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The task of obtaining and selecting plants which may not only survive under salty conditions, but grow and produce satisfactory yields is varied and complex. It is becoming necessary to select and breed crops for salt tolerance. Plants can be grown in artificially constructed salt basins to test their individual salt tolerance, but this takes time, at least one year. It would be desirable to develop a rapid test to determine the salt tolerance of a given plant. The use of a dye in conjunction with a series of salt solutions has been suggested for determining the salt tolerance of plants.

In arid end semiarid areas in the Western United States, soluble salts tend to accumulate in the soil in amounts harmful to crop production. A considerable portion of this land cannot be reclaimed due to the poor quality of permeability of the soil o Therefore, the wise selection of crops that will produce satisfactory yields on these soils in necessary.

Extensive land areas in arid and semi-arid regions contain excessive amounts of salt which inhibit or prevent the growth of desirable crop plants. This problem is increasing with the development of extensive irrigation projects. The future of irrigation agriculture will depend upon the development and the utilization of means of (1) preventing salt accumulation, (2) reclaiming salted soils. and (3) making the best use of those soils which by reason of drainage problems, soil texture and permeability, poor quality irrigation water, etc. are unsuited for economic reclamation. The utilization of land that is not feasible to completely reclaim, at least under present economic conditions, will depend upon the selection, development, and use of varieties and strains of crop plants which will produce economically under such conditions. The objectives of this research are to (1) test on a preliminary basis the salt tolerance of a number of promising crop plant materials. (2) observe the effect of salt upon plant behavior and growth, and (3) develop various methods of evaluating plant materials for salt tolerance.

Salt tolerance is a commercially important trait that affects plant species around the globe. Cellular response to saline environments is a well studied but complex system that is far from being completely understood. The SALT OVERLY SENSITIVE 1 (SOS1) gene is a critical component of salt tolerance in many species, encoding a plasma membrane Na+/H+ antiporter that plays an important role in germination and growth in saline environments. Here we report a preliminary investigation of salt tolerance in quinoa (Chenopodium quinoa Willd.). Quinoa is a halophytic grain crop of the Chenopodiaceae family with impressive nutritional content and an increasing world-wide market. Many quinoa varieties have impressive salt tolerance characteristics and research suggests quinoa may utilize novel mechanisms to confer salt tolerance. At this time there is no published data on the molecular characteristics of those mechanisms. We report the identification and sequencing of the SOS1 gene in quinoa, including a full length cDNA sequence of 3490 bp and a full length genomic clone of 21314 bp. Sequence analysis predicts the quinoa SOS1 homolog spans 23 exons and is comprised of 3474 bp of coding sequence (excluding the stop codon). Introns comprise 17840 bp of the genomic clone and range in size from 77 to 2123 bp. The predicted protein contains 1158 amino acid residues and aligns closely with SOS1 homologs of other species. The quinoa SOS1 homolog contains two putative domains, a Nhap cation-antiporter domain and a cyclic-nucleotide binding domain. Sequence analyses of both cDNA fragments and intron fragments suggest that two SOS1 loci are present in the quinoa genome that are likely orthologous loci derived from the ancestral diploid genomes of the modern allotetraploid quinoa genome. This report represents the first molecular characterization of a putative salt-tolerance gene in C. quinoa.

Halogeton (Halogeton glomeratus) is a halophytic, invasive species that displaces Gardner’s saltbush (Atriplex gardneri) on saline rangelands. Forage kochia (Bassia prostrata) is a potential species to rehabilitate these ecosystems. This study compared the salinity tolerance of these species and tall wheatgrass (Thinopyrum ponticum) and alfalfa (Medicago sativa). Plants were evaluated for 28 days in hydroponics where they were maintained at 0, 150, 200, 300, 400, 600, and 800 mM NaCl. Shoot growth and ion accumulation were determined. Alfalfa and tall wheatgrass were severely affected by salt with both species’ shoot mass just 32% of control at 150 mM NaCl. Alfalfa did not survive above 300 mM NaCl, while, tall wheatgrass did not survive at salt levels above 400 mM NaCl. In contrast, forage kochia survived to 600 mM, but produced little shoot mass at that level. Halogeton exhibited ‘halophytic’ shoot growth, reaching maximum mass at 141 mM, and not less mass than the control until salinity reached 400 mM. Gardner’s saltbush did not show a dramatic decrease in dry mass produced until it reached salt levels of 600 and 800 mM NaCl. Forage kochia yielded high amounts of dry mass in the absence of salt, but also managed to survive up to 600 mM NaCl. Salt tolerance ranking (GR50 = 50% reduction in shoot mass) was Gardner’s saltbush=halogeton>forage kochia> alfalfa>tall wheatgrass. Both halogeton and Gardner’s saltbush actively accumulated sodium in shoots, indicating that Na+ was the principle ion in osmotic adjustment. In contrast, forage kochia exhibited a linear increase (e.g. passive uptake) in Na+ accumulation as salinity increased. This study confirmed that halogeton is a halophytic species and thus well adapted to salt-desert shrubland ecosystems. Gardner’s saltbush, also a halophyte, was equally salt tolerant, suggesting other factors are responsible for halogeton displacement of Gardner’s saltbush. Forage kochia is a halophytic species that can survive salinity equal to seawater, but is not as salt tolerant as Gardner’s saltbush and halogeton.

Crop lands around the world are becoming more salt-affected due to natural processes and agricultural practices. Due to this increase of salinization, acquisition of saline tolerant germplasm for breeding purposes is becoming a priority. Although cultivated sunflower is classified as a moderately salt tolerant crop, highly tolerant germplasm may be of value. The goal of this study was to screen Helianthus spp. in order to determine the salt tolerance of different genotypes. To accomplish this goal, a novel method of rapid screening was developed. Screening for tolerance at initial growth stages was accomplished by germinating seeds in varying concentrations of NaCl solution in petri dishes. Radicle lengths were measured as an indicator of tolerance. This method identified genotypes that are more tolerant than others during germination. Greenhouse trials were also conducted to ascertain morphological measurements during vegetative stages. Two field locations were chosen to screen germplasm for tolerance through physiological maturity; College Station, TX with low salt concentrations and Pecos, TX with high concentrations of salt in the soil and water. Vegetative growth measurements showed a significant genotype by environment interaction. Due to insect infestation in both locations, yields could not be accurately measured and thus compared between sites in 2010. Yields between locations in 2011 showed significant differences and identified germplasm more suited for cropping in salt affected soil. Seed oil content was determined with Fourier Transform Near-Infrared Spectroscopy. Seed oil content was not significantly different between locations, but was highly significant between genotypes. These screenings identified genotypes that are more salt tolerant than others.

Salt glands are important structural adaptations in some plant and animal species that are involved in the excretion of excess salts. Zoysia matrella is a highly salt tolerant turf grass that has salt glands. Two cultivars of Z. matrella, ‘Diamond’ and ‘Cavalier’, were examined in this study to look for salt gland related factors responsible for the differences in their degree of salt tolerance. In addition to the adaxial salt gland density being higher in ‘Diamond’, the salt glands in salt treated (300 mM NaCl) plants of this cultivar were bigger than the ones in ‘Cavalier’. ‘Diamond’, as well as some of the ‘Diamond’ x ‘Cavalier’ hybrid lines, showed a significant induction in salt gland density in response to salt treatment. Examination of salt gland density in ‘Diamond’ x ‘Cavalier’ hybrid lines showed that salt gland density was a highly heritable trait in the salt-treated population. Ultrastructural modifications in the salt glands observed with Transmission Electron Microscopy (TEM), coupled with Cl- localization studies, suggested a preference for symplastic transport of saline ions in Z. matrella. Salt glands have been studied in several plant species; however, no studies have tried to associate the role of ion transporters with the functioning of salt glands in plants. RNA in situ studies with Na+ transporters showed localization of ZmatHKT1 transcripts in the adaxial salt glands, leaf mesophyll and bundle sheath cells for both cultivars. ZmatSOS1 expression was observed in the xylem parenchyma cells for leaves from both cultivars, but the expression was markedly different around the cells bordering the vascular tissue. The strongest expression of ZmatSOS1 for ‘Diamond’ was seen in the bundle sheath cells and the phloem, while for ‘Cavalier’ the signal was strongest in the mestome sheath cells and in cells surrounding the phloem. No expression of ZmatSOS1 was seen in the salt glands for either cultivars. ZmatNHX1 expression in both cultivars was very low, and observed in the salt glands and neighboring epidermal cells. Three alleles of ZmatNHX1 were identified in Z. matrella, along with three alternatively-spliced forms of ZmatNHX1, the expression of which were cultivar and treatment specific. Together, these results provide a model for salt transport in Z. matrella and signify potential roles of salt glands and select ion transporters in the salt tolerance of this species.

The accumulation of salts in soil is an important agricultural problem that limits crop productivity. Salts containing sodium (Na⁺) are particularly problematic, as cytosolic Na⁺ can interfere with cellular metabolism and lead to cell death. Maintaining low levels of cytosolic Na⁺, therefore, is critical for plant survival during growth in salt. Mechanisms to regulate Na⁺ accumulation in plant cells include extrusion of Na⁺ from the cell and sequestration of Na⁺ into intracellular compartments. Both of these processes are controlled in part through the action of Na⁺/H⁺ exchangers belonging to the Cation/Proton Antiporter-1 (CPA1) gene family. Genes belonging to this family have been identified in both salt-sensitive and salt-tolerant species, suggesting that salt-tolerant species may have evolved salt tolerance through modification of these existing pathways. The research presented here has focused on understanding how salt tolerance has evolved in Brassicaceae species, and particularly on the role that CPA1 genes have played in the adaptation to salinity of Eutrema salsugineum. Specific projects have sought to understand 1) how copy number variation and changes in coding sequences of CPA1 genes contribute to salt tolerance in E. salsugineum and its salt-tolerant relative Schrenkiella parvula, 2) whether functional or regulatory changes in Salt Overly Sensitive 1 (SOS1) from E. salsugineum (EsSOS1) contribute to its enhanced salt tolerance, and 3) whether accessions of Arabidopsis thaliana differ significantly in their response to salt stress.The results indicate that EsSOS1 and SOS1 from S. parvula (SpSOS1) both confer greater salt tolerance in yeast than SOS1 from A. thaliana (AtSOS1) when activated by the complex of the SOS2 kinase and SOS3 calcium-binding protein, whereas only EsSOS1 confers enhanced salt tolerance in the absence of activation. When expressed in A. thaliana, EsSOS1 also confers greater salt tolerance than AtSOS1 through regulatory changes that likely involve differences in expression pattern. Together, the results presented here suggest that mechanisms regulating cellular Na⁺ accumulation that exist in salt-sensitive crop species could be altered to enhance growth in salty soils. In addition, the 19 A. thaliana accessions used to create the MAGIC population were shown to differ significantly in their response to salt stress.

Salinity stress limits the growth and productivity of agricultural crops in many regions of the world. Whole plant tolerance to soil salinity involves numerous processes in many different tissues and cell types. For many cereals, sensitivity to salinity is due to the accumulation of sodium (Na⁺) to toxic concentrations in the leaves. This thesis investigates a mechanism of control of Na⁺ accumulation in leaves of wheat. Bread wheat excludes sodium from the leaves better than durum wheat. Bread wheat is hexaploid (AABBDD) whereas durum wheat is tetraploid (AABB). The D-genome in bread wheat carries a major locus for sodium exclusion, Kna1, which may contribute to the differences in sodium exclusion between bread wheat and durum wheat. An unusual durum wheat, Line 149, excludes sodium to a similar degree as bread wheat. Line 149 was derived from a cross between a Triticum monococcum (accession C68-101; AA) and a durum wheat (T. turgidum ssp. durum cv. Marrocos; AABB). Line 149 had been found to contain two major genes for sodium exclusion, named Nax1 and Nax2, which appeared to retrieve sodium from the xylem sap in the roots and so prevent it reaching the leaves. Line 149 had been crossed with the durum wheat cv. Tamaroi, which accumulates high concentrations of Na⁺ in the leaves, and near-isogenic single-gene mapping populations had been developed for Nax1 and Nax2. Nax1 had been located on chromosome 2A. The objective of this thesis was to map Nax2 and identify a candidate gene. Nax2 mapped to chromosome 5AL based on linkage to microsatellite markers. A high-affinity potassium (K⁺) transporter (HKT)-like gene, HKT1;5 was considered as a candidate gene for Nax2, based on similarity of the phenotype to a rice orthologue. Sequence information from a wheat HKT1;5-like expressed sequence tag in the public database was used to develop a probe for use in Southern hybridsation. A HKT1;5-like fragment was identified in Line 149 and T. monococcum C68-101, but was absent in Tamaroi. The HKT1;5-like gene, named TmHKT1;5-A, co-segregated with Nax2 in the Nax2 single-gene mapping population. The HKT1;5 probe identified three putative HKT1;5-like genes on the long arm of chromosome 4B, and one HKT1;5-like gene on the long arm of chromosome 4D, in Langdon (T. turgidum ssp. durum) substitution lines, and in Chinese Spring (T. aestivum) ditelomeric lines. No A-genome HKT1;5 like gene was identified in Langdon or Chinese Spring. The D-genome HKT1;5 gene, named TaHKT1;5-D, was found to co-locate with Kna1, the gene for sodium exclusion in bread wheat, in Chinese Spring chromosome 4D deletion lines. Nax2 (TmHKT1;5-A) was found to be homoeologous with the gene for sodium exclusion in bread wheat, Kna1 (TaHKT1;5-D). TmHKT1;5-A and TaHKT1;5-D, and their promoters, were 94% identical, and both were expressed in the roots of wheat plants. This is consistent with the genes being located in the stele of the roots and retrieving Na⁺ from the xylem sap as it flows towards the shoot, and so excluding Na⁺ from the leaves. A marker for TmHKT1;5-A was developed to track this gene in durum wheat breeding programs. A study of the HKT1;5 gene in diploid ancestors of wheat indicated that this gene is present in most Triticum monococcum accessions, some T. boeoticum accessions, but not present in any T. urartu accessions. T. urartu is the likely A genome ancestor of modern wheat. This may explain the absence of HKT1;5 in the A genome of modern wheat. The protein encoded by TaHKT1;5-D transported sodium when expressed in Xenopus laevis oocytes. The inward currents were specific to Na⁺, but at particular mole fractions of Na⁺ and K⁺ outward currents were observed that were consistent with outward K⁺ transport. These data were consistent with the putative physiological function, of retrieving Na⁺ from the xylem sap as it flows to the leaves, and resulting in a net exchange with K⁺. A construct designed to silence the expression of TaHKT1;5-D was introduced to bread wheat cv. Bob White. Nineteen putative transgenic plants were developed. The leaf Na⁺ concentrations and genotype of the T1 individuals were assayed. The data from two of the transgenic plants indicated that TaHKT1;5-D may have been silenced and that this may have lead to the increase in Na⁺ accumulation in the leaves. However, this data is not conclusive at this time. The information gained from this study will assist the introduction of the Na⁺ exclusion trait into current durum cultivars, which are poor at excluding Na⁺ and are salt sensitive. This information will also contribute to the body of knowledge of ion transport in plants and salinity tolerance in wheat. / Thesis (Ph.D.) - University of Adelaide, School of Agriculture, Food and Wine, 2008