatly advanced understanding of plant functional ecology, but it is unclear whether its predictions hold in wetland communities. Here, Pan and colleagues analyse leaf economics traits in wetland plants, showing that their trait relationships deviate from fully terrestrial plants, particularly by clustering towards acquisitive plant strategies.

Flooding is a common environmental variable with salinity. Submerged organs can suffer from O₂ deprivation and the resulting energy deficits can compromise ion transport processes essential for salinity tolerance. Tolerance of soil waterlogging in halophytes, as in glycophytes, is often associated with the production of adventitious roots containing aerenchyma, and the resultant internal O₂ supply. For some species, shallow rooting in aerobic upper soil layers appears to be the key to survival on frequently flooded soils, although little is known of the anoxia tolerance in halophytes. Halophytic species that inhabit waterlogged substrates are able to regulate their shoot ion concentrations in spite of the hypoxic (or anoxic) medium in which they are rooted, this being in stark contrast with most other plants which suffer when salinity and waterlogging occur in combination. Very few studies have addressed the consequences of submergence of the shoots by saline water; these have, however, demonstrated tolerance of temporary submergence in some halophytes.

Flash floods can submerge paddy field rice (Oryza sativa), with adverse effects on internal aeration, sugar status and survival. Here, we investigated the in situ aeration of roots of rice during complete submergence, and elucidated how underwater photosynthesis and floodwater pO2 influence root aeration in anoxic soil. In the field, root pO2 was measured using microelectrodes during 2 d of complete submergence. Leaf gas films that formed on the superhydrophobic leaves were left intact, or experimentally removed, to elucidate their effect on internal aeration. In darkness, root pO2 declined to very low concentrations (0.24 kPa) and was strongly correlated with floodwater pO2. In light, root pO2 was high (14 kPa) and primarily a function of the incident light determining the rates of underwater net photosynthesis. Plants with intact leaf gas films maintained higher underwater net photosynthesis relative to plants without gas films when the submerged shoots were in light. During complete submergence, internal aeration of rice in the field relies on underwater photosynthesis during the day and entry of O2 from the floodwater during the night. Leaf gas films enhance photosynthesis during submergence leading to improved O2 production and sugar status, and therefore contribute to the submergence tolerance of rice.