Consistent gains in grain yield in dry environments have been made by empirical breeding although there is disturbing evidence that these gains may have slowed. There are few examples where an understanding of the physiology and the genetics of putative important drought-related traits has led to improved yields. Success will first depend on identifying the most important traits in the target regions. It will then depend on accurate and fast phenotyping, which, in turn, will lead to: (1) trait-based selection being immediately transferable into breeding operations and (2) being able to identify the underlying genes or the important genomic regions (quantitative trait loci), perhaps leading to efficient marker-based selection (MBS). Genetic complexity, extent of genotype × environment (G × E) interaction and sampling cost per line will determine value of phenotyping over MBS methods. Here, we review traits of importance in dry environments and review whether molecular or phenotypic selection methods are likely to be the most effective in crop improvement programs and where the main bottlenecks to selection are. We also consider whether selection for these traits should be made in dry environments or environments where there is no soil water limitation. The development of lines/populations for trait validation studies and for varietal development is also described. We firstly conclude that despite the spectacular improvements in molecular technologies, fast and accurate phenotyping remains the major bottleneck to enhancing yield gains in water-limited environments. Secondly, for most traits of importance in dry environments, selection is generally conducted most effectively in favourable moisture environments.
Salinity affects plant growth by the osmotic stress of the salt around the roots as well as by toxicity caused by excessive accumulation of salt in leaves. The aim of this study was to determine whether there is significant genetic variation in tolerance to osmotic stress that can be useful in improving the salinity tolerance of crop plants. Durum wheat is a salt-sensitive crop whose yield is reduced by moderately saline soils. Genetic variation in tolerance to osmotic stress in durum wheat was examined in 50 international durum varieties and landraces by measuring the response of stomatal conductance to salt stress before salts built up in the leaf. Stomatal conductance is a sensitive indicator of the osmotic stress because it is reduced immediately with the onset of salinity, and is the initial and most profound cause of a decline in CO2 assimilation rate. Genetic differences of 2-3-fold were found in the magnitude of the response of stomatal conductance to salt-induced osmotic stress. Higher stomatal conductance in salt related to higher CO2 assimilation rate. There was a positive relationship between stomatal conductance and relative growth rate in salt. This study shows the potential for new genetic gains in salt tolerance in durum wheat.
Relative water content (RWC) is used extensively to determine the water status of plants relative to their fully turgid condition. However, plants often adjust osmotically to salinity or water deficit, which maintains turgor pressure and obscures the definition of 'full turgidity'. To explore this problem, turgor was measured by isopiestic psychrometry in mature leaf blades of barley (Hordeum vulgare) and durum wheat (Triticum turgidum ssp. durum) salinised to 150 mm NaCl, or bread wheat (Triticum aestivum) grown in soil dehydrated to varying degrees. Osmotic adjustment maintained turgor in all the plants but despite full maintenance in some of the salinised plants, their leaf RWC decreased substantially. This occurred because excess water was absorbed while the samples were floated on water as part of the RWC measurement. The absorption falsely increased the weight of the 'fully turgid' condition, causing RWC to be anomalously low by 10-15%. Cell solution was secreted into intercellular spaces and was seen under a microscope, which is a test encouraged for all RWC measurements. Several alternate methods are suggested for rehydrating tissues while minimising excess water absorption, but no simple definition of 'full turgidity' seems possible. In general, direct measurements of osmotic adjustment and turgor are preferred.
The capacity of plants to tolerate high levels of salinity depends on the ability to exclude salt from the shoot, or to tolerate high concentrations of salt in the leaf (tissue tolerance). It is widely held that a major component of tissue tolerance is the capacity to compartmentalize salt into safe storage places such as vacuoles. This mechanism would avoid toxic effects of salt on photosynthesis and other key metabolic processes. To test this, the relationship between photosynthetic capacity and the cellular and subcellular distribution of Na+, K+ and Cl- was studied in salt-sensitive durum wheat (cv. Wollaroi) and salt-tolerant barley (cv. Franklin) seedlings grown in a range of salinity treatments. Photosynthetic capacity parameters (Vcmax, Jmax) of salt-stressed Wollaroi decreased at a lower leaf Na+ concentration than in Franklin. Vacuolar concentrations of Na+, K+ and Cl- in mesophyll and epidermal cells were measured using cryo-scanning electron microscopy (SEM) X-ray microanalysis. In both species, the vacuolar Na+ concentration was similar in mesophyll and epidermal cells, whereas K+ was at higher concentrations in the mesophyll, and Cl- higher in the epidermis. The calculated cytoplasmic Na+ concentration increased to higher concentrations with increasing bulk leaf Na+ concentration in Wollaroi compared to Franklin. Vacuolar K+ concentration was lower in the epidermal cells of Franklin than Wollaroi, resulting in higher cytoplasmic K+ concentrations and a higher K+ : Na+ ratio. This study indicated that the maintenance of photosynthetic capacity (and the resulting greater salt tolerance) at higher leaf Na+ levels of barley compared to durum wheat was associated with the maintenance of higher K+, lower Na+ and the resulting higher K+ : Na+ in the cytoplasm of mesophyll cells of barley.
Coleoptile length in wheat (Triticum aestivum L.) can be affected by several factors, including genotype, height-reducing genes and environmental factors, including temperature. There is little information on how these factors influence rate and duration of coleoptile growth to determine final coleoptile length in wheat. Coleoptile growth was determined for eight genotypes representing four different height-reducing genes: gibberellic acid (GA)-sensitive, standard height (rht), GA-sensitive semidwarfs (Rht8); and GA-insensitive, semidwarfs (Rht2). These were grown in the dark at three temperatures (12, 16 and 20˚C) and coleoptile lengths measured every 12˚Cd. Logistic growth curves were fitted to coleoptile growth data for each genotype with thermal time as the explanatory variable. Differences in final coleoptile length were largely attributable to differences in rate of coleoptile elongation although there were small differences in duration of growth between genotypes. The longer coleoptile of the rht wheats was achieved through the fastest rate of coleoptile elongation. Coleoptiles of Rht8 wheats were equivalent in final length to rht wheats at 107 mm, but achieved this through a slower growth rate (2.10 mm ˚Cd–1) combined with an increased duration of growth (57˚Cd). In contrast, the shorter coleoptiles of Rht2 wheats resulted from 25% slower rates of elongation than either Rht8 or rht. There were no interactions between the components of coleoptile growth and temperature, although a longer duration and a fast rate of growth combined to increase coleoptile length at 12˚C compared with either 16 or 20˚C. In a second experiment, dry matter partitioning and length of coleoptile, subcrown internode (SCI), shoot and roots were determined after 200˚Cd. In Rht2, the SCI and shoot were short while roots were longer than either Rht8 or rht. Reduced dry matter (DM) partitioning to the coleoptile and SCI and DM retention in the seed reduced the endosperm-use efficiency (EUE) of Rht compared with rht. EUE was poor also in Rht8, apparently through increased respiratory losses. Reduced partitioning of dry matter to coleoptiles and the SCI in Rht2 increased the root : shoot ratio compared with rht or Rht8. We conclude that either increased rate or duration of coleoptile growth could be targeted in a breeding program that aims to increase coleoptile length in wheat.