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Water-saving attempts for rice cultivation often reduce yields. Maintaining productivity under drought is possible when rice genotypes are bred with improved metabolism and spikelet fertility. Although attempts have been made to introgress water mining and water use efficiency traits, combining acquired tolerance traits (ATTs), that is, specific traits induced or upregulated to better tolerate severe stress, appears equally important. In our study, we screened 90 rice germplasm accessions that represented the molecular and phenotypic variations of 851 lines of the 3 K rice panel. Utilising phenomics, we identified markers linked to ATTs through association analysis of over 0.2 million SNPs derived from whole-genome sequences. Propensity to respond to ‘induction’ stress varied significantly among genotypes, reflecting differences in cellular protection against oxidative stress. Among the ATTs, the hydroxyl radical and proline contents exhibited the highest variability. Furthermore, these significant variations in ATTs were strongly correlated with spikelet fertility. The 43 significant markers associated with ATTs were further validated using a different subset of contrasting genotypes. Gene expression studies and metabolomic profiling of two well-known contrasting genotypes, APO (tolerant) and IR64 (sensitive), identified two ATT genes: AdoMetDC and Di19. Our study highlights the relevance of polyamine biosynthesis in modulating ATTs in rice. Genotypes with superior ATTs and the associated markers can be effectively employed in breeding rice varieties with sustained spikelet fertility and grain yield under drought.

Drought poses a serious threat to crop production worldwide, and is expected particularly to affect rice production and hence food security. Given that wheat is known to tolerate drought better than rice, we compare rice and wheat (cv. Weebill) to understand the species level differences in drought adaptive mechanisms. We also compare two contrasting rice genotypes (IR64, drought susceptible, and Apo, drought tolerant) for such mechanisms under well-watered (100% field capacity, 100%FC) and water-limited (60%FC) conditions. The reduction in biomass of wheat under water limitation was smaller due to a higher rate of photosynthesis associated with maintenance of tissue turgor compared to rice genotypes. Drought caused greater inhibition of Photosystem II quantum efficiency, carboxylation efficiency, and photosynthetic capacity parameters in IR64 than in Apo. Transcript levels of photosynthesis-related genes were also significantly more repressed by water limitation in IR64, whilst the wheat genotype showed smaller reduction than Apo. Despite higher non-photochemical quenching (NPQ), a smaller increase in scavenging enzymes in IR64 resulted in more accumulation of reactive oxygen species (ROS) in 60%FC than in 100%FC compared to Apo. As a photoprotection mechanism, increased levels of NPQ resulted in lower ROS accumulation in wheat despite the similar increase in scavenging enzyme transcript levels as in Apo, signifying the importance of preventing oxidative burst for enhanced drought tolerance. In Apo, upregulation of the 9-cis-epoxycarotenoid dioxygenase 2 gene implies the use of xanthophyll pool for the ABA biosynthesis. Our data suggest that regulating photosynthesis and oxidative protection in the wheat genotype enhanced drought tolerance. Improving these traits for rice is crucial to develop drought-tolerant rice genotypes.

Drought significantly decreases crop productivity, especially in high water consuming crops like rice. Grain filling is one of the important critical growth phases in rice and drought during this phase leads to significant reduction in yield. In this study, a comparison was made between IR64 (drought susceptible) and Apo (drought tolerant) rice genotypes to capture the response to water limitation (50% field capacity (FC)) compared with the control (100%FC) during grain filling. Plants were grown in a high-throughput phenomics facility for precise imposition of moisture stress during grain filling. Apo performed better in water limited conditions with lower reduction of photosynthetic rate and maintenance of lower leaf temperature than IR64. Days from sowing to maturity, spikelet fertility and seed weight were more impeded by water limitation in IR64 than in Apo. Unlike Apo, IR64 did not show any decrease in transpiration rate at 50%FC compared with 100%FC. Metabolomic profiling of spikelets at grain filling showed distinct effects of water limitation on accumulation of metabolites, especially in Apo. Secondary metabolism, mainly the phenylpropanoid pathway involved in scavenging mechanisms, was upregulated in Apo. Accumulation of most amino acids was significantly higher in IR64 than in Apo. Due to higher rates of photosynthesis under stress, most carbohydrates accumulated more in Apo than in IR64 at 50%FC. Sucrose transporters were significantly upregulated in water limited conditions especially in Apo. Overall, thanks to higher source capacity, more source to sink transport and better scavenging, Apo showed a lower reduction in yield than IR64.

Plants have evolved several adaptive mechanisms to cope with water-limited conditions. While most of them are through constitutive traits, certain “acquired tolerance” traits also provide significant improvement in drought adaptation. Most abiotic stresses, especially drought, show a gradual progression of stress and hence provide an opportunity to upregulate specific protective mechanisms collectively referred to as “acquired tolerance” traits. Here, we demonstrate a significant genetic variability in acquired tolerance traits among rice germplasm accessions after standardizing a novel gradual stress progress protocol. Two contrasting genotypes, BPT 5204 (drought susceptible) and AC 39000 (tolerant), were used to standardize methodology for capturing acquired tolerance traits at seedling phase. Seedlings exposed to gradual progression of stress showed higher recovery with low free radical accumulation in both the genotypes compared to rapid stress. Further, the gradual stress progression protocol was used to examine the role of acquired tolerance at flowering phase using a set of 17 diverse rice genotypes. Significant diversity in free radical production and scavenging was observed among these genotypes. Association of these parameters with yield attributes showed that genotypes that managed free radical levels in cells were able to maintain high spikelet fertility and hence yield under stress. This study, besides emphasizing the importance of acquired tolerance, explains a high throughput phenotyping approach that significantly overcomes methodological constraints in assessing genetic variability in this important drought adaptive mechanism.

Drought tolerance is governed by constitutive and acquired traits. Combining them has relevance for sustaining crop productivity under drought. Mild levels of stress induce specific mechanisms that protect metabolism when stress becomes severe. Here, we report a comparative assessment of “acquired drought tolerance (ADT)” traits in two rice cultivars, IR64 (drought susceptible) and Apo (tolerant), and a drought-tolerant wheat cultivar, Weebill. Young seedlings were exposed to progressive concentrations of methyl viologen (MV), a stress inducer, before transferring to a severe concentration. “Induced” seedlings showed higher tolerance and recovery growth than seedlings exposed directly to severe stress. A novel phenomic platform with an automated irrigation system was used for precisely imposing soil moisture stress to capture ADT traits during the vegetative stage. Gradual progression of drought was achieved through a software-controlled automated irrigation facility. This facility allowed the maintenance of the same level of soil moisture irrespective of differences in transpiration, and hence, this platform provided the most appropriate method to assess ADT traits. Total biomass decreased more in IR64 than in Apo. The wheat cultivar showed lower levels of damage and higher recovery growth even compared to Apo. Expression of ROS-scavenging enzymes and drought-responsive genes was significantly higher in Apo than in IR64, but differences were only marginal between Apo and Weebill. The wheat cultivar showed significantly higher stomatal conductance, carbon gain, and biomass than the rice cultivars, under drought. These differences in ADT traits between cultivars as well as between species can be utilised for improving drought tolerance in crop plants.

Rice yield reduction due to water limitation depends on its severity and duration and on the phenological stage of its occurrence. We exposed three contrasting rice genotypes, IR64, UPLRi7 and Apo (adapted to lowland, upland and aerobic conditions, respectively), to three water regimes (puddle, 100% and 60% field capacity) in pots during the vegetative (GSI), flowering (GSII) and grain filling (GSIII) stages. Stress at all the three stages significantly reduced yield especially in lowland genotype IR64. Effect of water limitation was more severe at GSII than at the other two stages. Stress at GSI stage reduced both source activity (leaf area and photosynthetic rate) and sink capacity (tiller number or panicle number per pot). When stress was imposed at GSII, spikelet fertility was most affected in all the three genotypes. In both GSII and GSIII, although leaf area was constant in all the three water regimes, estimated relative whole-plant photosynthesis was strongly associated with yield reduction. Reduced photosynthesis due to stress at any given stage was found to have direct impact on yield. Compared to the other genotypes, Apo had deeper roots and maintained a better water relation, thus, higher carbon gain and spikelet viability, and ultimately, higher biomass and productivity under water-limited conditions. Therefore, screening for these stage-dependent adaptive mechanisms is crucial in breeding for sustained rice production under water limitation.

Finger millet (Eleusine coracana (L.) Gaertn) is an important crop for food security because of its tolerance to drought, which is expected to be exacerbated by global climate changes. Nevertheless, it is often classified as an orphan/underutilized crop because of the paucity of sci- entific attention. Among several small millets, finger millet is considered as an excellent source of essential nutrient elements, such as iron and zinc; hence, it has potential as an alternate coarse cereal. However, high-quality genome sequence data of finger millet are currently not available. One of the major problems encountered in the genome assembly of this species was its polyploidy, which hampers genome assembly compared with a diploid genome. To over- come this problem, we sequenced its genome using diverse technologies with sufficient cover- age and assembled it via a novel multiple hybrid assembly workflow that combines next- generation with single-molecule sequencing, followed by whole-genome optical mapping using the Bionano IrysVR system. The total number of scaffolds was 1,897 with an N50 length >2.6 Mb and detection of 96% of the universal single-copy orthologs. The majority of the homeologs were assembled separately. This indicates that the proposed workflow is applicable to the as- sembly of other allotetraploid genomes.