Climate change is a major threat to most of the agricultural crops grown in tropical and sub-tropical areas globally. Drought stress is one of the consequences of climate change that has a negative impact on crop growth and yield. In the past, many simulation models were proposed to predict climate change and drought occurrences, and it is extremely important to improve essential crops to meet the challenges of drought stress which limits crop productivity and production. Wheat and barley are among the most common and widely used crops due to their economic and social values. Many parts of the world depend on these two crops for food and feed, and both crops are vulnerable to drought stress. Improving drought stress tolerance is a very challenging task for wheat and barley researchers and more research is needed to better understand this stress. The progress made in understanding drought tolerance is due to advances in three main research areas: physiology, breeding, and genetic research. The physiology research focused on the physiological and biochemical metabolic pathways that plants use when exposed to drought stress. New wheat and barley genotypes having a high degree of drought tolerance are produced through breeding by making crosses from promising drought-tolerant genotypes and selecting among their progeny. Also, identifying genes contributing to drought tolerance is very important. Previous studies showed that drought tolerance is a polygenic trait and genetic constitution will help to dissect the gene network(s) controlling drought tolerance. This review explores the recent advances in these three research areas to improve drought tolerance in wheat and barley.

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Barley (Hordeum vulgare L.) is an important winter cereal crop grown in the semiarid Mediterranean, where late‐terminal drought stress during grain filling has recently become more common. The objectives of this study were to investigate the growth performance and grain yield of four barley cultivars under late‐terminal drought stress under both glasshouse and field conditions. At grain filling, four barley cultivars (Rum, ACSAD176, Athroh and Yarmouk) were exposed to three watering treatments: (1) well‐watered [soil maintained at 75 % field capacity (FC)], (2) mild drought stress at 50 % FC, (3) severe drought stress at 25 % FC in the glasshouse experiment and (1) well‐watered (irrigated once a week), (2) mild drought (irrigated once every 2 weeks), (3) severe drought (non‐irrigated; rainfed) in the field. As drought stress severity increased, gross photosynthetic rate, water potential, plant height, grain filling duration, spike number per plant, grain number per spike, 1000‐grain weight, straw yield, grain yield and harvest index decreased. In the glasshouse experiment, the six‐row barley cultivars (Rum, ACSAD176, and Athroh) had higher grain yield than the two‐row barley cultivar (Yarmouk), but the difference was not significant among the six‐row cultivars under all treatments. In the field experiment, Rum had the highest grain yield among all cultivars under the mild drought stress treatment. The two‐row cultivar (Yarmouk) had the lowest grain yield. In general, the traditional cultivar Rum had either similar or higher grain yield than the other three cultivars under all treatments. However, the yield response to drought differed between the cultivars. Those, Rum and ACSAD176, that were capable of maintaining a higher proportion of their spikes and grains per spike during drought also maintained a higher proportion of their yield compared with those in well‐watered treatment. In conclusion, cultivar differences in grain yield were related to spike number per plant and grain number per spike, but not days to heading or grain filling duration.

Heading time is a complex trait, and natural variation in photoperiod responses is a major factor controlling time to heading, adaptation and grain yield. In barley, previous heading time studies have been mainly conducted under field conditions to measure total days to heading. We followed a novel approach and studied the natural variation of time to heading in a world-wide spring barley collection (218 accessions), comprising of 95 photoperiod-sensitive (Ppd-H1) and 123 accessions with reduced photoperiod sensitivity (ppd-H1) to long-day (LD) through dissecting pre-anthesis development into four major stages and sub-phases. The study was conducted under greenhouse (GH) conditions (LD; 16/8 h; ∼20/∼16°C day/night). Genotyping was performed using a genome-wide high density 9K single nucleotide polymorphisms (SNPs) chip which assayed 7842 SNPs. We used the barley physical map to identify candidate genes underlying genome-wide association scans (GWAS). GWAS for pre-anthesis stages/sub-phases in each photoperiod group provided great power for partitioning genetic effects on floral initiation and heading time. In addition to major genes known to regulate heading time under field conditions, several novel QTL with medium to high effects, including new QTL having major effects on developmental stages/sub-phases were found to be associated in this study. For example, highly associated SNPs tagged the physical regions around HvCO1 (barley CONSTANS1) and BFL (BARLEY FLORICAULA/LEAFY) genes. Based upon our GWAS analysis, we propose a new genetic network model for each photoperiod group, which includes several newly identified genes, such as several HvCO-like genes, belonging to different heading time pathways in barley.

Increasing grain yield is still the main target of wheat breeding; yet today's wheat plants utilize less than half of their yield potential. Owing to the difficulty of determining grain yield potential in a large population, few genetic factors regulating floret fertility (i.e. the difference between grain yield potential and grain number) have been reported to date. In this study, we conducted a genome-wide association study (GWAS) by quantifying 54 traits (16 floret fertility traits and 38 traits for assimilate partitioning and spike morphology) in 210 European winter wheat accessions. The results of this GWAS experiment suggested potential associations between floret fertility, assimilate partitioning and spike morphology revealed by shared quantitative trait loci (QTLs). Several candidate genes involved in carbohydrate metabolism, phytohormones or floral development colocalized with such QTLs, thereby providing potential targets for selection. Based on our GWAS results we propose a genetic network underlying floret fertility and related traits, nominating determinants for improved yield performance.

Plant architecture has clear agronomic and economic implications for crops such as wheat and barley, as it is a critical factor for determining grain yield. Despite this, only limited molecular information is available about how grain-bearing inflorescences, called spikes, are formed and maintain their regular, distichous pattern. Here we elucidate the molecular and hormonal role of Six-rowed spike 2 (Vrs2), which encodes a SHORT INTERNODES (SHI) transcriptional regulator during barley inflorescence and shoot development. We show that Vrs2 is specifically involved in floral organ patterning and phase duration by maintaining hormonal homeostasis and gradients during normal spike development and similarly influences plant stature traits. Furthermore, we establish a link between the SHI protein family and sucrose metabolism during organ growth and development that may have implications for deeper molecular insights into inflorescence and plant architecture in crops.

Plant stature in temperate cereals is predominantly controlled by tillering and plant height as complex agronomic traits, representing important determinants of grain yield. This study was designed to reveal the genetic basis of tillering at five developmental stages and plant height at harvest in 218 worldwide spring barley (Hordeum vulgare L.) accessions under greenhouse conditions. The accessions were structured based on row-type classes [two- vs. six-rowed] and photoperiod response [photoperiod-sensitive (Ppd-H1) vs. reduced photoperiod sensitivity (ppd-H1)]. Phenotypic analyses of both factors revealed profound between group effects on tiller development. To further verify the row-type effect on the studied traits, Six-rowed spike 1 (vrs1) mutants and their two-rowed progenitors were examined for tiller number per plant and plant height. Here, wild-type (Vrs1) plants were significantly taller and had more tillers than mutants suggesting a negative pleiotropic effect of this row-type locus on both traits. Our genome-wide association scans further revealed highly significant associations, thereby establishing a link between the genetic control of row-type, heading time, tillering, and plant height. We further show that associations for tillering and plant height are co-localized with chromosomal segments harboring known plant stature-related phytohormone and sugar-related genes. This work demonstrates the feasibility of the GWAS approach for identifying putative candidate genes for improving plant architecture.

Abstract. In small-grain cereals, grain yield is closely associated with grain number. Improved spikelet survival is an important trait for increasing grain yield. We investigated spikelet number, spikelet survival and yield-related traits under greenhouse conditions, and pot-and soil-grown field conditions. Thirty-two spring barley (Hordeum vulgare L.) accessions (14 two-and 18 six-rowed accessions) were manually dissected to determine spikelet/floret number on the main culm spike (SNS) at awn primordium (AP), tipping (TIP), heading and anther extrusion. We observed a significant difference between two-and six-rowed barley for SNS and spikelet survival at all stages and growing conditions. Both traits were highly genetically controlled, with repeatability and broad-sense heritability values of 0.74-0.93. The rate of spikelet survival from AP to harvest was higher in two-(~70%) than in six-rowed (~58%) barley. Spikelet abortion, starting immediately after AP, was negatively affected by increased SNS and the thermal time required to reach the AP stage. The largest proportion of spikelet reduction happened during the AP-TIP phase, which was the most critical period for spikelet survival. The duration between AP and the end of stem elongation correlated better with spikelet survival and yield-related characters than the estimated duration of stem elongation using leaf height measurements. Our observations indicate that the main spike plays an important role in single-plant grain yield. Extending the length of the critical AP-TIP phase is promising for improving yield through increased spikelet development and survival. The results also demonstrate that greenhouse conditions are appropriate for studying traits such as phase duration and spikelet survival in barley.